UNIVERSITY OF CALIFORNIA ANDREW SMITH HALLlDIft LECTURES ELEMENTS OF BOTANY. PART I. CONTAINING THE DESCRIPTIVE ANATOMY OF THOSE ORGANS, ON WHICH THE GROWTH AND PRESERVATION OF THE VEGETABLE DEPEND. BY ANTHONY TODD THOMSON, F. L. S. MEM. OF THE ROY. COLL. OF SURGEONS, AND THE MEDICO-CHIRURG. SOC. OF LONDON: FELLOW OF THE MEDICAL SOCIETY, OF THE ROY. PHY- SICAL SOCIETY, AND THE SPECULATIVE SOCIETY, OF EDINBURGH: MEM. OF THE SOCIETE DE MEDECINE DE MARSEILLES, AND OF THE SOCIETE MEDICALE D*EMULATION DE PARIS. VOL. I. LONDON : PRINTED FOR LONGMAN, HURST, REES, ORME, AND BROWN, PATERNOSTER ROW. S. GOSNELL, Printer, Little Queen Street, London. TO THE PRESIDENT AND THE FELLOWS OP THE ROYAL COLLEGE OF PHYSICIANS, OP LONDON; IN THE HOPE THAT THEIR AVOWED PATRONAGE OF AN ELEMENTARY WORK ON BOTANICAL SCIENCE, WILL PROMOTE THE FUTURE STUDY OF THAT BRANCH OF KNOWLEDGE AS A PART OF MEDICAL EDUCATION; , AND THAT, BY THUS DISPLAYING THEIR APPROBATION OF WHATEVER CAN ENLARGE THE VIEWS OF THE STUDENT IN THE INVESTIGATION OF ORGANIC LIFE, THE CHARACTER OF THE MEDICAL PHILOSOPHER MAY BE ELEVATED TO ITS DUE RANK IN SOCIETY; THESE LECTURES, WITH THEIR GRACIOUS PERMISSION, ARE INSCRIBED BY THE AUTHOR. u2 PREFACE. NUMEROUS works on the elementary principles of Botany, and many of them very excellent pro- ductions, have been published, both in this coun- try and on the continent, within the last twenty years: and, by pointing out the mode of inves- tigating the laws of vegetable life, have done much to remove an objection to the study of Bo- tanical science, which had long prevailed, " that " it is a pursuit that amuses the fancy and ex- " ercises the memory, without improving the mind " or advancing any real knowledge *." Under these circumstances, the publication of a new work on this subject may require some apology; and it was not my intention to have placed the fol- lowing Lectures before the public, had I not ac- cidentally met with a manuscript copy of them, * White** Nat. Hist, of Sclborne, 8vo. London, 1822, vol. ii. p. 38. VI PREFACE. as they were delivered to my pupils, exposed for sale in a bookseller's shop*. Reflecting, therefore, that any peculiar theories connected with vege- table physiology, which I had taught, and of which little more than outlines had been sketched for the class-room, were likely to be much mis- represented, and even that many of the facts taken from authors might be misstated, justice to my reputation required that I should rather pub- lish my own opinions, than run the hazard of their getting into the press in a mutilated condi- tion. In revising my manuscript, however, for this purpose, I found that the view of the sub- ject opened before me, that one investigation led on to another; and that a frequent appeal to Na- ture forced me to reject much of what I had for- merly regarded as truth ; so that the work im- perceptibly extended far beyond the limits I had allotted to it; and, now, retains little more than the name and the arrangement of the original Lectures. The present volume treats of the forms and the anatomy of those organs which are necessary for the growth and the preservation of the vege- PREFACE. Vll table individual; and, although it does not pro- fess to enter fully into the explanation of the laws which regulate the functions of these organic structures, yet, much of physiology has been in- troduced, both to illustrate the descriptions, and to relieve the dry ness of the anatomical details. The necessity of an accurate knowledge of struc- ture, will be fully perceived in the perusal of the physiological discussions which are intended to form the subject of the second volume. I trust that the plan of illustrating the descriptions by the introduction of cuts into the body of the letter-press, will be found of considerable assist- ance to the student; and that the engraved plates, although not all of equal merit in point of execu- tion, will, nevertheless, be found sufficient for conveying correct ideas of those parts, which, from their minuteness, are necessarily microscopic objects. As an apology for errors which may be detected in the volume, I might plead the interruptions, anxieties, and unremitting duties of very exten- sive professional occupations ; but, as I am aware that no author is dragged before the tribunal of viii PREFACE. the public against his inclination, and until he declares himself prepared for trial, I do not think such circumstances valid reasons for lenity or favour. The work, therefore, is published with- out any claim of indulgence, but with a convic- tion, that, although many objections may be raised to the doctrines it contains, yet, if it have merit, that its faults will be lightly handled ; and if it be undeserving of approbation, that even the countenance of the learned body, under whose patronage it is sent forth, cannot alter the sen- tence which Justice should dictate. ANTHONY TODD THOMSON. 91, Sloane Street, May 30, 1822, CONTENTS. LECTURE I. -Page I. INTRODUCTION SKETCH OF THE RISE AND PROGRESS OF BOTANY UTILITY OF THE SCIENCE METHOD OF STUDYING IT, AND PLAN OF THE COURSE. LECTURE II.~Page 39. DEFINITION OF A PLANT GENERAL VIEW OF THE VEGETABLE FUNCTIONS. LECTURE III. Page 69. GENERAL COMPONENTS OF THE VEGETABLE STRUCTURE. LECTURE IV. Page 125. VEGETABLE ORGANIZATION. THE ROOT ITS SITUA- TION, SPECIES, AND VARIETIES DIRECTION AND DURATION. LECTURE V. Page 189. THE SUBJECT OF THE FORMER LECTURE CONTI- NUED. OF SOILS AND MANURES. OF THE MEDI- CINAL AND DIETETIC AL PROPERTIES OF ROOTS, LECTURE VI. Page 24 1. THE STEM ITS DIRECTION DIVISIONS AND BRANCH- ING COVERING COLOUR FIGURE. CLASSIFICA- TION OF STEMS. VOL. I. b CONTENTS. LECTURE VII. Page 284. SUBSTANCE AND ORGANIZATION OF THE STEM AND BRANCHES : ANATOMICAL DEMONSTRATION OF THE COMPONENT PARTS OF THESE ORGANS THEIR FORMATION, INCREASE, AND REPRODUCTION. LECTURE VIII. Page 383. ORIGIN AND ATTACHMENT OF BRANCHES ; STRUC- TURE OF LIGNEOUS ROOTS; AND OF HERBACEOUS DICOTYLEDONOUS STEMS. LECTURE IX. Page 447. OF LEAVES IN THEIR UNEXPANDED STATE, OR AS THEY ARE CONTAINED IN THE GEM : IN THEIR EXPANDED STATE, OR AS CONSTITUTING FOLIAGE. LECTURE X. Page 5 13. OF THE PETIOLE. COMPOUND LEAVES. THE SITUA- TION, POSITION, INSERTION, DIRECTION, MAGNI- TUDE, AND EXTENSION OF LEAVES. ANATOMY OF THE LEAF. LECTURE XL Page 599. OF THE CUTICULAR SYSTEM OF LEAVES: USE OF THE CUTICULAR APERTURES. OF THE APPEND- AGES OF THE STEM AND LEAVES PUBESCENCE THORNS PRICKLES GLANDS PROPS t USES OF THESE APPENDAGES. EXPLANATION OF THE PLATES. PLATE I. Fig. 1. Section of the tubers of Orchis mascula: a. the new tuber; b. the old ; e. the base of the stem of the present year's herbage; d. the plantule or embryon; e. a sheath cover- ing the plantule ; /. the point of conjunction of the ves- sels of the stem of the old tuber, with those of the plan* tule and the new tuber; g. a root. (p. 159, 161). 2. A transverse section of the new tuber, to display the vessels injected, (p. 160). 3. A transverse section of the old tuber, to demonstrate the effect of the absorption of the nutritious fluids on the cells in the immediate vicinity of the vessels, (p. 160). 4. Longitudinal section of a potatoe : a. the central part or pith ; b. b. b. b. the cortical part ; c. c. c. points where the eyes or gems are situated ; d. point where the tuber was attached to the runner, (p. 163). 5. Bulbs of Crocus sativus; a. b. the new bulbs adhering closely to c. the old or parent bulb much shrivelled ; d. the radical plate of the old bulb. (p. 167). 6. Section of fig. 5 ; d. two cords of vessels, which separate from the general bundle immediately on entering the new bulb, and pass ,on to supply nutriment to the embryons e. which are forming on a. b. (p. 167, 169). 7. Bulbs of Ixia polystachia ; a. b. the bulbs bearing the pre- sent year's herbage, partly covered with c. the reticulated b 2 Xli EXPLANATION OF THE PLATES. coat of the old bulb d.; e. e. e. e. e. roots of the recent bulbs protruded between them and the old bulb; /. a young bulb appended by a runner, (p. 167, 168). Fig. 8. Bulbs of Colchicum autumnale decorticated ; a. b. the new bulbs ; c. c. a flattened process on each ; d. the old or pa- rent bulb shrivelled up ; e. the remains of its radical plate ; /. the remains of its flattened process. 9. Another view of figure 8. (p. 170, 178). 10. Section of fig. 8 ; a. the old bulb ; b. c. the new bulbs (no lines should have been drawn from these letters) ; d. d, the opaque portion of the new bulb ; e. e. semi-trans- parent portion ; /. g. points of conjunction of the old and the new bulbs h. h. herbage of the present season, (p, 170, 178). PLATE II. 1. Entire bulb of Lilium candidum. (p. 173). 2. Transverse slice of one of the scales of the bulb, fig. 1. to display the vascular bundles. 3. A. the bulb, fig. 1. denuded of its scales, to show the young bulb : . the stem of the present year's flower; b. the young bulb; *c. c. fragments of the scales of the parent bulb; d. d. d. d. roots of the parent bulb ; e. e. remains of those of last year's bulb. B. Section of A.: a the remains of the caudex of last year's bulb ; b. caudex of this year's bulb. (p. 175). 4. Bulbs of Lilium superbum: a. the bulb of the present year ; b. that of last year ; c. the remains of that of the year be- fore last ; d. the embryo of next year's bulb ; e. e. e. e. roots of the bulb; /./. roots of the stem. (p. 175.) 5. The bulb, fig. 4, denuded of its scales : a. the caudex of the present year's bulb ; b. succulent .runner bearing the em- bryon bulb ; c. d. old decaying bulbs ; e. base of the stem of this year's herbage;/./, roots of this year's bulb. (p. 175). 6. Section of fig. 4.: . the caudex of this year's bulb; b. EXPLANATION OP THE PLATES. the base of the stem ; c. the runner ; d. the embryon bulb ; e. the place where the runner of the adult bulb has sepa- rated ; g. roots, (p. 176). Fig. 7. A scale of a bulb of Lilium pomponium: a.b. two young bulbs formed on it. (p. 176). 8. A portion of a scale of Lilium superbum, with a bulb formed on its margin; a. root of the young bulb. (p. 176). 9. Congregated bulbs of Saxifraga granulata ; a. the bases of the leaves and stem. 10. a. The plantule taken from a bulb of fig. 9. ; b. the plantule in a state of vegetation, (p. 178). PLATE III. (in two parts.) 1. A transverse section of the bulb of Narcissus Jonquilla : a. a. two young bulbs rising between the second and third layers ; b. the roots protruded from the radical plate which is hid by the position of the bulb. (p. 180). 2. A bulb of Allium victorialis: a. the outer layer of the bulb extended into the sheathing stem ; b. the reticulated layers ; c. the caudex; d. d. the roots, (p. 182). 3. A vertical section of the bulb, fig. 2. ; a. the succulent layers extending into the stem ; b. the old reticulated layers ; c. the caudex ; d. d. the young bulbs, (p. 183). 4. A tulip bulb : a. a portion of the flower stalk ; b. the new flower bulb ; c. c. the exterior tunic of the exhausted bulb ; d. a lateral leaf bulb. (p. 182). 5. Transverse section of a tulip bulb ; a. the base of the flower stalk ; b. remains of the layers of the exhausted bulb ; e. the new bulb. (p. 180). 6. (Middle of the plate.) Longitudinal section of a tulip bulb; a. the flower stem ; b. b. b. tunics of the exhausted bulb : c. the young flower bulb in its tunic d. d. d. ; e. the point of conjunction of the new bulb, with the parent, (p. 182). 6, 7. Bulbs of Hyacinthus orienialis: a. the body of the bulb; b. the manner in which the layers overlap one another ; c. b3 EXPLANATION OF THE PLATES. the radical plate; d. young lateral leaf bulbs; e. (fig. 6.), a bulb which has produced leaves this season, formed with- in the coats of the parent bulb. (p. 183). Fig. 8. A longitudinal section of fig. 6. : a. the caudex; b. the young bulb ; c. c. the remains of this year's plant, (p. 184). 9. A transverse section of fig. 6. (bottom of the plate): a. the young bulb ; b. the old or parent bulb ; c. c. roots, (p. 184). 10. A bulb of Allium sativum, denuded of its outer or sheathing tunics: a. the young bulbs; b. c. d. e. remains of the sheathing layers;/, the caudex and roots, (p. 187). 11. A longitudinal section of fig. 10.: a. the caudex; b.c. d. the sheathing layers which terminate in leaves, (p. 186). 12. A. The bulb of Fritillaria Pyreniaca : a. a. the two hemi- spheres, which are conjoined at the caudex ; b. the flower stalk rising between them ; c. the roots : B. a. a. one of the hemispheres separated from the other; b.b. the young bulbs; c, the flower stalk ; d. the roots, (p. 172). PLATE IV. t. A twig of Cullumia ciliaris. (p. 267). 2. A twig of Acacia decipiens. (p. 267). 3. A twig of Acacia alata. (p. 267), 4. A portion of the stem of Ruta graveolens. (p. 265). 6. The flower bud of CEnothera grandiflora, which is macu- lated, (p. 266). 6. A perfoliate stem, as demonstrated in Chlora perfoliata, (p. 267). 7. The stem of Lathy rus latifolius; illustrative of the winged and the stipulated stem ; a. a. a. a* stipules (p, 267). 8. A portion of the stem of Passiflora quadrangularis : a. a. tendrils (p. 653); b.b> stipules; c.c.c. glands. 9. A portion of the stem of Lilium superbum: a. a. bulbs, (p. 268). 10, A portion of the stem of Cactus flagellifonnis. (p. 268). EXPLANATION OF THE PLATES. XV PLATE V. Fig. 1. Longitudinal section of Agaricus procerus: a. the stem; b. the lax pith ; c. a fragment of the pileus. (p. 293). 2. Portion of the stem of Ptychosperma gracilis (from Mirbel): a. exterior ligneous and vascular bundles; I. interior lig- neous and vascular bundles, (p. 299). 3. A. A transverse slice of the stem of Trigridia pavonia, of the natural size : B. a portion of the same magnified, (p. 294). 4. A longitudinal section of the scape of Typha latifolia. (p. 294). 6. A. A transverse slice of the stem of Allium molly. B. a portion of the above slice magnified ; a. the cutis ; b. vascular bundles : C. a magnified vertical section of the above: a. the cutis; b. vascular cords ; c. c. cellular mat- ter: D. the natural size of the part, of which C. is one half, highly magnified. 6. Portion of a transverse slice of the stem of Juncus conglo- meratus, highly magnified ; a. the pith ; b. the cortex ; c. d. bundles of spiral vessels ; e. air cells dividing the vascular cords ; /. bundles of proper vessels, forming the striae on the surface of the stem : g. g. lacunae between the cortex and the pith. 7. Section of a transverse slice of the stem of Hippuris vulgaris, highly magnified: a. the exterior or cellular part; b. the centre, which is vascular; c. an uncoiled vessel. 8. A. A transverse slice of the culm of Triticum astivum, slightly magnified. B. a vertical section of one side of the tube, highly magnified: a. longitudinal cells, under the epidermis; b. the pith; c. a spiral vessel. C. a transverse section of a portion of the above slice, highly magnified : a. the cutis; b.c. vascular bundles, (p. 301). 9. Vertical section of a portion of the stem of Tradescantia virffinica: a. a. a. a. a. a. a. vascular cords; b. b. fragments of the base of the leaf, which in this plant is sheathing. b4 EXPLANATION OF THE PLATES. Fig. 10. A small portion of the above, highly magnified, to display the nature of the vessels : a . the kind of sheath in which the rings forming the vessels, b. b. b. are enclosed. 11. The articulation of the culm of Triticum testivum split: . the lower joint terminating in a sheathing leaf; b. the new joint, (p. 300). 12. The cuticle of the culm of Triticum cestivum, to demonstrate its respiratory pores, (p. 302). 13. The cuticle of Juncus conglomerate: a. a. portions which cover the striae, and which are devoid of respiratory pores ; b. portion between the striae thickly studded with pores, (p. 302). 14. Transverse slice of an apple twig (copied from Mr. Knight's plate in the Philosophical Transactions} ; a. the bark ; b. the alburnum or young wood ; c. the pith. 15. A terminal twig of ^Esculus Hippocastanum split down the middle ; a. the pith ; b. the wood ; c. the cortex ; d. e. buds or gems ; /. g. the bases of the petioles of two leaves, which are cut off; h i. cords of vessels connecting the pe- tiole with the stem ; k. terminal leaf bud. (p. 316.) PLATE VI. 2. A. A thin transverse slice of a twig of ^Esculus Hippocas* tanum: B. the dark-coloured portion of the slice A, sepa- rated, and very highly magnified; a. the cuticle; b. ex- terior layer of the cellular integument ; c. interior layer of the cellular integument; d. vascular layer of the cortex c. liber ; * alburnum in its first stage ; /. perfect wood ; g. a divergent layer; A. large vessels of the wood; t. medullary sheath; k. pith. (p. 319 406). 3. A transverse slice of a twig of Sambucus nigra in the first year of its growth, (p. 336). 4. Slice of a two years' old branch of Sambucus nigra. (p. 336). 5. Slice of a three years' old branch of Sambucus nigra. (p, 336). EXPLANATION OF THE PLATES. XV11 Fig. 6. Longitudinal section of a portion of the stem, and a branch of Sambucus nigra; a. the upper part of the trunk, in which c. the pith, occupies the centre ; b. the branch ; d. its pith; e. the wood produced by the branch; /./. two abor- tive buds. (p. 368). 7. A vertical section of the dark portion of A . fig. 2. : a. the cortex, consisting of, 1 . the cuticle ; 2. the cellular integu- ment ; 3. the vascular layer, in which the character of the proper vessels is displayed ; and, 4. the liber or inner bark ; b. the half-organized alburnum ; e. to the Polygons, contain acrid and several deleterious 3 LKCT. I.] SCIENCE OF BOTANY. 21 species. It also informs us, that some plants are acrid and poisonous when growing in water, which appears to be their natural element, although they are inert when they vegetate on dry land; and that some inert land plants become acrid when they accidentally spring up in water or in marshy places. Do we wish to discover the probable me- dicinal properties of the plants in any new situa- tion, before we venture to try their effects upon the animal frame? Botany informs us how to do so, by arranging the plants with which we are un- acquainted, into their natural families. Thus we know that the Solanece are narcotic ; the Gentiance yield a bitter, and sometimes a purgative principle; the Laurel tribe a stimulant, which is in some in- stances highly deleterious; the Gorymbiferce are emmenagogue; the Rubiacea?, to which Cinchona belongs, diuretic and tonic; the Crucif'erce antiscor- butic; and the Malvaceae emollient. A medical Bo- tanist, believing that a certain plant yields a pecu- liar medicinal principle, is led to examine whether the species of the same genus which are indigenous to the clime that he inhabits may not contain some- thing similar, if not exactly the same ; and thence discoveries valuable to his country, and sometimes to the human race, are effected. But this import- ant method of generalizing can be practised only by the Botanist; for, to one ignorant of Botany, not only is the language foreign and unintelligible, r 3 22 UTILITY OP THB (JLECT. I, but the resemblances which characterize the va- rious members of the same family, and which are perfectly obvious and striking to the Botanist, are overlooked and cannot be perceived by an ordinary observer. The advantages of a knowledge of Bo- tany, also, and of the habits of plants, to the phy- sician, are equally evident in the assistance they afford in the cultivation of a branch of the profes- sion, which has lately been much and properly at- tended to; I refer to medical topography ; a subject important to all, but absolutely requisite to the military medical practitioner. Suppose, for ex- ample, that an army is about to encamp in an enemy's country, and in a situation where circum- stances may require that it should remain for a considerable time. The season of the year, and the kind of weather prevailing at the moment, may render it difficult for the medical staff to pro- nounce whether the place be healthy or otherwise; ^nd the information afforded by experience is too }ate to prevent the impending evil, should it prove unhealthy. But the very plants which cover the soil clothe with a prophetic character the Botanical physician, and enable him to anticipate the danger which it is requisite to avoid. Yet how little has this branch of study been attended to in the edu- cation of professional men; who, before they pre- sume to commence the performance of the duties expected of them, ought at least to be acquainted LECT. I.] SCIENCE OF BOTANY. 23 with the nature and qualities of the implements they are about to employ in the cure of diseases. A remark made to me by an old and respect- able physician, the late Dr. Denman, explains in some degree the cause of this neglect: " I be- " lieve/' said he, * that the practitioners of the " present period are sensible of the inadequacy " of the old method of bringing up young men " to the profession, and are therefore anxious to *' send out those brought up under them better " instructed, and with a firmer foundation of prin- " ciples than was the case forty years ago : but " the ardour of youth to take an early, active " share in the bustle of life, places many obstacles " in the way of accomplishing their good inten- " tions." The justness of the observation is too obvious ; and as medical men, after they have once entered upon the busy stage of life, cannot well retrace their steps, happy would it be for stu- dents were they to take advantage of the expe- rience of their predecessors, all of whom, I will venture to assert, believe that much anxiety, much trouble, would have been saved to them- selves, and much more pleasure and reputation gained in the exercise of their profession, had a few more years been spent in their education, and a few more collateral studies been attended to during that period. The practice of medicine is the study of a lifetime ; it is entered upon the moment a man c 4 24 UTILITY OF THE [bECT. I. begins to prescribe, and it must be unremittingly continued*; but little satisfaction can be derived from it, if the mind be not previously well stored with principles, and enlightened by a liberal edu- cation. By the force of natural genius and a good address., an ignorant physician may float for a while in honour's atmosphere ; if fortunate, he may even be respected, and may attain wealth and reputation ; but, when an unexpected difficulty oc- curs, his deficiencies appear, the bubble bursts, his reputation is dissipated, and he sinks neglected and despised. The properly educated physician, on the contrary, may rise slowly at first ; but, like the sun, his strength increases as he rises ; and^ although he must naturally decline with gathering years, yet, when he sets, it is with equal dignity, only with a milder lustre. And surely, to use the language of a celebrated moralist, " to desire the " esteem of others for the sake of its effects, is not " only allowable, but in many cases our duty; " and to be totally indifferent to praise or censure " is so far from being a virtue, that it is a real " defect in character "f-." Botany is one of those collateral sciences, which is not only useful, but * The following words of Linnaeus, contained in a letter to Haller, should be impressed on the mind of every student: " Disco adhuc ; ignoscas quod doctus, etiamnum non eva. " serim." f Dr. Blairj Sermon VI, vol. ii. LECT. I.] SCIENCE OF BOTANY. 25 adds grace to the medical character. Did I wish to select examples in support of this remark, I need only point to the works of Prosper Alpinus, Sir Hans Sloane, Malpighi, Haller, Alston, Lewis, and of our own contemporary, the indefatigable Orfila. A practitioner, indeed, unacquainted with Botany, may know the names of many plants and their uses ; he may even gain a knowledge of the physiognomy of a few of them ; but his ideas are obscure and confused; his ignorance may often be rendered conspicuous where he would most desire to conceal it ; and it lays him open to the arts of the designing, and of those who would wish to ex- pose him. Of the advantages which the profession has derived from the labours of Botanists, I need mention a very few only of many examples that might be adduced : the reintroduction of the Fox- glove by Dr. Withering, as a remedy for dropsies, and the recent extension to this country of the Pyrola umbellata, and the Gurn Acaroides. As many of the medicinal plants appear as common weeds, a medical man ought to be able to distin- guish these when required ; and, in the case of ve- getable poisons, nothing will sink him more in the opinion of others, than his appearing ignorant of the plant which has occasioned the mischief; while nothing will raise him more in their esteem than his being able to point out its distinguishing characteristics^ by which it may be known and 26 UTILITY OF THK [LECT.-J. Avoided in future. But a more important consider- ation still, to a reflecting mind, is, that by the degree of acquaintance which a practitioner has with plants that are poisonous to the animal eco- nomy, the life of a fellow-creature may be lost or saved. All poisonous plants do not produce the same effects, and these, consequently, require dif- ferent modes of treatment ; but if the plant which has caused the mischief cannot be ascertained, how is the remedy to be selected ? The utility of Botany to many of the other arts is not less obvious ; and we are indebted to it for a variety of our comforts, both as to food and the luxuries of life. The grains so indispensable for pur existence, the greater number of the fruits, and the most beautiful flowers, that enrich our orchards and ornament our gardens, are of foreign origin ; and many of them have been brought to us by Botanists whose inquiries had led them to visit remote countries. The Horse Chestnut, for example, now so common in our plantations, was conveyed to Europe from the north of Asia, by Clusius, a Botanist, in the year 1550. The Kidney Bean, Phaseolus vitlgaris, was brought from the East Indies : and the Nol-kol, the root of which affords a large supply of wholesome nutriment, has just been introduced from the same place. The Crown Imperial, Fritillaria imperialis, was transported from Constantinople ; the Camellia, 4 LECT. I.] SCIENCE OF BOTANY. 27 from Japan ; many of the Roses from China , the Nasturtium from South America ; and the Pelar- gonium, or Geranium, as it is improperly called, from the coeist of Caffreria. The Potatoe, the chief support of a great majority of our poor, was first described by Caspar Bauhiri in 1590 ; and afterwards brought into this country, whence it was dispersed over Europe*. In our own times we have seen the West Indies enriched with the Bread-fruit by the scientific skill of Sir Joseph Banks ; and every day new plants are brought home and naturalised to our climate, of great im- portance both in an economical and political point of view *f~. Even the arts may be benefited by a knowledge of Botany. Thus, many fine statues might have been preserved, had the fact been sooner * It is very generally believed that this useful vegetable was brought from Virginia by Sir Walter Raleigh ; and Willdc- now states that, " in the year 1623, he distributed the first " which he brought from Virginia in Ireland." Doctor Smith Barton has pointed out the errors of this statement : in the first place, Sir Walter never was in Virginia ; secondly, he was not living in 1623; having lost his head in October 1618; and, thirdly, it is by no means certain that the first exclusive depot of the Potatoe was Ireland. f The number of plants now known and systematically ar- ranged amounts to 44,000; although those known by the Greeks, Romans, and Arabians, did not exceed 1400; and in Caspar Bauhin's time, all that indefatigable Botanist could col- lect for his Pinax Theatri Botanici, a work of forty years' la- bour, did not exceed six thousand species. 28 UTILITY OF THE [LECT. I. known, that the black spots which appear upon them is a vegetable fungus, the Lichen niger of Linnaeus. The dry rot in wood, also, is a fungus. It is related that Mahomet Bey, King of Tunis, was dethroned by his subjects for having the repu- tation of possessing the philosopher's stone. He was restored by the Dey of Algiers, upon promising to communicate to him the secret. Mahomet sent a plough, with great pomp and ceremony, in- timating that agriculture is the strength of a na- tion, and that the only philosopher's stone is a good crop, which may be easily converted into gold. I mention this anecdote, because it conveys an impressive idea of the importance of Agricul- ture ; between which and Botany the connexion is so natural, and the advantages to be derived by the farmer from a knowledge of it so apparent, that the neglect of it as an essential part of his education is, indeed, wonderful. But so blind are men, often, to their true interests, that agriculture in this country has, till within a few years past, been regarded as an employment fit only for the most uninformed part of society. Following stu- pidly in the footsteps of his predecessors, and guided by a few rules, which had been handed down to him from the rudest ages, the agriculturist was ignorant that a knowledge of the theory of his operations was necessary for enabling him to overcome unexpected obstacles ; to guard against LECT. I.] SCIENCE OP BOTANY. 29 the uncertainty of seasons ; and to multiply the means of supplying the wants of the community and enriching- himself. These mistaken notions are now happily vanishing ; and the utility of Che- mistry and Botany is beginning to be felt and ac- knowledged by the farmer. From the more ge- neral study of the economy of vegetable life, how many improvements might be suggested in the cul- tivation of even the most common of our useful plants ! How many of those which are still re- garded as useless weeds, might be found to be of great importance if their properties were accu- rately investigated! And certainly the nature of soils would be better and more easily known by an acquaintance with the different kinds of plants which each variety of surface produces. But, besides these objects of utility, to the ad- vancement of which Botany unquestionably con- tributes, it is likely to render a still greater service to social life, by the cultivation of that branch of it which assigns to each tribe of plants its altitude, its limits, and its climate ; and which the French designate by the tei-mGeographie Botanique. By its aid nature may be in some degree subdued by art, the mountains of Europe may be girt by the Cin- chona, the vine may cluster upon our rocks, and the high Palmetto and Coco wave on the sun- ward sides of our native vales. These examples are sufficient to show the uti- 30 UTILITY OP THE [LECT. I. lity of Botany in advancing- many of the more im- portant arts ; and the study of it is no less bene- ficial as a branch of general education. If the intellectual enjoyments of a well-informed mind be a valuable possession, whatever can augment these must be considered as of great importance. Many branches of knowledge have this effect; and, although they cannot be considered as directly advancing our interests or fortune in our inter- course with mankind, yet the possession of them affords a more permanent satisfaction than either wealth or honour can bestow*. Botany is one of * If we apply these observations to the fair sex, we shall perhaps immediately hear of the danger of producing literary women, of having wives that would leave the management of their houses to pore over the page of the philosopher, and be solving a mathematical problem instead of making a custard. We shall be told that learned women are insupportable, and use their acquirements as tyrants do their authority, making them weapons of oppression rather than the instruments of happiness. But a female pedant and a woman with a well-in- formed mind and liberal education are two different beings ; and I would answer remarks such as these in the words of one of the sexf, which I think are completely convincing. * c The fragile nature of female friendships," says she, " and " the petty jealousies that break out at the ball-room, have " been, from time immemorial, the jest of mankind. Trifles " light as air will necessarily excite, not only the jealousy, but " the envy of those who think only of trifles. Give them " more employment for their thoughts ; give them a nobler f Mrs. Barbauld. LECT. I.] SCIENCE OP BOTANY. 31 these; and the pleasure to be derived from the knowledge of it is not confined to any period of life, or any rank in society. In youth, when the affections are warm and the imagination is vivid ; in more advanced life, when sober judgment as- sumes the reins ; in the sunshine of fortune and the obscurity of poverty ; it can be equally en- joyed. The opening buds of spring, the warm luxuriant blossoms of full-blown summer, the yel- low bower of autumn, and the leafless, desolate groves of winter, equally afford a supply of men- tal amusement and gratification to the Botanist. I have thought it necessary to state these ex- amples of the usefulness of > Botany and the plea- sures to be derived from the study of it, in order to satisfy the demands of those who consider that nothing ought to be attended to which does not present some immediate object of profit or of uti- " spirit of emulation; and we shall hear no more of these paltry " feuds : give them more useful and more interesting subjects " of conversation, and they become not only more agreeable, " but safer companions for each other." I would add, that men who exclaim against learned women, often allow the sex an excessive and unrestrained pursuit of pleasure ; and think so meanly of their powers of mind, as to believe them fit only to be amused with the fictions of romance. But, if pleasure be essential for the happiness of woman, it can be obtained from the pursuit of science, that knowledge which is founded on truth and utility, provided an early taste for it be implanted in the mind. 32 GENERAL DIVISION [LEOT. I. lity. What is the use of Botany? What can you gain by it? are constant questions. We have al- ready stated our opinion as to what may be gained by it ; but I would further answer such inquirers by an anecdote told of the Greek philosopher He- raclitus. This philosopher was very poor, but much respected, and visited by individuals of the highest rank. One day, when certain persons came to consult him, they found him paring tur- nips for his supper, and warming himself in a kitchen: the meanness of the place occasioned them to stop ; upon which the philosopher thus accosted them: " Enter/' said he, " boldly, for " here, too, there are gods." We have now to consider what mode of prose- cuting the study of the science upon which we are about to enter is likely to be productive of the greatest benefit. BOTANY, as we before observed, is that science which teaches the knowledge of vegetables, their structure, habits, and properties; and to distin- guish different plants from each other ; or, it com- prehends Phytology and Systematic Botany*. * Vegetable physiology, indeed, cannot be separated from systematic Botany ; for, to use the words of a French writer, M. Aubert du Petit-Thouars, " it is essentially the basis upon " which that science is raised ; and the more this basis shall be " known, the more Botany, already so attractive in herself, * will see the number of her disciples increase." LECT. f.] OF BOTANICAL SCIENCE. 33 Vegetables, like animals, are organized living bodies. To the superficial observer there appears no difficulty in distinguishing them from animals and fossils ; but those who have examined the sub- ject more minutely, find many obstacles to prevent them from drawing the exact line of distinction be- tween the three kingdoms of Nature. Still, how- ever, vegetables possess peculiarities of structure, habit, and functions, which characterize them; and these are found in every plant. As plants are living beings, so are they also perishable: death, as is the case in animals, may either proceed from innate causes, depending upon their organiza- tion, or be produced by external causes. It is ob- vious to our senses, that vegetables derive nourish- ment from the soil in which they are fixed, and in which they grow, and perfect seed capable of repro- ducing the species. The researches of philosophy have further informed us, that they possess irritabi- lity, by which the nutriment they imbibe is progres- sively moved through every part of their bodies, con- verted into various secretions, and assimilated into the substance itself of the plant; and that, like animals, they produce certain changes on the at- mosphere, and can accommodate themselves to the vicissitudes of heat and cold. In the func- tions of generation, also, plants have many of the peculiarities of the most perfect animals. In stat- ing, however, the close analogy between plants VOL. i. D 34 GENERAL DIVISION [LBCT. t. and animals, we must always bear in memory that they have one important function less than ani- mals sensibility *. The losing sight of these cir- cumstances, particularly the former, led the inge- nious Darwin into a labyrinth of error ; and has exposed his memory to the sarcasm of malevolent wit and the derision of ignorance. That part of our subject therefore, which refers to the economy of the vegetable system, should first engage the attention of the student. It constitutes Phyto- logy, and comprehends the Anatomy and Physio- logy of plants; and is the most amusing, and cer- tainly not the least instructive part of the science. The anatomy of plants is more difficult than that of animals, from the minuteness of their parts, the union of them, and the extreme difficulty of se- parating them without destroying their texture. If, however, it be more difficult, it is less disgust- ing, and the microscope very much facilitates our inquiries. Without it we can have no idea of the structure of plants, and consequently no correct notions of their functions can be obtained. In studying the Anatomy the Terminology is acquir- ed, an acquaintance with which is absolutely ne- cessary for securing a knowledge of systematic arrangement. By combining with these the study * " Vegetabilia, sensatione licet destituantur, aeque taroen ac *' animalia vivcre probat ortus> nutritio, v\v$uv (kotuledon), hollo\v. G 3 86 VASCULAR TEXTURE. [LECT. III. which exist in the stems of Grasses, and in Palms. They are numerous also in most herbaceous plants ; and particularly in aquatics of a lax texture. They are seldom detected in the root, and never in the bark ; but are situated round the medulla of the young shoots of trees and shrubs ; whence bundles of them are given off,, and enter the middle rib of leaves, to be distributed through them under their upper surface. They have been detected, also, in the calyx, or flower-cup, and other parts of the flower ; and Gsertner asserts that they are evident even in the seed-lobes. The spiral vessels, in their course, proceed always in straight lines, without any deviation; whereas all the other vegetable vessels often take a curved direction. It is into these vessels that coloured injections most easily enter ; and when an annual twig of the Fig is thus injected, they are seen in a transverse section of it, like red dots around the pith, placed within an external circle of the vessels, which contain the succus proprius, or milky juice of the plant. It cannot be affirmed that the varieties of form, which we have pointed out in the vegetable vessels, is of the same importance as the differ- ence which exists in the structure of the arteries and veins of animals. There are, indeed, some plants in which three of the modifications of struc- ture, according to Mirbel's observation, are found in the same tube. In the Butomus unibellatus, LECT. Hi.] VASCULAR TEXTURK. 87 Flowering Rush, says that author, "I have seen long portions of vessels present, at intervals, the appearance of an unrolled trachea, a transversely cleft vessel, and a porous vessel." Besides these vessels, Hedwig imagined there existed lymphatic, or absorbing vessels, opening upon the cuticle, and forming a circle round the exhaling pores : a doctrine, which has been adopted by Willdenow and others ; but which Mirbel justly combats, alleging " that the sides of the cells which terminate in the cuticle, and the fragments of which remain fixed to that pellicle when it is detached, have been mistaken for lymphatic ves- sels by Hedwig *." Mirbel mentions another set of vessels, which he denominates little tubes ; but they may rather be regarded as tubular cells, than vessels, being closed at the extremities. They resemble, very much, stretched cellular substance, except that the membrane composing them is less transparent, and of a greater consistence. The solidity of plants depends very much on the quantity and density of these cells, which are filled with thick and coloured, or thin and colourless juices, ac- cording to the nature of the plants in which they exist. Such is the vascular system of vegetables, * Elem. de Phys. veget. 1* Partic, p. 40. c4 88 GLANDULAR TEXTURE. [LECT. III. As we do not intend, at present, to enter into any account of the uses of the vessels we have described, we will only observe, that as all vegetables take up nourishment from the soil, and change it into juices different from each other, and which must be preserved from mingling together during the life of the plant, we might (a priori) suppose that plants must necessarily possess a vascular system : microscopic anatomy proves the fact, displays the numerous ramifica- tions, and general distribution of the vessels : ob- servation shows that their elongation increases the bulk and growth of the plant, and that they per- form the most important functions in the vegetable system. I may here also remark, that whilst the tubes, or vessels, which have been described, are intended chiefly for the longitudinal progression of fluids, the lateral transmission of the vegetable juices is performed, perhaps solely, by organized pores and slits, such as have been described in the perforated vessels, and in the sides of the cells. But the account of the manner in which this function is performed, must be postponed for our future consideration. The structure of the Glandular texture, as far as relates to the interior of the vegetable body, is much more difficult of demonstration than that of any of the general solid components which have been already noticed : but, when the im- 2 LECT. III.] GLANDULAR TEXTURE. 89 possibility of attaining an accurate knowledge of the glands of the animal body, which are large and visible to the naked eye, is considered, it will not appear wonderful that our remarks on this part of our subject are drawn rather from analogy than from actual observation. When, however, we reflect on the nature and diversity of the vege- table secretions, and that plants possessing the most opposite properties rise from the same soil, there appears to be no medium by which the absorbed aliment can be so altered in its characters, except by that of a glandular system. I am willing to admit that the simplicity of the vegetable structure is astonishing ; and that effects are pro- duced in plants, by means which are apparently very inadequate, when we regard them with a reference to the animal economy : yet, still, when the eye glances over the number and variety of vegetable products, there is much reason for sup- posing, that the simple transfusion of fluids can scarcely be sufficient for the production of these changes. We know that the laws of chemical affinity, in the temperature in which they take place, are inadequate to the effect ; and, besides, many of the changes produced, particularly those which fit the sap to be assimilated into the sub- stance of the plant itself, are directly contrary to the laws of chemical affinity, which operate in destroying these combinations, as soon as the vital 90 GLANDULAR TEXTURE. [LKCT. III. principle of the plant ceases to act. Although, therefore, we cannot by demonstration prove the existence of internal glands in vegetables, yet we have the strongest analogical proof in favour of the supposition that they do exist. The pores and clefts of the cells and the vessels which have been described are surrounded by opaque regular borders ; and even the flat thread which forms the spiral vessels is edged with a similar border. These bodies are regarded by M. Mirbel as glands ; and he conceives the opinion receives weight from the circumstance of the mucilage, which is changed into the organized tissue, being found always col- lected in greatest quantity around those vessels whjch are most studded with these opaque borders. This supposition is extremely probable, and is one of those which, if they cannot be confirmed, cannot be positively denied. If, as we suppose, vegetable glands exist, they must necessarily enter as a general component into the structure of every plant. Besides the glands, the existence of which in the interior of the plant, if not demonstrable is too probable to admit of much doubt ; there are also external bodies, which all Botanists have agreed in considering as glands, and which, in general, separate, as an excretion, some peculiar fluid *. Thus, honey, or a nectarious fluid, is * " Glandula," says Linnaeus, " est papilla humorem excer- nens.*' Philosophia Botanica, 84r,- 6. LECt. HI.] LJGN f EOU8 FIBRE. 91 secreted at the base of the petals or coloured floral envelopes, in the greater number of plants ; on the stalks of others, a viscid substance is thrown out ; and on some, little sharp bristles are planted, which are -perforated, and through which a very acrid fluid is ejected into the wound which they make in the cuticles of animals. Examples of the first are to be found in almost all flowers ; we observe the second in the species of the genus Silene, called Catch-fly, and in many other plants ; and the bristles of the Stinging Nettle supply a well-known instance of the last. Of the structure of these glands, although they are external, yet very little is known ; and microscopes of the greatest magnifying powers present them as masses of cellular substance only, with vessels passing on to their centre, without developing any other particular organization, which might lead to the formation of a theory explaining the mode in which they perform their functions. These, however, are, in some degree, obvious from their effects ; and afford more than probability to the idea that vegetables possess a glandular system. We shall have occasion to point out, and explain, the particular forms and functions of these external glands in different plants, when we come to examine the parts on which they are situated. The Ligneous Jibre is a very minute, firm, 92 LIGNEOUS FIBRE. [LECT. III. elastic, semi-opaque filament, which, by its cohesion with other filaments of the same kind., forms the proper fibres, or layers of longitudinal fibres, that constitute the grain or solid part of wood. It enters, also, into the composition of another set of layers, that traverse the longitudinal, named divergent. It is intended to give support and firmness to the vegetable body, and hence is found in greater abundance in trees and other perennial plants ; and according to the number of the ligneous fibres in each bundle of layers and the force of their cohesion, the wood of different trees possesses a greater or less degree of hardness. But, although wood is found of various degrees of consistence, yet, as Count Rumford has suggested, it is probable that the ultimate fibre is the same in all plants*. Whether the ligneous fibre be of original forma- tion similar to the muscular fibre of animals, or condensed membranous or cellular texture, or an obsolete obstructed vessel as Hedwig supposes, is yet undetermined. It is so intimately united with the cellular texture containing the vegetable secretions, that it cannot be procured pure for examination, without the aid of chemical agents to separate these adjuncts. If a thin shaving of well- dried wood be first digested in boiling water, then * Nicholson's Journal, vol. xxxiv. p. 319. LECT. III.] EPIDERMIS. 93 in alcohol, and lastly in ether, every thing soluble in it will be extracted by these liquids, and the insoluble part which remains be found to be com- posed of interlaced fibres, easily subdivided and having- some degree of transparency : these are the ligneous fibres. They have neither taste nor odour, and remain unaltered by exposure to the atmosphere : but although insoluble in water, alcohol, or ether, the fixed alkalies and mineral acids dissolve and decompose ligneous fibre. The relative quantity of this fibre in any plant may be pretty accurately ascertained, by exposing a given quantity of the wood to a moderate fire, in close vessels, for a number of hours sufficient to convert it into charcoal ; for, as the wood only becomes charcoal and the other parts are dissipated, the proportional weight of the charcoal obtained shows the quantity of the ligneous fibre contained in the wood. By experiments of this kind, carefully per- formed on the wood of the Poplar, the Lime, the Fir, the Maple, the Elm and the Oak, Count Rumford ascertained that the quantity of ligneous fibre in each of these trees was equal to nearly nine twentieths of their wood in its natural state *. The Epidermis is that portion of the vege- table structure which is exterior to all the others ; * Gilberts Annalen der Physick, - xiv. p. 25; and Thomson's System of Chemistry, 5th edit. vol. iv. p. 186. 94 EPIDERMIS. [LECT. in. at least to those which retain their vitality in the vegetating state of the plant : or it is that part which is interposed between the living organs of the individual, and all extraneous substances. In this respect it resembles the cuticle of animals ; but Botanists have been too fond of tracing this analogy, which has, not unfrequently, biassed their observations, and led to erroneous conclu- sions. It extends over the surface of every part of the plant; from that of the delicate petal of the flower, to that of the leaves, the branches, the stem, and the root; but, except in young stems and roots, it is not the exterior part of those organs of the plant ; the coarse rugged surface of older roots and stems being exterior to the real epidermis. It is common to every kind of plant, nor can we conceive that any one can exist without it. Botanists, as I have already stated, are very fond of drawing an analogy be- tween the epidermis of plants, and the animal cuticle ; and the resemblance, in many respects, is conceived to be closer than it really is ; but there is, nevertheless, in some circumstances a very striking analogy. The vegetable epidermis may be separated from the parts which it covers, by raising it cautiously with a knife ; but this is more easily effected by maceration and boil- ing. It is more readily separated from the cellu- lar substance it covers in the leaf, than in any LECT. III.] EPIDERMIS. 95 other part of the plant ; and for this purpose I would recommend to the student the leaf of any of the Lily tribe, before the stem shoots up ; or of the Lettuce (Lactuca sativa}, or that of Sorrel (Rumex acetosa) ; but even in these, some of the cellular matter is always detached in separating it ; and to this circumstance is perhaps to be at- tributed the variety of opinions which phytologists have advanced regarding its structure. The epidermis appears at first of a green co- lour on the young stems and branches of almost all plants ; but it changes to different hues, ac- cording to the age of the part it covers. Accord- ing to Du Hamel, it is composed of fine, but tough fibres, which are interwoven "together ; and every where interspersed with pores, which per- mit the mouths of the absorbing, transpiratory and air vessels to open to the atmosphere. Com- paretti also describes it as composed of fibres, in- terwoven so as to form hexagonal meshes, the areas of which are filled up with opaque or dia- phanous vesicles, inflated as if extended with air or water, and having a small black point in the centre. Mr. Bauer, on the contrary, conceives the structure to be altogether cellular, and vary- ing in different plants*. My own observations lead me to adopt the opinion of the elder Saussure, * Tracts relative to Botany. Lond. 1805. 96 EPIDERMIS. [LECT. in. that the true epidermis is a fine, transparent, un- organized pellicle *. The pores, by which the in- sensible perspiration escapes, are so minute, that they are quite invisible, and with difficulty permit the passage of air through them. Thus, if an apple be put under the receiver of an air-pump, and the air withdrawn, the cuticle of the apple will be lacerated by the dilatation of the air con- tained in the pulp of the fruit. There are oblong- pores also in the cuticle of herbaceous plants in particular, as was first observed by Decandolle, who named them cortical pores. The size of these is considerably greater than that of the former ; and varies in different plants. The epidermis seems to be entirely destitute of longitudinal vessels. In herbaceous plants, and in young and succulent twigs, it is, with a few exceptions, colourless and transparent ; the apparent colour being produced by that of the juices in the cellular substance immediately be- neath it, in the same manner as that of the hu- man cuticle is produced by the colour of the capillary web which it covers, and which is filled with different coloured fluids in different races of men ; white in the inhabitants of the temperate zones, and black, or brown, in those of the torrid * Obs. stir VEcorce des Feuilles. LECT. III.] EPIDERMIS. 97 regions of the globe *. When the epidermis is applied very closely to the cellular layer below it, which is the case in herbaceous plants, and in the young twigs of trees and shrubs, the greater por- tion of the light is transmitted through the cuticle, and reflected from the cellular layer, and not from the substance of the cuticle ; so that the colour of the herbaceous stem, or of the twig, is, in this case, that of the cellular layer, and not of the cuticle itself; yet in trees and shrubs, which an- nually renew the cuticle, as the Plane, the Birch, the Currant and many others, the epidermis, when it begins to peel off, becomes more opaque and does not transmit the light, but reflects it from its own surface. Thus the old cuticle of the Plane is dark coloured, while the new is of a light green hue ; the stem of the Birch, from which layers of epidermis are continually peeling off, is white, while the young branches are brown ; and the old branches of the Currant are dark brown, while the young shoots are of a very light green hue. In some plants, instead of being thrown off in * " On peut done concevoir le corps reticulaire comrae " un systeme capillaire general, entourant 1'organe cutane, et " formant avec les papilles une couche intermediate au corion " et a 1'epiderme. Ce systeme ne contient, chez la plupart des " hommes, que des fluides blancs. Chez les negres, ces fluides " sont noirs. Us ont une teinte intermediate chez les nations " basanees." Bichat, Anat. Gcnerale, $c. tome 4 me , p. 657. VOL. I. H 98 EPIDERMIS. [LECT. HI. plates^ or in layers, the old cuticle is reduced into powder* Although the epidermis is not cast off from all plants in this manner, yet, it is constantly renewed ; and, where it remains, the old cuticle cracks as the diameter of the stem of the tree, or of the branch, increases: it is then gradually pushed outwards, and the accumulation of suc- cessive layers, in this manner, forms the rugged coats which characterize many trees^ as the Elm and the Oak. This renewal of the epidermis in vegetables is similar to what takes place in ani- mals. The snake, for example, casts his skin an- nually, as do also the crab, the lobster, the spi- der and many other insects ; and the cuticle of the human body often peels off, particularly after some diseases, as scarlet fever for instance, and on the application of acrid matters to the skin. In animals, however, when destroyed, it is again regenerated ; but in vegetables, this occurs on the stems and branches of perennial plants only ; for on annual plants, and on the leaf and flower, it is not renewed after being destroyed. The ve- getable epidermis is capable of extension ; but this is less considerable than has been supposed ; and as there is a constant renewal, there must be a proportional increase or growth of its parts, so that it is not simply extended to enable it to cover LECT. III.] EPIDERMIS* 99 a greater portion of surface ; but new cuticle is added to produce this effect. The use of the epidermis is to keep the parts beneath it together ; and to regulate the perspira- tion and absorption of the plant. It is calculated also to defend the parts it covers from humidity; for which purpose, it is covered with a waxy secre- tion. The powers of the cuticle in regulating these functions is fixed according to the nature of the plant. In succulent plants, which require much moisture to be retained in their leaves, the cuticle is so constructed as to assist absorption, but rather to prevent transpiration. Thus, if a leaf of the Aloe be cut off, it will remain a very long time, even when exposed to the sun's rays, before it shrivels ; but, if in this state it be exposed to damp air, or thrown into water, the absorption is so rapid that it will regain its original plumpness and size in a few hours. A fine proof of that overruling Wisdom which fits every thing for the fulfilment of the purposes of its creation : for, in this instance, the Aloe being a native of a dry arid soil, and a warm climate, it could not long exist if the per- spiration from the surfaces of its leaves were con- siderable, but on the contrary it draws a great deal of moisture from the atmosphere by absorption. Another use of the epidermis is to prevent the destruction of the parts it covers : for, as it is in the vessels of the inner bark, as I will afterwards H 2 100 EPIDERMIS. [LECT. nr. demonstrate, that the greatest activity, irritability, and degree of vital energy reside, if that part be wounded to any considerable extent, so that the external air gets access to it, exfoliation, and the death of the part, and sometimes that of the whole plant, follow ; the cuticle forming, as Sir J. E. Smith elegantly expresses it, " a fine, but essential 66 barrier between life and destruction*." Mirbel-f- combats the idea of the epidermis being a distinct organ, and supposes it to be the external layer of the cellular membrane con- densed, and altered by exposure to the air and light. But although I admit that the cuticle be nearly the same as the parietes of the cellular tissue which it covers, yet, it is nevertheless a distinct organ. The simple exposure of the cellular mem- brane will not form epidermis ; but, on the contrary, when the cellular substance is exposed, it is more apt to exfoliate ; and when the wound becomes healthy, it is then only that cuticle is reproduced. During this process the new epidermis proceeds from the sides of the wound, gradually extending over it, in the same manner as in a wound of the human body. The very close connexion of the epidermis and the cellular substance can be no argument against our opinion ; for, although the * Introduction to physiological and systematical Botany, 2dedit. p. 18. f Eltmem de Pkys.^veget. l ere Partie, p. 35. LECT. III.] EPIDERMIS. 101 flakes of epidermis which are cast off annually by the Plane tree, and some other trees, consist of cellular substance also, yet, the cuticle is already formed under the flakes before they fall, and therefore it cannot be in these cases produced by the action of the air, and light. On examination of these flakes, the epidermis appears to be dis- tinct from the plate of cellular substance which separates with it. Mirbel himself, indeed, is obliged to modify his objection, and adds, But, although the epidermis of vegetables does not resemble that of animals, and is certainly the external part of the cellular substance, yet, it is no less true that secondary causes modify its nature, and it consequently becomes an organ the functions of which are very distinct and im- portant *. Such a concession is all that can be demanded. That the human cuticle is a distinct organ has never been denied, and yet we know that it is equally without vessels and nerves. On the same principle, therefore, the vegetable epider- mis must be admitted to be a distinct organ ; and * " Mais, quoique Pepiderme des vegetaux ne ressemble " pas a celui des animaux, et qu'il soit forme certainement par " la partid exterieure du tissu cellulaire, il n'est pas moins vrai " que des causes secondaires modifient sa nature, et qu'il " devient par le fait un organe dont les fonctions sont tres- " distinctes et tres-importantes." Physiologic vegetate, vol. i. p. 89. H 3 102 EPIDERMIS. [LECT. in. the modifications of it in different plants are more numerous than among animals. The illustration of these varieties must be deferred, till we come to consider the structure of the bark, of which the epidermis is usually considered a layer. Such are the principal solid components of the vegetable body. Other solid matters certainly enter into the structure of some plants ; but, as those are not common to the vegetable race, they cannot be classed amongst the general compo- nents. Perhaps, also, all the parts which have been examined may be resolved into modifications of the membranous and cellular textures, which might consequently be regarded as the only real solid vegetable components ; but, although we allow that the vessels, ligneous fibre, glands, and epidermis most probably are composed of mem- branous, or cellular texture, differently modified, yet, as each of these parts possesses very distinct functions, such a refinement would only throw obstacles in our way towards the attainment of truth; I have, therefore, considered it preferable to regard them as distinct components. LECT. III.] THE SAP. 103 GENERAL FLUID COMPONENTS. Vegetables, by their vital energy, increase in bulk, and augment the quantity of solid matter they contain, consequently the principles of the solids must be contained in the particular fluids which they select and imbibe from the soil ; but in what manner the fluids are changed into solids, and whether any of the solid matters be taken up ready formed, or whether they result from a trans- formation effected solely by the action of the vegetable vessels ; are subjects of consideration upon which, in the present stage of our inquiries, it would be premature to enter. These fluids, however, after being absorbed by the roots, enter into and fill the cells and vessels of the plant, and form a very considerable portion of the bulk of the vegetable body. As soon as they enter the plant, they constitute its SAP, or common juice, to the nature of which, as it is one of the general components of vegetables, we must now direct our attention. Were we about to examine the moving powers by which the fluids selected from the soil, and absorbed by the roots of plants, are carried for- ward through the vessels, we would demonstrate that although these moving powers operate at all times during the life of the vegetable, yet, that H 4 104 THE SAP. [LECT. in. their action is most energetic in spring and at midsummer, at which periods, therefore, a much greater quantity of fluid is found in the vegetable vessels. As, however, the simple examination of the sap itself is our present object, it is sufficient to state the fact ; and to know that, at these sea- sons, when an incision is made through the bark and part of the wood of most kinds of trees, or a hole is bored in the trunk, a fluid exudes in con- siderable quantity. This is the sap, or common juice. It is in the same situation, for the purposes of the plants, as the chyle of animals is, while it is yet in the thoracic duct, and before it is mingled with the blood, and exposed in the lungs to be fitted for the purposes of the animal. Neither is in a proper state for yielding the various secretions, and adding, by the process of assimilation, to the growth of the plant, or of the animal ; but the analogy goes no farther. In the animal, the di- gestive powers of the stomach and the action of the mesenteric glands so change the food taken into it, that no chemical analysis of the chyle produced from it could lead to an accurate know- ledge of the kind of food, which had been em- ployed by the animal ; but, in plants, the food is already prepared in the ground before it is ab- sorbed by the roots, and, therefore, were it pos- sible to obtain the sap from the vessels very near to the extremities of the roots, we should be en- LECT. III.] THE SAP. 105 abled to discover, with a considerable degree of accuracy, the real food of plants *. This, how- ever, cannot be accomplished ; and as the sap, in its progress, dissolves some ready-formed vege- table matter, which had been deposited at the end of the former autumn, in the upper part of the root and at the base of the stem, its original pro- perties are thus altered ; and the farther the part, which is bored in order to procure the sap, is from the root, the more vegetable matter this fluid is found to contain. This fact was first no- ticed by Mr. Knight, who ascertained that, owing to this deposition, the wood of the stem, and the large branches of trees, have a greater specific gravity, and contain more soluble extractive mat- ter when cut down in winter, than in spring, or early in summer : on this, account, although there is reason for believing that the food of almost all vegetables is the same, yet, the sap, in the state in which we can obtain it, differs in different species of plants ; and, therefore, no just idea can be formed of its nature from the most accurate analysis of it, when procured from any single plant. Were it possible, however, to obtain the * It has been supposed that the roots of plants absorb, in- discriminatelyj all the soluble matter contained in the soil on which they grow ; but were this the fact, many more sub- stances would 5e found in the vegetable body, than have yet been discovered in it. 106 THE SAP. [LECT. HI. sap completely free from the peculiar juice of the plant, it would be very probably found nearly the same in all plants. From an examination of it, such as it can be obtained, we are enabled to draw some general conclusions; and by comparing the analysis of the sap of many different plants together, we discover those components which are most frequently present, and consequently form an opinion, approximating to the truth, of the real nature of the sap. When a tree is wounded, in the manner we have described, in the spring, it is said to bleed ; and if the wound be considerable, and in the principal stem, the tree being thus drained of its fluid, soon dies *. Any quantity of sap may be collected by this mode of wounding trees, and the * When we reflect how early this fact must have been known, it is remarkable that so. little progress has been made in developing the power by which the sap is carried forward through the plants. The rudest nations are acquainted with the fact that trees bleed when wounded ; and from a know- ledge of its consequences, the Asiatic nations, in their wars, commit the greatest injuries which their opponents can suffer. The Palms in Asia being as necessary for supplying the ordi- nary food of the natives, as grain is in Europe ; when an hostile army enters the territory of an enemy, they cut notches with hatchets in all the Palms which they meet with ; which occasions the sap, and the other juices of the plants to run out ; and the Palms either die altogether, or are rendered abortive for that season. LECT. III.] THE SAP. 107 less progress vegetation has made, it is obtained in a purer state. It should, therefore, be drawn very early in the spring, before the leaves expand, and as near to the root of the plant as it can easily be obtained, if we wish to examine its chemical properties. When the sap is thus drawn from a tree, it usually appears nearly as colourless and limpid as water, has scarcely any taste and no particular odour. A phial containing a certain quantity of sap weighs heavier than the same phial, contain- ing an equal portion of distilled water ; so that the specific gravity of sap is greater than that of water. If it be kept for some time in a warm place, it undergoes sometimes the acetous, at other times the vinous, and in some instances even the putrefactive fermentation. These differences would indicate a great disparity in the compo- nents of the sap of different vegetables ; but there is every reason for thinking that they depend more on the admixture of the proper juices, which, as I have already stated, are always more or less mixed with the sap, as we can obtain it ; and it is probable, that the sap of different plants differs more in the proportional quantity of these juices mixed with it, than in the nature of its components. The rapid vinous fermentation of some kinds of sap is taken advantage of in warm climates for economical purposed. From the top of the Cocoa-nut tree 108 THE SAP. [LECT. in. the natives of India extract the sap, mixed un- doubtedly with the proper juice, by making an incision with a sharp knife overnight, and suspend- ing under it a vessel to receive the fluid as it exudes. This liquor, next morning before the sun is hot, is a pleasant, mild, cooling beverage ; but before evening, it ferments and becomes pow- erfully intoxicating. In Ceylon, arrack is distilled from this fluid, which is named toddy ; ' and it, also, yields a coarse black sugar, called jaggery. As, however, in this case the sap is combined with the proper juice of the tree, the extraordinary effects of the rapid fermentation must, in a great degree, be ascribed to it. It is to the same cause also, as I before noticed, that we must ascribe the dif- ference in different saps, particularly the saccha- rine and acid qualities, which they sometimes pre- sent even when newly drawn. Thus we are told that sugar is extracted in the proportion of ten pounds from every two hundred of the sap of the Acer saccharinum,, Sugar Maple. But the sap is so mixed with the peculiar secreted juices of the plant, when it is drawn from the tree for this purpose, that it can scarcely be considered as yielding the sugar. According to the observations of Mr. Knight, sap always contains a considerable por- tion of air. It, also, differs in its specific gravity according to the distance from the root at which it is taken, the gravity increasing in the direct LECT. III.] THE SAP. 109 ratio of the distance, which appears in some degree to arise from the solution of deposited matter in its progress, but, perhaps, more from the transpi- ration of the plant throwing off a large proportion of the watery part of the matter taken up from the soil. Such are the sensible qualities of sap ; its chemical properties and composition are dis- covered by tests and analysis by heat. I shall first mention some experiments I made on the sap of the Vine, and then detail the results of those made on some other saps by Vauquelin a celebrated chemist of the French school. On the eighth of April, when the thermometer was at 68, and a few leaves of the Vine, which was the subject of experiment, had already ex- panded, I cut off the extremity of the lowest of the branches, and introduced the cut end of the part which remained fixed to the Vine, into a six ounce Apothecaries' phial. On the following morning, about three ounces of a clear, limpid, colourless fluid, like water, was found in it. It had no perceptible taste, except a slight mucila- ginous feeling on the tongue ; no odour ; and weighed rather heavier than distilled water. On pouring a little of the Tincture of Litmus into it, it was very slightly reddened, thereby indicating the presence of an acid *. The oxalic acid almost , * Dr. Prout, who examined the sap of the Vine, found that the specimen which he procured did not differ in specific gravity from pure water ; nor did it alter Litmus paper. 110 THE SAP. [LECT. m. immediately rendered it milky, and threw down a white precipitate. The acetate of barytes threw down a white flocculent precipitate; and by acetate of lead (Goulard's extract), a white curdy preci- pitate was also produced. No change was effected on it by the solution of ammonia, nor by that of gelatin : nor was any dark hue communicated by the sulphate of iron. The addition of alcohol to this sap threw down a light flocculent precipitate, of a mucous nature. The sulphuric acid added to it, occasioned a slight effervescence, and evolved the odour of acetous acid. The conclusions which may be drawn from these appearances is, that this specimen of the sap of the Vine contained acetate of potash with perhaps an excess of acid ; carbonate of lime, vegetable mucilage, some albuminous matter and water. As no effect was produced by the addi- tions of the sulphate of iron and the solution of gelatin, we conclude that it contained neither tannin nor gallic acid, and therefore possessed no astrin- gent property : a proof that the sap which we tried was pretty pure, for some of the secreted fluids of the Vine are both very acid, and considerably astringent. The small quantity of sap which was obtained, owing to the rather advanced state of the season, for such an experiment, prevented the pro- portions of the different ingredients from being ascertained. M. Vauquelin, as I have already mentioned, made the most interesting experiments, which LECT. HI.] THE SAP. Ill yet, been attempted on sap. As he does not, how- ever, mention the state of the trees, from which the sap for his experiments was taken, the result of the analyses affords some reason for believing, that the sap was not in the purest state. He examined the sap of the Elm, Ulmus campestris, collected to- wards the end of April, the beginning of May, and the end of May. The result of the first analysis was, that 1039 parts of this sap consisted of 1027-904 of water and volatile matter, 9-240 of acetate of potash, 1-060 of vegetable matter, and 0-796 carbonate of lime. The second analysis of the sap collected at the beginning of May afforded a greater proportion of vegetable matter, less acetate of potash, and also less carbonate of lime : and in the third analysis of that collected at the end of May, the quantity of the acetate of potash was still more diminished, and also that of the carbonate of lime *. In all he found slight traces of sulphate and of muriate of potash. From two different ana- lyses of the sap of the Beech, Fagus sylvatica, procured also at different periods of the same season, he obtained water, acetate of lime, free acetic acid, gallic acid, and tannin, with some vegetable extractive and mucous matter -}-. In the same manner he examined the sap of the Common Hornbeam, Carpinus sylvestris, collected in March and April, and found in it, acetate of potash, * Annales de Chimie, t. xxxi. p. 21. f Ibid, p. 26. 112 THE SAP. [LECT. HI. acetate of lime, sugar, mucilage, vegetable ex- tract, and water. In the sap of the Common Birch, Betula alba, he found acetate of lime, acetate of potash, acetate of alumina, sugar, vege- table extract, and water. In all the specimens thus analysed the quantity of vegetable matter was found to be greater in the sap drawn late in the season, than in that collected at an earlier period of it. If we are to consider these results of Vauque- lin's experiments as pointing out the real composi- tion of sap, we can gain no information from his labours ; as from the results of them we should suppose that every different kind of tree must have sap of a description peculiar to itself, which, both the analogy of the animal kingdom, and the knowledge we have of the nature of soils, inform us cannot be the case. It is probable that the water, the acetate of potash, and the carbonate of lime, are taken up from the soil, and enter as general constituents into the composition of sap ; but the mucilage, the sugar, the extractive, the gallic acid, the tannin, &c. are undoubtedly produced by the vegetable system itself, and must, therefore, vary in different plants. As we shall afterwards find that all the productions of the vegetable system can be resolved into carbon, hydrogen, and oxygen ; each production' differing in its composition from another in the proportion only of its components ; it is easy to comprehend that, as from the water LECT. III.] THE SAP. 113 the hydrogen and oxygen may be obtained, and the carbon from the acetate of potash and carbonate of lime ; Vauquelin was authorized in concluding that the acetate and carbonate, which are found in a diminished quantity in sap drawn at an advanced season, may be decomposed by the vital action of the growing plant, and the carbon they yield with the hydrogen and oxygen go to form the vegetable matter. But although the acetates are found in soils, yet they are not in any considerable quantity, nor sufficient to supply all the carbon wanted for the purposes of plants : it is, therefore, probable that if sap could be examined in its purest state, it would be found to consist of water, holding car- bonaceous matter in solution, acetate of potash, carbonate of lime, and now and then some siliceous and aluminous particles, suspended in the so- lution, and of sufficient minuteness to enter the mouth of the absorbents of the root. It is pro- bable, also, that there is little difference in the pure sap of all plants ; but as the first changes take place undoubtedly in the roots, and the modi- fying power of these parts must be different in different kinds of plants, the changes which occa- sion the varieties of the sap, are sooner produced in some plants than in others. On these differences of the ascending fluid, however, the secretions certainly do not depend ; for, if that were the case, the grafted branch would not bear fruit and leaves, VOL. I. I 114 THE SAP. [LECT. in. and have secretions deposited in its cells similar to those of the parent, but to those of the stock. We know, however, that this is not the case ; for, if the branch of a Pear-tree be engrafted on an Apple-tree, the Pear branch will produce leaves, flowers, and fruit, and have the new wood formed on it exactly the same as the Pear-tree from which it was cut, although the first modifications of the nutriment imbibed by the roots of the Apple stock are dif- ferent from those of the parent Pear : but as soon as it arrives at the secreting organs of the Pear branch the alterations it undergoes are exactly the same as if the branch had remained attached to its parent stem ; and consequently the wood, leaves, flowers and fruit, will have the characteristics of its original. The modifications which take place in the roots of plants, throw considerable obstacles in the way of obtaining a perfect knowledge of this part of the vegetable economy; for, to obtain such a knowledge of the nature of sap would require an examination of that fluid in a greater number of different species of plants, than the opportunities, and the period of any life, permit. All that we can aim at, therefore, in the present state of our knowledge, is the formation of a probable hypothesis, rather than the attainment of truth deduced from certain experiments. In this mode of viewing the subject, we may regard the sap LECT. III.] THE SAP. 115 of plants as consisting of water which is its prin- cipal component, carbonaceous mattery acetate of potash, and carbonate of lime*; which ingredients are decomposed by the vital powers of plants, and new combinations of their constituents produced by the same powers, so as to form the different parts of which a plant consists. The large portion of vegetable matter contained in the first sap, as we have already noticed, must have been previ- ously deposited in the cells of the root, and taken up by the water of the sap in its progress upwards : and air which is also found in sap, as Mr. Knight has demonstrated, is either the produce of vege- tation, or is taken in by the roots dissolved in the water of the soil-f~. Such is the nature of the sap. In spring and at midsummer it forms a large portion of the vegetable body ; and is carried forwards through the vessels, with an impetus sufficient to raise it to the summits of the highest trees, until arriving at the leaves, in * The enumeration of these ingredients as the general com- ponents of sap, cannot be objected to because many other saline and earthy matters are occasionally found in sap ; those depending altogether on local circumstances affecting the soil. f Sir H. Davy, in his Lectures on Agriculture, has adopted the opinion of Feburier, that the sap is found in two states ; one kind in the vessels of the alburnum, containing chiefly saccha- rine matter, mucus and albuminous matter, and another in the bark, containing tannin and extract ; but I am not inclined to regard any juice found in the bark as ascending sap ; and) there- fore, cannot subscribe to this opinion. i 2 116 GENERAL FLUID COMPONENTS. [LECT. III. which it is exposed to the action of the air, and light, the great quantity of water it contains be- comes no longer necessary, and is thrown off by perspiration ; whilst the succus proprius, or pecu- liar juice of the plant, from which all its secretions are formed, is produced by the changes resulting chiefly from this exposure. We have, therefore, next to proceed to examine the nature of this pe- culiar juice, as one of the general components of plants. GENERAL FLUID COMPONENTS THE PROPER JUICE. I should be anticipating the inquiries we have to make into the physiology of leaves, were I now to attempt to explain to you how the sap is con- veyed into these organs, to be exposed to the action of light and air ; or, by what means those changes are effected in them, by which it is converted into the proper juice, succus proprius : at present we have to examine this fluid merely as a general component. It is, however, necessary to remark, that although we may admit with Malpighi that the proper juice is to the vegetable system, what the blood is to the animal, yet, the functions by which it is prepared from the sap must be modified in different kinds of plants, since it exhibits some peculiar characteristics in each kind. When a plant is cut through transversely, the proper juice is seen issuing from both divided sur- LECT. III.] GENERAL FLUID COMPONENTS. 117 faces, but in greatest quantity from the open ori- fices of the divided vessels in the part farthest from the root ; a fact which is ascribable to the progres- sion of the proper juice being invertedly to that of the sap, or from the leaves towards the roots. It is very often mixed with sap, and cannot be distin- guished from it by colour ; but in many instances it is coloured or milky. Thus, if a twig of any of the species of Spurge (Euphorbia) be cut, the proper juice issues from the wound in the form of a resinous milky emulsion, and may be obtained in considerable quantity. This juice in the majority of plants is, as I have said, colourless ; it is, however, yellow in some, as in the Celandine (Chelidonium); red in others, as in the Bloody Dock (Rumex san- guinea) and the Logwood tree (Hsematoxylon) ; deep orange in the Artichoke (Cynara Scolymus) ; white, as in the Spurges (Euphorbia), the Dande- lion (Leontodon Taraxacum)., the Fig (Ficus), &c. ; blue in the root of Pimpernell (Pimpinella nigra) ; and green in the Periwinkle (Vinca). The colour is sometimes changed by the exposure of the exuded juice to the air. Thus Opium, which is the proper juice of the white Poppy, is white and milky when it exudes from the incisions made in the plant for the purpose of obtaining it; but changes to a yellowish brown hue by exposure to the air. The plantule also of the French or Ha- ricot Bean, when wounded, emits a reddish proper i 3 118 GENERAL FLUID COMPONENTS. [LECT. III. juice, which after being exposed to the air for a short time assumes a deep indigo blue tint : and the juice which exudes from incisions in the leaves of the Soccotrine Aloe, yields, by simple exposure, according to the statement of M. Fabroni, a very deep and lively purple dye, so permanent, and re- sisting so completely the action of acids, alkalies, and oxygen gas, that he thinks it may be used as a pigment in miniature painting ; or as a dye for silk, which it will effect without the use of any mordaunt*. It is necessary to mention that Mirbel, and some other Botanists, have fallen into an error, in confounding together the proper juice, and the secretions of plants. It is from the proper juice that the secretions are formed ; but it must un- dergo another elaboration, something similar to that which the blood of animals undergoes in their glands, before it is changed into the different se- cretions ; and assimilated into the substance of the plant. Thus, both essential and aromatic oils are found in some parts of the same plant ; muci- lage, resin, tannin, extract, acids and alkalies, and even silica, in other parts ; but these various pro- ductions cannot be considered as the proper juice : they are secretions formed from it. The proper juice of plants is, therefore, " that changed state of * Monthly Mg. 1798. LECT. III.] GENERAL FLUID COMPONENTS. 119 " the sap, after it has been exposed to the air, and " light, in the leaf, and is returning from it to " form the different secretions." The organs by which the secretion is performed are probably glands, which we have already endeavoured to prove exist in vegetables ; and the secreted fluids themselves are deposited in cells in different parts of the plant, particularly in the bark, and the roots ; these parts acquiring different medical vir- tues, from the matters thus lodged in them. It is almost as impossible to obtain the proper juice of plants free from sap, as it is to procure the sap free from the proper juice ; this, however, in the season in which it can be obtained in most abundance, is not so liable to be diluted or mixed with sap as at other times ; and therefore it is in the warmest times in summer, that it ought to be taken for the purpose of examining its properties- Some naturalists have, rather fancifully, drawn a very close analogy between it and the blood of animals. Thus Rafn, with a microscope magnifying 135 times, supposed that he could detect round globules, resembling the red globules of the blood, swimming in a clear fluid, in the juice of Euphorbia palustris; and Fontana thought he observed them in the sap of Rhus toxicodendron. But such obser- vations, which are often the effect of optical decep- tion, are of little value, even admitting their va- lidity, in a physiological point of view. In an i 4 120 GENERAL FLUID COMPONENTS. [LECT. III. accurate examination of the proper juice of plants, Mons. Chaptal found that in no two kinds of plants does it agree as far as its sensible qualities are con- sidered ; but as it is in the leaf that the change from sap into the proper juice occurs, so its sensible qualities are modified according to the action which takes place in that organ ; and that this should differ is not surprising if we consider the great difference of the structure of leaves. In one particular, however, Chaptal found that all the specimens he examined agreed. When he poured into them oxygenated muriatic acid, a very con- siderable white precipitate fell down ; which had the appearance of fine starch, when washed and dried, and did not change when kept for a length of time. It was insoluble in water, and was not af- fected by alkalies. Two thirds of it were dissolved in heated alcohol ; and these were evidently resinous, as they were again precipitated from the spiritous solvent by water. The third part, which continued insoluble in both alcohol and water, was found by Chaptal to possess all the properties of the ligneous fibre. In the seed lobes a greater quan- tity of this woody fibre was found than in the proper juice of the plant itself; a fact which ac- counts for the rapid growth and increase of parts of the young plant, before the roots are able to take up from the earth the principles of nutriment. The proper juices of plants, both in the seed, and LECT. III.] GENERAL FLUID COMPONENTS. 121 in the perfected plant, contain nourishment al- ready properly adapted to be immediately assi- milated into the substance of the plant. But this preparation takes place, either during the time, or after, the sap has been exposed to the action of the light and air in the leaf; as no woody fibre is found in the ascending sap, although the prin- ciples of it are undoubtedly contained in that fluid. A new chemical combination of these principles takes" place ; but how this is effected, or by what means the change is produced, we know not ; and it is one of those mysteries of nature from which human ingenuity will never perhaps be able to remove the veil. In the same manner the blood of animals contains the components of the mus- cular fibres already formed ; and an assimilation of it is constantly going on, without our being able to perceive it, or even to form the most distant conception of the manner in which it is performed. The elementary principles of the proper juice of plants and of the sap are the same ; but differ in the relative proportions. These principles are carbon, hydrogen, and oxygen. The same prin- ciples, differently modified, form all the secretions and the solid materials of the plant itself. The extraneous ingredients which some plants are found to contain, as part of their substance, such as the alkaline and neutral salts, metallic oxyds, silex and other earths, are probably obtained ready 122 GENERAL FLUID COMPONENTS. [LECT. III. formed in the soil, in a state of division sufficiently minute to be suspended in water, and drunk in by the absorbent vessels of the roots. This is in some degree proved by the effect of change of situation on plants which naturally grow near the sea. Most of these plants, when burned, yield soda ; but, when they are removed from the sea-shore, and cultivated in an inland situation, potash in- stead of soda is procured from their ashes. As the sap undergoes the same exposure to the air and light in all plants, and one product only can be formed in each plant by this exposure, the difference of the proper juice in different plants, is a strong argument in favour of the existence of vegetable glands, independent of the undeniable proof afforded by the formation of the very dif- ferent products, which are deposited in different parts of the same plant. Unless there were glan- dular organs, one product only could be produced in each plant by the function of the leaves, and the action of light and of air on the sap. The secre- tions of plants formed from the proper juice are very numerous, and known under the names of gum, fecula or starch, sugar, gluten, albumen, gelatin, caoutchouc, wax, Jixed oil, volatile oil, camphor, resin, gum resin, balsam, extract, tan- nin, acids, aroma, the bitter, the acrid and the narcotic principles, and ligneous Jibre. These are found in different parts of plants without any uni- LECT. III.] GENERAL FLUID COMPONENTS. 123 fonnity of distribution ; and although so nume- rous and different from each other in their sen- sible qualities and chemical properties., yet are they all composed of different modifications of the same principles, carbon, hydrogen, and oxygen. Thus 100 parts of gum, according to the experiments of Gay Lussac and Thenard *, consist of 42.23 of carbon, 6.93 of hydrogen, and 50.84 of oxygen, the oxygen and hydrogen being nearly in the same relative 100.00 proportions as they are contained in water. 100 parts of common resin consist of 75.944 of carbon, 15.156 of a combination of oxygen and hy- drogen in the same proportions as they exist in water, and 8.900 of hydrogen in excess. 100. 100 parts of olive oil consist of 77.213 of carbon, 10.712 oxygen and hydrogen, as in water, and 12.075 of hydrogen in excess. 100. * Recherches physico-chimigues, t. ii. p. 290. 124 GENERAL FLUID COMPONENTS. [LECT. III. The solids, also, except the earths and salts, are formed from the same principles : 100 parts of the ligneous fibre of the Beech and the Oak, for example, consisting of Beech. -Oak. Carbon 51.45 . . . . 52.53 Oxygen 42.73 .... 41.78 Hydrogen 5.82 .... 5.69 100.00 100.00 * and thus almost the whole of vegetable matter may be resolved into these three simple elements. Such are the general components of vegetables. The investigation of them is yet in its commence- ment only, and much must be done before their real properties be fully understood. It is requisite that the student keep them constantly in view, for otherwise much of the more detailed part of our subject will appear obscure and confused. * Recherches physico-chimiques, t. ii. p. 294. Thomsons Chemistry, fifth edition, vol. iv. p. 183. 125 LECTURE IV. VEGETABLE ORGANIZATION. THE ROOT ITS SITUA- TION, SPECIES, AND VARIETIES DIRECTION AND DURATION. HAVING endeavoured to give you some idea of the general components of the vegetable body, I have now to describe to you the organs formed by the combination of these constituents. Every plant, as I have already stated, possesses two sets of organs. One of these is intended merely for the growth and preservation of the in- dividual ; the other for the propagation, and, con- sequently, the continuation of the species : or' all plants are endowed with CONSERVATIVE and RE- PRODUCTIVE organs. If we dig up a tree or a shrub in the summer, when it is in flower, and some of the fruit is al- ready formed, an Orange-tree for example, which bears flowers and fruit at the same time, we have, in one plant, a complete display of these parts. In the first set are comprehended the root, the trunk, the branches, and the leaves, with their appendages ; in the second, the flower and the fruit, with their appendages. Many phytologists regard those species of plants only, which possess 126 CONSERVATIVE ORGANS. [LECT. IV. all, or the greater part of these organs, as perfect ; and those in which some of the more conspicuous of either kind are not present, or are apparently deficient, as imperfect. But we need not hesitate in pronouncing this opinion erroneous ; for, as the perfection of a plant consists in the power of its organs to carry on its functions, and to continue the species, individuals only can be imperfect. The different organs, in whatever manner they are present, assume a considerable variety of form ; and as it is on that variation that Botanists have founded specific distinctions ; and as it is of im- portance in a physiological point of view also, the student ought to make himself well acquainted with the distinguishing characteristics which it constitutes. In examining these, the best method is to take the organs in the order in which they have been enumerated: let us, therefore, com- mence with the CONSERVATIVE. a. The ROOT (Radix) is defined by Linnaeus to be that part of a plant which imbibes its nu- triment, producing the herbaceous part and the fructification ; and which consists of a caudex, or body, and radicles *. Simple as this definition ap- pears to be, it is, nevertheless, objectionable, inas- * " Radix alimentum hauriens, herbamque cum fructifica- " tione producens, componitur medulla, libro, cortice; constat- " que caudice et radicula" Phil. Botanica: LECT. IV.] THE ROOT. 127 much as many succulent plants in arid situations do not receive their nutriment by the root ; nor is the root the nutritious organ in the tribes named Hepaticee *, Confervse *j~, and Fuci ; in all of which it can be regarded as an attaching organ only, designed to secure the individual to the soil, or to the substance on which it is fixed. A less exceptionable definition is that adopted by Mr. Keith, which characterizes the root as " that part " of the plant by which it attaches itself to the soil " in which it grows, or to the substance on which " it feeds, and is the principal organ of nutrition ||." It might be stated, as an objection to this defini- tion, that plants exist which have roots, and, yet, are not fixed by them either to the soil or to any other substance ; as for example, Duckweed (Lemna), a small green lenticular plant, which floats abundantly on the surface of our stagnant pools in summer, and has unattached roots which hang perpendicularly loose in the water. But all plants are at first attached ; and as it is impossible to frame any definition to which no exception * A tribe of small herbaceous plants resembling the Mosses found in damp shacfed places. (- An aquatic genus, consisting of fibrous, or threadlike jointed and branched parts, closely matted together. Marine plants of various forms attached to rocks and stones. || System of physiological Botany, vol. i. p. 33. 128 CONSERVATIVE ORGANS. [LECT. IV. could be advanced, this " is perhaps/' as Mr. Keith remarks, " as comprehensive as any one " that can be given." In all plants the primary root is a simple elon- gation of that part which, during the germina- tion of the seed, is first protruded, and is deno- minated the radicle ; but as the plant continues to grow, the root gradually assumes a determinate form and structure, which differs materially in different plants, but is found always similar in all the individuals of the same species. Botanists have taken advantage of this fact, and have classed roots according to their forms, as seen in the adult, or fully grown root ; and have availed themselves of these diversities for fixing specific distinctions. The classification of roots, which I have ventured to arrange, differs from that which is usually found in works on the elements of Bo- tany ; but it is one which, I trust, will enable you to form accurate ideas regarding the structure and functions of these important organs. Before, how- ever, entering upon the consideration of it, you ought to be informed that every root, whatever may be its form, consists of two distinct parts, the body, caudex, and the rootlet, radicula. The main part of the root, caudex *, in trees and plants that * " Caudex descendens sub terra sensim subducit, et radicu- " las profert, a Botanicis ex varia structura variis nominibus " distinctus." Phil. Bot. 80. LECT. IV.] THE ROOT. 1'29 live for several yeais, is in general woody, and is to the other parts what the trunk or stem is to the branches and leaves ; enlarging progressively in a similar manner, and giving off lateral branching shoots, which spread horizontally, or in a direc- tion that forms nearly a right angle with the caudex. In those plants, however, the herbace- ous part of which dies annually whilst the root survives, and in many annuals, the caudex is also a reservoir of nutriment, which is intended for the renewal of the herbaceous part in the fol- lowing season, or to be expended in perfecting the flower and the seed. The rootlets, radiculce *, are small, or threadlike productions of the caudex, terminating in Jibrils, which are extremely mi- nute, the real absorbing organs of the root ; and are supposed by Du Hamel-f~, Willdenow^, Sir E. J. Smith and others, to die annually in the au- tumn with the foliage, and to be reproduced in the spring ; an opinion which Mr. Knight || re- gards as incorrect, as far at least as concerns the terminal fibres of woody plants. My own ex- * " Radicula est pars radicis fibrosa, in quam terminatur " caudex descendens, et qua radix nutrimentum haurit pro " vegetabilis sustentatione." Phil. 'Bot. 80. f Phys. des Arb. liv. i. chap. v. J Principles of Botany > 11. } Introduction to physical and systematical Botany, p. 104. Jl Phil. Trans. 1809. VOL. I. K 130 CONSERVATIVE ORGANS. [LECT. IV. perience and observations, however, lead me to believe that, even in woody plants, the fibrils are annual productions. Every root may be arranged under one of the three following classes, i. SIMPLE ROOTS ; ii. BRANCHED ROOTS ; iii. ARTICULATED ROOTS. i. The SIMPLE ROOT consists either of a single caudex furnished with fibrils only, or of one or more rootlets with fibrils*. Considered as a genus, it comprehends three species, the conical, the sufr- globular, and ihejibrous. 1. The Conical root (Radix conica) (fig. a) is a a. tapering caudex furnished with lateral fibrils, which are situated chiefly towards its smaller extremity. It is generally a reservoir of nutritious matter which is pre- pared in the leaves, and is the proper juice of the plant, to be exhausted in the pro- duction of the flower and the seed. The change which takes place at the flowering season in culinary roots of this descrip- tion, as for example, in the Carrot, is rendered very evident; by the nutritious matter it contains becoming less saccha- rine, diminishing in quantity, and the whole root acquiring a woody consistence. The wedgelike form of the conical root seems to * " Simplex qua? non subdividitur." Phil. Bot. 80. b. 3. LECT. IV.] THE ROOT. 131 be particularly well adapted for penetrating per- pendicularly into the ground ; but that this is not the sole intention of nature in giving it this form would appear from the fact, that tihe direction of some conical roots is horizontal ; as, for example, the Blood Root, Sanguinaria Canademis*, a North American plant, which is conical, truncated, and horizontal ; the root forming a right angle nearly with the stem. The roots of the Carrot, Daucus carota; the Parsnip, Pastinaca sativa ; the Horse- b. radish, Cochlearia Armor acia ; and the Dandelion, Leontodon Taraxacum ; are familiar examples of the conical root. The following may be regarded as its vari- eties. a. The Spindle-shaped root (Radix fusiformls^c) (fig. b) differs from the real conical root only, in not tapering equally throughout its length, but swelling out a little below its summit, like the spindle or wooden pin employed by ancient ma- trons in formmg the thread, which they drew from the flax wrapped round the distaff; whence its name. Like the pro- per conical root, it is a reservoir of nu- * .Vegetable Mat. Med. of the United States, p. 31. f " Fusiformis quae oblonga, crassa, attenuata : ut Daucus, " Pastinaca." Phil. Bot. 80. This is the definition rather of the conical root than of the spindle-shaped. K 2 132 CONSERVATIVE ORGANS. tritious secreted mattei [LECT. iv. and by its shape, it is well calculated to penetrate the ground. The Ra- dish, Raphanus sativus ; and the Beet, Beta vul- garis ; are the best examples of this form of root. b. The Abrupt or truncated root (Radix c. prcemorsa *) (fig. c ) is ori- ginally an entire conical root ; but after some time the lower extremity decays and drops, as if it had been bitten off, while numerous lateral rootlets are protruded from the remain- ing portion. Such is the case in the root of the larger Plan- tain, Plantago major, and in that of Scabiosa succisa, which, according to Gerarde-f*, received its common appellation Devil's Bite, Morsus Diaboli (fig. c), from a su- perstitious opinion connected with this appear- ance of the root. A curious modification of the Abrupt root * " Pramorsa, quae deorsum truncata est, nee attenuate " apice terminatur ; ut Scabiosa, Plantago, Valeriana" Phil. Bot. 80. 9. f The old herbalist's words are, " The greater part of the " root seemeth to be bitten away : old fantasticke charmers re- " port, that the devil did bite it for envie, because it is an herbe " that hath so many good vertues, and is so beneficial to man- kinde." Vide Herbal, p. 726. LECT. IV.] THE ROOT. which occurs in a few herbaceous plants, and which has misled some Botanists to describe it erroneously in these instances, as an articulated or scaly jointed root, depends on the following circumstance, which was first noticed and de- scribed by Dr. Grew*. In the Primrose, Pri- mula veris, for example, the root is abrupt ; but as the lower leaves of the plant annually decay and fall, they leave a small portion of their basis d. at the place of their attachment, which swells and becomes more succulent ; and the plant sinking in the ground, lateral fi- bres are protruded above each of these portions ; so that the buried part of the root, owing to a similar de- cay and sinking annu- ally taking place, gra- dually assumes the character of a long caudex, and the whole bears a strong resem- blance to a notched or articulated root. See (fig. d.) a, the present year's * Anatomy of Plants. K3 134 CONSERVATIVE ORGANS. [LECT. IV. foliage ; b. the remains of last year's, with "a rootlet protruded above it ; c. c. bases of old leaves converted into firm toothlike scales or processes ; d. the decayed, or truncated root. The cause of this decay of the lower extremity of some conical roots shall be explained, when treating of the physiology of these organs. 2. The Subglobular root (Radix subrotunda) is an almost spherical caudex, terminating in one or more small tapering points. Like the conical root, it is a reservoir of nutritious matterlntended for the production of the flower and seed. The best example of it is the black Radish, Ra- phanus sativus, var. B. niger. There are two va- rieties of the subglobular root : a. The Turnip-shaped root (Radix napiformis) (fig. e) is a caudex, the shape of which is the intermediate of the spindle- shaped and the subglobular roots, bellying out suddenly above, and terminating belowin along taper- ing point furnished with fibrils. It is scarcely necessary to quote the Turnip, Brassica Rapa, as an example. I. The Flattened subglobular root (Radix placenttf'ormis) has the appearance of a globular caudex which is compressed both above and be- LECT. IV.] THE ROOT. 135 low. It has not the tapering point of the two former, but a number of long fibrils which hang from the centre of the lower depression. The Sow-bread, Cyclamen europceum (fig. y), which is an example of this root, does not attain its de- cisive character in the first year of its growth ; and al- though a reservoir of nu- triment, yet it is not ex- hausted in perfecting the flower and fruit, but con- tinues to increase for se- veral years. The nutriment which is deposited in the cau- dices of all these species of the simple root, is not that which is directly absorbed from the soil by the fibrils ; but the proper juice of the plant, pre- pared from the sap exposed to the action of the light and' air in the leaves. The necessity of a luxuriant and healthy state of the herbaceous part of the plant, therefore, for increasing the quantity of nutritious matter in the simple roots, which are cultivated for food, is very obvious. It may be thought that the practice pursued to produce earlier and larger Radishes, " which is by sowing them in " hot-beds in the early spring, and exposing the the shrivelled one is that of the preceding year, which having its nu- tritious contents exhausted in formation of the herb and the flowers, shrivels towards autumn ; and either withers away altogether and disappears in the succeeding winter, or remains a mere ske- leton in the third year. The ovate conjoined tuber is therefore biennial, being formed and perfected in one year, and performing its functions and dy- ing in the second. In fig. q are seen these three states of tuber as displayed in Orchis acuminatum: L3 150 CONSERVATIVE ORGANS. [LECT. IV. 1. shows the tuber nearly shrivelled to a skin; 2. the tuber of last year after the plant it bears has flowered ; 3. the new tuber, the production of the present year ; 4. are the real roots of the plant*. r. Our conjoined ovate tu- ber is the Radix testiculata of 'authors-}-. , 2. The Conjoined club- shaped tuber (Tuber clavce- 1 forme conjunctum) (fig. r ) is of an oblong shape, thicker at the loose extremity, and resembling in some degree a short club. As in the former species of tuber, it is generated every * There formerly existed a very absurd superstition con- nected with the double state of these tubers. If a pair of them be separated and thrown into water, the new tuber, owing to its greater gravity, sinks, whilst the older one, being lighter, swims. The swimming tuber, when prepared in a particular manner and worn round the neck, was believed to possess the magic pro- perty of securing to the wearer the strongest attachment of any one he pleased ; and this belief still continues to prevail to some extent among the ignorant. A more rational and useful purpose to which the new tuber of the Orchis mono, and some other spe- cies of the Orchidece is applied, is the manufacture of salop. For this purpose the new tuber, which has attained its perfect growth, is prepared by first scalding it in boiling, water to detract the skin, then placing it in an oven for ten or twelve minutes to give it semitransparency, and finally drying it in a moderate heat. In this state it resembles sago, and constitutes a nutritious wholesome article of diet. f Quasi testiculis animalium similis. LECT. IV.] THB ROOT. TUBERS. 151 year; but there are always two or more tubers in a state of advancement towards perfection, and as many in that of decay. Pile-wort, Ranunculus Jicaria, affords the best example of this form of tuber : vide fig. r, in which 1. 1. indicates the more recent tubers ; 2. 2. those in a state of decay. . 3. The Fingered tuber (Tuber digitatum) (fig.*) receives its name from the tubers resembling fingers. In Satyrium albidum, the plant exhausts two tubers in one season; see fig. s, in which b. b. represents a pair of old tubers, the origin of the existing plant ; a. a pair of tubers for that of the next year, with a still younger pair attached to them : c. c. are the real roots of the plant. 4. The Palmated tuber (Tuber palmatum) (fig. t) also receives its name from its form, which somewhat re- sembles the human hand. There are rarely more than two palmated tubers con- joined, as in Orchis 152 CONSERVATIVE ORGANS. [t,ECT. IV. maculata. In fig. t, 1.1. represents the roots; 2. the withering tuber, the origin of the present plant; 3. the new tuber, the pyramidal eleva- tion on the top of which marks the point, whence shall germinate the plant of the ensuing year. 5. The Bundled tuber (Tuber fasciculatum) u. (fig. u) is spindle- shaped or nearly cylindrical. Tu- bers of this de- scription " are " "formed,andalso " wither away in " parcels," each parcel being equi- valent to a single tuber in the species already de- scribed : they are well exemplified in Bird's-nest Ophrys, Ophrys nidus avis (fig. u). Such are the closely attached tubers. From the circumstance of the new tuber, or the sets of tubers, being always formed laterally to the old, the plant appears to perform a certain degree of progression ; so that in a few years it is found at some distance from the spot where it originally flourished*. I have used the term, " the plant," * The cultivation of tuberiferous plants with closely at- tached tubers, which are chiefly of the Orchis tribe, has always been considered very difficult ; but it may be successfully ma- naged as practised by the late Mr. Crowe, and thus described LECT. IV.] THE ROOT. TUBERS. 153 because, although a new plant actually rises every year from the new tuber, or that one in the regu r lar order of succession ; yet each is only a conti- nuation of the plant raised from the seed perpe- tuated by lateral production, as the buds on an ungrafted tree are continuations of the original tree ; and differ essentially from the seedling plant, which is a real renewal of the species. b. FJLIPENDULOUS TUBERS are attached to the parent plant by underground runners, or cords, which spring not from the roots but the lower part of "the stem ; the roots being in this instance, as in the plants with closely attached tubers, truly v. fibrous. There are two species only of the fill- pen dulo us tubers ; soli- tary and congregated. I- The 5otorpen- dulous tuber (Tuber so- lit arium) (fig. v ) is egg- shaped and attached, as by Sir J. E. Smith : " They are best removed when in full flower, " the earth being cleared completely away from the roots " (tubers), which are then to be replanted in their natural " soil previously dried and sifted. Afterwards they must be " well watered. The bulb (tuber) for the following year has " not at the flowering period begun to throw out its fibres, for " after that happens it will not bear removal." Introd. to phy- siological and systematic Botany t p. 110. 154 CONSERVATIVE ORGANS. [LECT. IV. at b. to the extremity of a lateral runner, proceed- ing from the collar of the stem, immediately above a. the old, or plant-bearing tuber. As the new tu- ber makes very little progress until after the flow- ering of the parent plant is over, it may escape the observation of the student, if looked for be- fore that period. The Musk Orchis, Ophrys monorchis (fig. v), which has received its spe- cific name Monorchis*, from having apparently but one tuber, is the only example of this species of tuber with which I am acquainted. 2. Congregated pendulous tubers (Tubera con- gregata) are more generally either globular or ovate, surrounding the stem, attached by runners or fibres which connect them with it at different distances, and in indefinite numbers. Like all other tubers, they are reservoirs of nutriment and moisture ; whilst the runners to which they are attached perform an office similar to that of the umbilical cord in animals ; conveying the pro- per juice from the parent plant to the tubers, to be deposited in them for the support of the suc- ceeding plants; and maintaining the connexion between the plant and its lateral progeny, the gems on the surface of the tubers, until these are perfected and endued with that vitality which en- ables them to exist as independent beings. It is a * From /uo'vos (monos), one. LECT. IV.] THE HOOT. TUBERS. 155 conjecture which is not improbable, that the final cause of the length of the runners is to place the tubers at a convenient distance from the parent plant and from each other, in order that the new plants may enjoy the advantages of a fresh and unexhausted soil. The Potatoe, which affords the best exemplifica- tion of this species of tuber, has many gems on its surface, all of which shoot up into stems ; and from each, besides the runners for the production of new tubers, rootlets are given off for the ab- sorption of that portion of nutriment which is required to be supplied from the soil for the sup- port of^the vegetating plants. Such are all the species of tubers which I think it necessary to particularize ; and to one or other of them, every known tuber may be re- ferred. Other species are undoubtedly noticed by authors, under the head of tuberous roots; but these will be found to be varieties only of some of those we have examined : even the granules attached to the roots of the White Saxifrage, Saxifraga granulata, are sometimes quoted* as tubers, although they actually belong to the next of the appendages of the root, which we have to take under consideration, the bulbs. The roots of several species of the Iris tribe have some resem- * Keith's Syst. of physiological Botany > vol. i. p. 39. 4 156 CONSERVATIVE ORGANS. [LECT. IV. blance to the tuber ; but although the new caudexes of these roots bear buds, which may be correctly regarded as lateral progeny, yet, these caudexes are nevertheless real roots, and do not accord with the definition of tubers. I shall have to notice them more particularly when we examine the di- rection of roots. As in tuberiferous plants, the tuber is formed by the plant of the present year, and itself forms that of the succeeding, which again forms a new tuber or tubers, it becomes a subject of rational inquiry to ascertain the part which the real roots of the plant act, in producing these results. That the plant is sustained to a cer- tain degree by the absorbing powers of the real roots, is obvious ; but it has not, yet, been deter- mined to what extent or at what period it might be deprived of the tuber and be supported solely by the roots. In the early stage of its growth it is altogether nourished by the tuber, the nutriment passing directly from the tuber into the vessels of the stem, and ascending through them to nou- rish the plant. When the roots begin to absorb from the soil, the fluid matters they take in are probably mingled, in the leaf, with the already formed nutriment brought from the tuber ; and it is not unlikely that it is from the proper juice pro- duced by the exposure of this mixture to the air and light in the leaf, that the new tubers are formed. That the new tubers, however, may be LECT. IV.] THE ROOT. TUBERS. 157 formed independent altogether of the nutriment obtained from the tuber bearing the plant, is evi- dent ; for, the first tubers of the plant raised from seed are formed altogether from the nutriment obtained from the soil. It should, however, be re- collected, that the plant raised from seed differs from that evolved from the gem on a tuber ; the former being an entire new being, a renewal of the species, the latter a mere continuation of the in- dividual from which it springs. It may be sup- posed, that although the new tubers be formed from the nutriment obtained from the soil, yet that the contents of the old tuber are intended chiefly for perfecting the flower and the seed *, in the same manner as the saccharine matter depo- sited in the caudexes of the Turnip and Carrot ; but, admitting this supposition, it is not the less true that this nutriment may be diverted to the use of the new tubers ; for those on the Potatoe plant, are both enlarged and multiplied by nipping off the flowers, to prevent the formation of the seed. From whatever source the nutriment is obtained, the healthy state of the leaf is absolutely requisite to perfect the tuber ; for, the partial destruction of * This supposition certainly obtains some support from the fact, that Potatoes produced on plants raised from seed do not flower in the second or third year ; the tubers apparently re- quiring a greater magnitude and higher degree of perfection before they can form the flower. 158 CONSERVATIVE ORGANS. [LECT. IV. these by insects or frost, or the impeding of their functions by disease, as for example the curling of the leaves of the Potatoe plant, is followed by a decrease both in the quantity and in the size of the new tubers, and also by a deterioration of the nutriment deposited in them. The runners to which pendulous tubers are attached are bundles of vessels, produced from the bark of the stem, and intended solely to convey the proper juice from the descending vessels to these knobs ; and that there are no returning vessels, or, in other words, that the progression of fluids through these runners is in a direction from the stem to the tubers only, is rendered pro- bable from the following experiment made by Mr. Knight. He divided the runners connecting the tubers with the stem in a Potatoe plant, and immersed both portions in a decoction of Logwood. The colouring matter passed along in both direc- tions, but did not enter the stem, which it would have done had there been any returning vessels in the runners, whilst it was found to have entered the tubers and filled an elaborate assemblage of vessels which are situated between the bark and fleshy parenchyma *. The anatomy of tubers displays a considerable difference in the structure of those which belong * Phil. Transactions, 1803. LECT. IV.] THE ROOT. TUBERS. 159 to the tribe of plants named Monocotyledons and that .denominated Dicotyledons; it demonstrates also, that the organization of the tuber has the closest affinity to that of the stems of the tribe to which it belongs. Thus, for example, the stem of the Orchis mascula, which is an annual, tuberiferous, monocotyledonous plant, consists of a lax cellular texture enclosed in a cutis vera and epidermis, with threadlike bundles of vessels running longitudi- nally through it, at equal distances. The same appearance is evident in the tuber, the vascular bundles or cords being larger only, and inosculating with each other at the lower extremity of the tu- ber, whilst at the upper they pass into a transverse bundle, which seems to unite with one cord of vessels descending from the stem of the plant, and two cords which apparently connect the vascular system of the new tuber with that of the parent stem and the old tuber. This organization is quite obvious to the naked eye in the half-decayed tuber, and can be made so in the new tuber if the stem of the plant be cut transversely through about an inch from the tubers, and the portion to which they are attached inverted, and allowed to remain for twenty-four hours in a decoction of Brazil wood. When the vessels are thus filled with colouring matter, a longitudinal section of the new tuber dis- plays the vessels running in cords through the cel- lular texture, which is now swelled with nutritious fluids (vide plate 1. fig. 1. a), whilst a transverse 160 CONSERVATIVE ORGANS. [LECT. IV. section shows them like coloured points, set in a diamond form, in the white parenchyma (Ib. fig. 2). In the old tuber this appearance varies ; the lon- gitudinal section showing the cords of vessels sur- rounded with the cells still turgid with fluids to a certain distance on every side, and these enclosed as it were with a wall of empty cells, which in a transverse section assumes a circular or irregular hexagonal figure ; or each cord of vessels appears as a point in the centre of a large circle or hexagon, the interior of which consists of moist cellular matter, and the walls of dry or empty cells (Ib. fig. 3). This appearance of the old tuber can be explained only on the supposition that during the exhaustion of the nutritious matter it contains, the fluids pass laterally through the cells towards each cord of vessels, by which they are taken up and conveyed to the vessels of the stem and those of the new tuber ; and consequently, the more distant cells being those soonest emptied, and each cord of vessels acting as the attractive centre of a given space, the cells on the periphery of each space, owing to the positions of the vas- cular cords, must necessarily be emptied in such a manner as to give the circular or hexagonal ap- pearance to the transverse section. If this opinion be well founded, it would appear, that the vessels of the tubers of monocotyledonous plants, which during the growth of the tuber convey to and LECT. IV.] THE ROOT. TUBERS. J61 deposite the nutritious fluids in the cells of the parenchyma, take on a retrograde movement, and acquire the functions of absorbents as soon as the tuber begins to vegetate. A knowledge of the dis- tribution of the vascular system of these tubers, and of the manner in which the new tuber is at- tached, with the appearance of the embryon plant at its apex, can be readily obtained by examining a longitudinal section of the base of any Orchis, after the new tuber is formed ; particularly if the plant be injected with a coloured fluid, as has been already described *. From such an examination it is evident, that, however close the attachment of the new and old tubers is in the closely attached tubers, the attaching organ performs a function very similar to that of the umbilical cord of ani- mals ; and resembles in every respect, except in length, the runners which connect the pendulous * Vide Plate I. fig. 1. which represents a longitudinal slice of the tuber of the male Orchis, Orchis mascula ; a. the new tuber with its longitudinal vessels, and the intermediate cel- lular texture turgid with nutritious fluid ; b. the old tuber in a half-exhausted state; c. the stem of the plant broken short; d. t\\e plantule, which will form the next year's plant ; e. a kind of sheath over the plantule> which terminates in a membranous cone, and appears to be formed of a doubling of the cuticle of the stem, reflected so as to unite with that of the new- tuber ;f, the point of conjunction of the vessels of the stem and of the old tuber, with those of the plantule and the new tuber ; g. a root produced from the stem* VOL. I. M 162 CONSERVATIVE ORGANS. [LECT. IV. tubers with the parent plant. In fact, the attach- ment, when the tuber is viewed in its natural state, appears much shorter than it actually is, owing to the shortness of the collar of the stem: but circumstances occasionally occur which lengthen the organ, so as to give it the real character of a runner, and render the new tuber pendulous (fig. w) *. If we take the Potatoe to exemplify the organization of the tubers of Dicotyledons, we shall find that they resemble, in an equal degree, the stems of the tribe. When a tuber of this plant is cut, either transversely or longitudinally, it appears evidently to be composed of two distinct parts ; one of which is internal and cellular, somewhat like the pith or central part of the stern of the plant, and surrounded on every side by the other, which is cellular also but more dense, and bor- dered with a complicated system of vessels, branches from which stretch into the pith ; and that it has a close affinity to the cortical part of the stem. * The plant from which this figure was drawn was raised in a common garden pot, with the old tuber very near the surface. a. The old tuber ; b. the new tuber ; e. the attaching process extended so as to assume the form of a runner. LECT. IV.] THE ROOT. TUBERS. 163 Wherever a gem is formed on the tuber, the pith or central part advances to the surface^ and gives a small thread of its substance to the em- bryon ; and this appears to furnish its pith : whilst, in the same manner, the cortical substance, which, although attenuated almost to a pellicle^ yet still surrounds the thread of pith, seems to supply its bark*. The vessels also of the tuber tend to each point where a gem rises on its surface ; and, when this begins to vegetate, they enlarge so much as to become very perceptible to the naked eye ; and acquire a retrograde movement, taking on the functions of absorbents, and conveying the nutri- tious matter deposited in the tuber to the vascular system of the germe for its evolution and support. The epidermis of tubers, to whichsoever na- tural order of plants they belong, is utterly devoid of either absorbing or exhaling organs ; and it is this circumstance which fits tubers so admirably for preserving, in an unaltered state, the nutri- ment deposited in them ; the preservation of this in its natural state being absolutely necessary for * Vide Plate 1, fig. 4-. A longitudinal slice of a Potatoe : a. the central part, or pith ; b. the cortical part ; c. the change in the position of these parts where a gem is given off; d. the point where the tuber was attached to the runner. It may not, perhaps, be unnecessary to say that the points on the sur- face of a Potatoe, called eyes in common language, are the gems, the rudiments of the expected plants. Mi 164 CONSERVATIVE ORGANS. [LECT. IV. enabling the germe to maintain its vitality. Finally we may, with Sprengel, regard the tuber as a hybernaculum, or winter habitation, for the pre- servation of the embryon plants yet latent in the gems which it bears *. The BULB, the next of the radical appendages which we have to examine, is a globular or pyri- form, coated body, solid or formed of fleshy scales or layers, and of a more or less complex structure. It is not peculiarly an appendage of the root, but is found also on the stem and branches of some plants, and even mingled with the flowers of others. As, however, the bulbs which are attached to the root, have several peculiarities which dis- tinguish them from those situated on the other parts of the plant, it is necessary to treat of them in this stage of our inquiries. The roots of bulbs are simple fleshy fibres, annual productions, which issue either from the circumference of the flattened basis of a kind of caudex on which the scaly and laminated bulbs are seated, or from the substance of the bulb itself, * Linnaeus defines the hybernaculum thus : " Hybernaculum " est pars plantae includens herbam embryonem ab externis " injuriis ;" and adds, " estque Bulbus vel Gemma." Sprengel, in the edition of the Philosophia Botanica, which he has edited, very properly adds the tuber. " Tuber est hybernaculum solidum, " substantia marginali molliori cinctum'* LECT. IV.] THB ROOT. BULBS. 165 or from a simple scale, when it is solid ; and these situations of the roots form the chief characteristic of the bulb, distinguishing it from the tuber, and marking out those limits between the bulb and the tuber, ,the existence of which is denied by Mirbel * and some other authors : for the roots of the plant borne on a tuber, whether of the closely attached or pendulous kind, always proceed from the stem. Bulbs, like tubers, are reservoirs of nutriment for the developement and temporary support of the plants formed within them ; a fact which is illustrated to common observation by the growing of onions, in the storeroom of the housewife. Young radical bulbs, as I shall soon have occasion to show you, are produced in some instances above, in others below, and in others again within, or at the sides of an old bulb, during the adult vegeta- tion of its plant ; and are, like the gems formed on the surface of a tuber, the lateral progeny of the plant itself, not of the bulb. They are more frequently formed previously to the flowering of the parent plant; and as they preserve the plantule within them, until the ensuing spring, they are re- garded as winter habitations (hybernacula) by Lin- * " II n'y a point de limites entrela bulbe et le tubercule; la " transition se fait de 1'une a 1'autre, par la bulbe solide qui " participe de toutes deux." Elem. de Phys. veg. l ere partie, p. 135, nota. M 3 166 CONSERVATIVE ORGANS. [LECT. IV. neeus, who points out the close affinity, except in situation*, which exists between them and the buds of trees. Bulbs may be divided into solid, scaly, and laminated. I. The SOLID BULB ^ is a mass of cellular substance, filled with nutritious fluids, and en- closed within a thin epidermis, with vessels run- ning through it from the basis to the apex. It is covered with one or more coats, either of a mem- branous, or a fibrous, or a reticulated texture J. The solid bulb resembles the tuber in several re- spects ; but differs from it in being coated, and, as has been already noticed, in giving off the roots from a radical plate, or from a scale at the basis of the bulb, the point opposite to that from which the shoot is produced. Solid bulbs may be arranged under three spe- cies, taken from the situation on the old bulb where the new one is produced ; and thence they may be denominated superincumbent, lateral, and enclosing. 1. The Superincumbent solid bulb (Bulbus so- lidus superpositus) I have so named from the new bulbs being produced on the summit of the old one, and appearing, even when they are fully * " Bulbus est hybernaculum caudiei descendenti insidens." Phil. Bot. 85. f " Solidus constans substantia solida.'* Ib. J Vide Plate L fig. 7. IrECT. IV.] THE ROOT. BULBS. 167 grown,, incumbent on it ; as exemplified in Crocus*, Ixia^y &c. The old bulb withers, whilst those which it bears are attaining maturity; but it does not entirely decay away, so as to leave its progeny as distinct and separate individuals, until the new bulbs be formed on them. In some in- stances, Crocus for example, the plants of the present year, besides exhausting the old bulb for their support, appear to obtain that portion also of their nourishment, which is drawn from the soil, through it ; the vessels that proceed from its ra- dical plate passing through the centre of the present year's bulbs, and sending canals even to the rudiments of those formed at the basis of their foliage. This is rendered evident to the naked eye by dividing bulbs of Crocus sativus, taken up in the latter end of May ^. In other plants, as * Vide Plate 1 . fig. 5. Bulbs of Crocus sativus dug up in May. a. b. the perfected bulbs, adhering closely to c. the old bulb much shrivelled ; d. the radical plate of the old bulb with roots protruded from its circumference. f Fig 7. Bulbs of Ixia polystachia taken up, also, in May. a. b. the new bulbs which bear the present year's foliage and flower, partly covered with c. the reticulated coat of the old bulb d. ; e. e. e. e. e. the roots of the recent bulbs, protruded between them and the old bulb;,/, a small bulb appended by a runner as described in the text. J Fig. 6. The bulbs of fig. 5 dissected : a. b. the re- cent bulbs ; c. the old bulb, with its vessels seen passing up from its radical plate, and continued through the new bulb to the basis of its foliage ; d. two cords of vessels which M4 168 CONSERVATIVE ORGANS. [LECT. IV. Ixia for instance, although the same course of vessels be perceptible, yet, very thick roots are thrown out from the superincumbent recent bulbs, by which the nutriment is immediately taken up from the soil, without passing through the de- caying bulb *. Under certain circumstances of soil, also, wires are protruded from between the more recent bulbs and the old bulb, with small bulbs attached to them-f-, and these are some- times so elongated as, to give to the lateral pro- geny a pendulous character. On dissecting this species of bulb, we find that, besides the outermost coat, which is in general either fibrous or reticulated, it has two others more succulent and fleshy, which appear to form the bases of the sheathing of the leaves, and to cover the whole of the bulb terminating at the plate whence the roots are protruded. Within these is the solid homogeneous mass of the bulb, covered with a thin, beautiful, transparent epider- mis, and depressed considerably in the centre, where the stem and part of the foliage are at- tached ; and having cavities also at those points where the young bulbs are formed. The exterior leaves terminate at a kind of shoulder, which sur- separate from the others immediately after they enter the new bulb, and pass on to the embryon bulbs, forming on a. 6. e* the embryon bulbs. * Vide Plate I. fig. 7. f Ibfd. fig. 7. LECT. IV.] THE ROOT. BULBS. 169 rounds the general cavity ; and within the ex- panded bases of these leaves the young bulbs are seated. As in the case of tubers, although the young bulbs are the production of the existing plant, yet, as has been already remarked, they have a vascular communication with the soil through the medium both of their direct parent and the old decaying bulb*. From this examina- tion the following physiological conclusions may be drawn ; that, in this species of the solid bulb, the old bulb is exhausted in the production of the herbaceous part of the plant by the functions of which the bulbs immediately incumbent upon it, and which bear that plant, are perfected, and by which the embryon bulbs, seated on each of them, are also formed. Towards the autumn, the exhaustion of the old bulb is completed ; and the vinculum which connects together the bulbs seated on it, and which are now perfected, being thus broken, they separate ; and each remains as a distinct being, to be in its turn exhausted, in the production of the plant of the ensuing year, and of its series of bulbs. 2. The Lateral solid bulb (Bulbus solidus la- teralis) is named from the new bulbs being formed on each side of the old, as exemplified in Colchi- cum autumnale. In this species of solid bulb, the * Vide Plate 1. fig. 6. d. 170 CONSERVATIVE ORGANS. [LECT. IV. old bulb is completely exhausted, and the new bulbs separated from it before the rudiments of the next series are perceptible. The points of union between the parent and its progeny are near the basis of the bulb on both sides. On dissection, the perfect bulb is found to consist of a mass of cellular substance turgid with moist farinaceous matter ; but it is not, as in the former species, per- fectly homogeneous, one part being opaque, and the other semitransparent. The vessels run lon- gitudinally through the bulb, from the radical plate to the part whence the foliage springs ; and, in their dried up state, in conjunction with the emptied cellular texture and the epidermis, give a tough, spongy character to the exhausted bulb ; which, at the time of the separation of the recent bulbs, exhibits the appearance of a shrivelled leathery bag, the roots having dropped from the radical plate *. * Vide Plate 1. fig. 8, 9, 10, which represent the bulbs of Colchicum autumnale, dug up in May, in different points of view. Fig. 8. a. b. the recent bulbs (denuded so as to show their true appearance) of an irregular Pear shape, being con- siderably longer on the side opposite to that by which they are attached to the old bulb, and terminating in c. c. a flattened process ; on the inner and upper part of which, under a pro- jection formed by the termination of the shorter side of the bulb, is the radical plate, whence the roots protrude. It is half way above this plate, on the one side, and behind the flattened process, on the other, that the new bulbs are formed. d. The LECT. IV.] THE ROOT. BULBS. 171 The physiology of this species of solid bulb closely resembles that of the other. The new bulbs are produced by the plant of the bulb to which they are attached, and which is exhausted in the support of the foliage and fructification they bear : and in partly supplying also the nutriment which is deposited in them : I say partly, because it must be kept in mind, that the recent bulbs, of both species, have a direct communication with the soil through their roots ; and consequently it is probable that by far the greater part of the matter deposited in them, proceeds from the nutriment obtained from the soil, changed into the proper juice of the plant by the functions of the leaves. 3. The Enclosing solid bulb (Bulbus solidus includens), I have named from the circumstance of the young bulbs being enclosed between their d. The old bulb shrivelled up ; e. the remains of its radical plate ;f. the remains of its terminal process. Fig. 9. the back view of fig. 8. a. b. the recent bulbs ; c. c. the flattened processes seen from behind ; d. the old bulb ; e. the remains of its radical plate. Fig. 10. a longitudinal slice of the bulbs, a. The old shri- velled bulb ; b. c. the recent bulbs, in which is seen the dis- similar structure of their mass ; d. the opaque portion ; e. the semitransparent : the longitudinal lines in d. show the course of the vessels, proceeding from the point of attachment behind the process of the old bulb ; at^ and on the opposite side, from that above its radical plate at g. h. h. the part of the bulbs on which the foliage is seated. 172 CONSERVATIVE ORGANS. fLBCT. IV. \ - common parents; which, in this species of bulb, are two hemispheres, opposed to each other by their flat surfaces, and united at their bases to the radical plate, whence the stem proceeds *. Its physiology is the same as that of the other solid bulbs. II. SCALY BULBS consist of fleshy scales at- tached to a radical plate, and so arranged as to lie over each other like the tiles of a roof. v Each scale is a homogeneous mass of cellular substance, thick and fleshy in the middle ; but in some in- stances nearly membranous at the edge, concave on one side, and necessarily convex on the other. Each scale, also, is a distinct reservoir of nutri- ment, and is endued with such a share of vitality as not only enables it to live if detached from the bulb ; but, when placed in a proper soil, to ve- getate and produce an entire new bulb. There are two species only of scaly bulbs, the squamous and the granulated. 1 . The Squamous bulb (Bulbus squamosus ^) * Vide Plate 3. fig. 12. The twofold bulb of cluster-flowered Fritillary (F. Pyrenaica). A. a. a. the two hemispheres united at the caudex ; b. the stem rising between them ; c. the roots : B. a. one hemisphere separated from the other, in order to show the two young bulbs b. b. produced on each side of the stem c. ; the longitudinal line on each marking the place at which the di- vision will take place: d. the caudex. f " Squamosus constans imbricatis lamellis. Lilium.*' Phil. Botan. 85. 1. 4 LECT. IV.] THE ROOT. BULBS. 173 is well exemplified in the LiJy tribe. If we dig up a plant of one of these, say the White Lily, Lilium candidum, in summer, and examine its bulb, we shall find that it consists of concave, fleshy, overlapping scales, each of which is at- tached at the base to a radical caudex, but is loose at the apex*. A scale, when separately examined, appears to be a homogeneous mass of cellular texture enclosed in a cuticle ; the middle being thick and fleshy, but the edges and the apex nearly membranous; a construction, which, on the convex surface, gives it a gibbous or keeled character. The vessels run in longitudinal cords, arranged at equal distances, through the scale ; as may be seen in a transverse section of it, if the bulb Have been placed for some hours in a coloured so- lution-^. Removing the scales till the stem be ex- posed, the new bulb is seen formed at its basis ; and, by making a longitudinal section of the whole, the manner in which the lateral offspring is connected with the parent plant, through the medium of the vascular system of the stem and of the caudex, is * Plate 2. fig. 1. The entire bulb of the Lilium candidum, as it appears, when taken up in summer, during the vegetation of the stem. f Fig. 2. A transverse slice of one of the scales, to show the arrangement of the vessels in its substance. The vascular bundles appear like dots ; and owing to the cellular texture being more condensed in the immediate vicinity of these, each bundle seems as if it were enclosed in a sheath. 174 CONSERVATIVE ORGANS. [LECT. IV. made apparent*. When the new bulb is per- fected after the flowering 1 season is over, the ex- terior scales of the old one decay ; and by degrees the stem and a large portion of the old caudex also separate ; but as the whole of the caudex does not slough off, and a new portion is added by each successive bulb, it becomes gradually elon- gated, and at length bears some resemblance, when the scales are taken off, to a cylindrical, toothed, praemorse root. The new bulb, how- ever, is not always found seated close to the stem, in every species of this natural genus of bulbife- rous plants. In Lilium superbum, for instance, a very thick, succulent, lateral runner is projected from the caudex of the bulb ; and, pushing aside the neighbouring scales, advances considerably beyond them, bearing the rudiments of the new bulb on its extremity. Owing to this circumstance, the plant of each successive year rises at some distance from the site of its predecessor ; and as the old bulbs do not soon die away, one or two of these, in a decaying state, are generally found * Plate 2. fig. 3. A. The young bulb as seen at the basis of the stem, when the scales of the old bulb are removed : a. the stem ; b. the young bulb ; c. the attached fragments of the scales ; d. the roots of the present year's bulb ; e. the remains of those of last year. B. The interior of A. divided by. a longitudinal section ; a. the remains of last year's caudex ; b. the caudex of the bulb of the present year, with the vessels which nourish it anastomosing with those of the parent stem. LECT. IV.] THE ROOT. BULBS. 175 appended to the present year's bulb *. On taking off the scales of the bulb, the manner in which the runner, which bears the new bulb, is sent off, becomes extremely apparent -f- ; and the connexion between it and the stem and caudex of the adult bulb is rendered still more obvious, by inverting the cut end of the stem in a coloured solution, and making a longitudinal section of the whole ; for, as the vessels in this species of Lily are com- paratively large, they can be readily traced, by the naked eye, when filled with the colouring matter, passing from the stem into the caudex, thence into the runner, and through it to the small scales forming on its apex J. * Plate 2. fig. 4-. The bulb of the superb Lily, with a small portion of the stem of the plant, a. The bulb of the present year ; b. last year's bulb in a decaying state, with c. the remains of the bulb of the prior year still attached to it : d. the rudiments of the new bulb attached to the extremity of the succulent runner, shooting forward between the scales; e. the roots of the bulb ; f. roots given off from the stem above the bulb. f Fig. 5. a. The caudex of the bulb denuded of its scales. b. the runner projected from it, and bearing the rudiments of the young bulb on its extremity ; c. d. the remains of former years' bulbs ; e. the cut stem of the vegetating plant ; f. the roots of the adult bulb. J Fig. 6. a longitudinal section of the caudex and lateral runner of fig. 5. injected by placing the cut end of the stem in a solution of extract of Logwood : a. the caudex, the coloured spots in which show where the bundles of vessels passed off to 176 CONSERVATIVE ORGANS. [LECT. IV. It has been already stated, that each scale is not only a distinct reservoir of nutriment ; but that it is endued with such a share of vital energy, as enables it, when detached from the bulb and placed in a proper soil, to produce an entire new bulb. In this respect, the scale resembles some leaves, which I shall particularize when we treat of those organs. The scales of the Canadian, and those of the scarlet Pompone Lily, both of which are natives of high latitudes, exhibit this property more readily than any other of the tribe. The young bulb rises either from a callus formed at the base, or the margin of the scale * ; which gra- dually decays away as the bulb enlarges ; and finally separates from it as soon as a sufficient num- ber of roots are produced to support this lateral production as an independent being. The bulbs, however, which are thus propagated, differ from those formed in a more direct manner, as they pro- duce leaves only ; and several successions of leaf bulbs are propagated in a direct line from them, the scales of the adult bulb; b. the stem with the vessels injected; c. the runner, with the vessels running through it to supply the rudiments of the young bulb on its extremity ; d. a scale of the young bulb ; e. the place where the runner that supported the adult bulb was broken off. ' * Fig. 7. a scale of Lilium pomponium, with two bulbs, a. and b. forming on it. Fig. 8. a portion of a scale of Lilium su- perbum with a bulb formed on its edge ; a. a root already pro- truded from the bulb. LECT. IV.] THE ROOT. BULBS. 177 before a flower bulb is produced. It may seem ex- traordinary that no formation of young bulbs oc- curs on the scales individually, when they are al- lowed to remain attached to the old bulb ; but when we consider that in this case their vital energy, and the nutriment they contain, are ex- hausted in supplying the growing plant, and aid- ing the formation of its lateral progeny ; and that, on the contrary, when a scale is separated, the formation of the new tyulb is merely the em- ployment of the same agencies, which would have acted simultaneously with the other scales had the separated scale been left in its original state, only differently directed, the explanation is obvious. Another question, however, arises on this point. If, as has been stated, the young bulb be the production of the joint action of the bulb and the plant, how is the bulb on the solitary scale formed ? I confess my inability to solve this question ; and can only remark, that the bulb formed on the separated scale is, comparatively, a very imperfect being; and the formation of a series of bulbs in a direct line from it, each pro- gressively more perfect than its precursor, is requisite before a bulb can be produced as perfect as that one, which is the effect of the joint func- tions of the plant and of the vipe or adult bulb : and in this progression to perfection the plant of each season necessarily plays its part. VOL. I. N 178 CONSERVATIVE ORGANS. [LECT. IV. 2. The Granulated bulb (Bulbus granulatus) is named from its appearance being that of a small, globular body, or grain. It is found usually asso- ciated with many others, studding, as it were, the root to which they are affixed ; as is beautifully ex- emplified in that of grain-rooted Saxifrage, Saxi- fraga granulata *. On examining a single bulb, we find that it is composed of slightly curved granular scales, covered with two coats -j~, and enclosing the plantule, which, on vegetating, bursts the coats and shoots up between the scales. As it advances, these are gradually emptied, until the epidermis only remains like a small shrivelled leaf; whilst fresh bulbs are generated upon runners, sent off from the basis of the herbage. The minuteness of the scales prevents their structure being ex- amined by the unassisted eye ; but, with the aid of the microscope, it is perceived to be nearly the same as that of those of the squamous bulb. III. LAMINATED BULBS are composed of fleshy layers, attached at the base to a solid radical caudex. Each layer consists of a plate of cel- lular substance, filled with secreted juices, and enclosed between two cuticles. Bundles of ves- * Vide Plate 2. fig. 9. which represents the bulbs congre- gated on the runner of the plant a, f Fig. 10. a. a bulb denuded of its coats, with the plan- tule in a state of vegetation ; b. a plantuie separated from the enclosing scales, attached to the radical plate, and a small fragment of the tunic. LECT. IV.] THE ROOT. BULBS. 179 sels run vertically through each layer, communi- cating with those of the caudex, the stem, and the roots : these are entire and porous vessels ; but in some of the laminated bulbs, the spiral vessels are so numerous as apparently to make up the greater part of the substance of the layers. There are two species of laminated bulbs ; the concen- tric and the nestling. 1. The Concentric laminated bulb (Bulbuscon- centricus tunicosus) consists of sheathing laminae, each enclosing, or nearly enclosing, the other from the centre to the exterior coat or tunic. This spe- cies of laminated bulb may be subdivided into two sections ; the first comprehending those bulbs in which the layers are entire, or in which each layer is the sheath, as it were, of those within it ; and the second, those in which the layers are divided, or only overlap those within them, but do not form an entire sheath, as in the former instance. * With entire Layers. The most familiar examples of this division are the common Onion, Allium Cepa, and the mem- bers of the Narcissus tribe. In these, the cau- dex is of an irregular, serniorbicular form ; and consists of a central cellular part or pith, covered by a more solid envelop or cortex, which is inter- posed betwixt the basis of the layers and the pith, and forms also the proper radical plate, whence N'2 UNIVERSITY OF .. 180 CONSERVATIVE ORGANS. [LECT. IV. the roots are protruded. The number of the layers varies : the exterior are thin and semipellu- cid, being 1 formed simply of two membranous cu- ticles, with equi-distant bundles of vessels running longitudinally through them : the interior are more fleshy, the cells being filled with nutritious juices. A transverse section of the bulb exhibits the continuous character of each layer *. The new flower bulb is formed nearly in the centre of the old bulb, at the base of the stem ; but in general, there are several leaf bulbs formed : one or more within the second or the third layer from the surface, and others laterally and altogether exterior to the old bulb. The outer layers only of the old bulb are completely exhausted in one sea- son ; hence the bulb in several subsequent seasons appears to be the same which originally flowered, only much enlarged. The lateral bulbs separate, , and, as in similar cases, maintain an independent existence. In the Tulip, the bulb consists usually of four concentric layers only (independent of the common coat), the outermost of which may be termed fleshy ; but being nearly diaphanous, the longitudinal vessels are seen running through it at equal distances, with the intervening spaces on * Vide Plate 3. fig. 1. which represents the transverse section of the bulb of Narcissus Jonquitta : a. a. two young bulbs rising betwixt the second and third layers ; b. the roots pro- truded from the radical plate, which is hid by the position of the bulb. 4 LECT. IV.] THE ROOT. BUJUBS. 181 the inner surface studded with opaque slightly elevated glands, which produce correspondent depressions on the outer surface of the next layer. The other layers are fleshy and thick in proportion as they approach the enclosed plantule, or centre ; and, although the depressions of the glands be visible on the exterior of each, yet, from the tur- gidity of the cellular texture, neither these nor the vessels are perceptible in them ; but by placing the bulb in a coloured solution for twenty-four hours, and then making a transverse section of it, the latter are seen like coloured points arranged in a regular series, and nearly equi-distant from the enclosing epidermis of each layer*. By tearing the layer longitudinally opposite to any of these points, the vessels can be traced downwards through the substance of the layer, and even into the radical plate. In this bulb, the stem, which bears both the foliage and the flower, rises from the centre of the bulb ; but as soon as the new bulb, which is formed close to its base, enlarges, the enclosing laminae of the old one Jbegin to lose their suc- culency and plumpness ; and this exhaustion con- * Plate 3. fig. 5. a transverse section of a Tulip bulb, du- ring the flowering season, a. Section of the stem ; b. remains of the exhausted layers of the old bulb ; c. the new bulb, which should flower next season ; the dots in the layers showing the positions of the longitudinal bundles of vessels. N3 182 CONSERVATIVE ORGANS. [LECT. IV. tinues to keep pace with the enlargement of the offspring 1 , until the layers are completely emp- tied. They then decay away, and leave the stem partially covered with the void cuticles on the outside of the new bulb *. The stem itself, with the old radical plate, next die away ; while the new bulb, after remaining as it were in a state of inactivity during the winter, shoots out fresh roots from its radical plate in the spring, and runs the same course as its predecessor. Under fa- vourable circumstances, two or three new bulbs are sometimes formed in the same season ; but these are usually only leaf bulbs -f~. In long-rooted Allium (Alliurn victor tails) the concentric sheathing layers (each of which, as in several other species of the genus, is expanded into a leaf) are of a reticulated texture^; and * Fig. 6. a longitudinal section of a Tulip bulb, during the flowering season, a. Section of the stem ; b. the remains of the old bulb ; c. the new flower bulb enveloped in d. its sheath or exterior coat ; e. the rudiment of the caudex of the new bulb, where it separated from the parent stem. -p Plate 3. fig. 4. a bulb of the common Tulip taken up in the flowering season, a. Part of the stem, which is cut off; b. the exterior layers of the exhausted bulb, enveloping both the new flower bulb c. and an offset, or lateral leaf bulb, the stem from which is seen at d. t Fig. 2. a bulb of long-rooted Allium. a. The outer layer of the present year's bulb, extended into the sheathing stem ; b. the reticulated layers, loose both at the caudex c. and above ; d. the roots of the existing bulb piercing the network of the reticulated layers. LECT. IV.J THE ROOT. BULBS. 183 when exhausted of the nutriment contained in them, assume the appearance of lacework. In this bulb, also, the young bulbs are formed within the second layer of the present year's bulb *. * * With divided Layers. In this division of the concentric laminated bulbs, the layers, as has been already stated, do not form an entire sheath. In the garden Hyacinth (Hyacinthus orient alls), which I select as an ex- ample because it can be easily procured, they comprehend two thirds only of the circumference of the circle. Each layer is fleshy and thick in the centre, and becomes gradually thinner towards its edges until it terminates in a membranous film, embracing the layers within it in the same manner as the hand holds a ball which it cannot completely enclose. The deficient portion of each layer is always on the opposite side to that of the layer immediately beneath it. The caudex has the character of a segment of a hollow sphere, on the convex surface of which the layers are attached ; whilst a considerable number of fleshy roots protrude from the circumference -j~. The * Fig. 3. a vertical section of the same bulb. a. the suc- cessive succulent layers extended into the stem ; b. the ex- hausted reticulated layers ; c. the caudex ; d. the young bulb. f Vide Plate 3. fig. 6 and 7. a. The body of the bulb ; b. the manner in which the layers overlap each other ; c. the radical plate, with the roots separated from one side of the cir- N4 184 CONSERVATIVE ORGANS. [LECT. IV. vessels run in longitudinal bundles through the layers ; and so numerous are the spiral vessels in this description of bulb, that, in the Amaryllis tribe, the Squill, and some other of the larger bulbs, on breaking a layer transversely, they may be drawn out to the distance of three or four inches ; and from the strength of their component fibres and their number, they are capable of sup- porting the pendulous portion of the layer. I have also been informed that the fibres of the spiral vessels of the bulb of some species of the Blood-flower (Haemanthus) have been spun into thread and manufactured, which I can readily be- lieve, from the strength of a thread which I have formed from the spiral fibres of the layer of a bulb of Brunsvigia toxicaria, Poison- bulb, by merely twisting them in my fingers. On making a trans- verse, or a longitudinal section of the Hyacinth' bulb, we find one or more young bulbs seated be- tween the layers * ; and that one which is nearest cumference, to show its concavity ; d. young lateral bulbs, which produce leaves, and si new bulb when detached from the parent ; e. a bulb, which has produced leaves this season, formed within the coats of the old bulb. * Vide Plate 3. fig. 8. and 9.-- Fig. 8. a. The caudex, as it appears in the longitudinal section of a full-grown bulb of Hya- cinthus orientalis ; b. a young bulb forming with the outermost fleshy layer; c. the remains of the stem and foliage of the pre- sent year's plant. Fig. 9. a transverse section of another b*lb of the same species of plant, showing the terminations of the LECT. IV.] THE ROOT. BULBS. 185 to the centre of the old bulb is, usually,, the real flower bulb for the next season ; for, although, ow- ing to a few of the outer layers only decaying, the old bulb appears still to yield the flower, yet, this actually springs every year from a new bulb formed within the old one. Young leaf bulbs are also protruded exteriorly from the base of the old bulb ; and these appear to be really the production of the bulb itself ; for the Dutch cut their bulbs trans- versely, after they have attained a certain size ; and plant the lower half, in order to multiply the number of the lateral bulbs ; whilst the flower bulb, as has been already stated, is evidently the result of the combined functions of the plant and the bulb. No experiments, at least that I know of, have been made, to ascertain whether a layer of any of the concentric laminated bulbs, when separated from its caudex and planted, will produce young bulbs, as is the case with the scales of the squa- rnous bulbs ; but I am of opinion that if an en- layers, and the divided bundles of longitudinal vessels, a. the lower portion of the flower stem ; b. the bulb which will flower next year, gradually expanding the old bulb by its growth ; c. the roots attached to the caudex, which are hid by the position of the superincumbent parts. Fig. 7. another bulb of the garden Hyacinth in which four young lateral leaf bulbs d. are seen seated on the caudex, to show the manner in which these lateral offsets generally occur. 186 CONSERVATIVE ORGANS. [LECT. IV. tire layer were carefully separated from its caudex, and planted, it would produce young bulbs upon its basis. 2. The Nestling bulb (Bulbus nidulans) is com- posed of entire concentric layers, which termi- nate in sheathing leaves, and enclose within each, from the circumference to the centre, several young- bulbs ; so that, when the layers are partially rup- tured, the whole seems to consist of small bulbs, and has a nestlike appearance *. The layers are more membranous than those of the other lami- nated bulbs we have examined ; and consequently display, more evidently, the longitudinal equi-dis- tant bundles of spiral and entire vessels. The caudex is very vascular ; in form nearly an inverted truncated cone, and protruding a great number of long threadlike roots. On dissecting the young bulbs, enclosed within the old one, we find that those only which have already shot up into leaf, contain small bulbs or their rudiments within the layers ; hence, there is reason for concluding that the young bulbs are in this instance, chiefly the result of the functions of the plant. In the bulb of Garlic, Allium sativum, * Vide Plate 3. fig. 10. the recent bulb of Garlic, Allium sativum, with the sheathing layers partially removed to exhibit the young bulbs, a. The young bulbs ; b. c. d. e. remains of the sheathing layers \f. the caudex. Fig. 1 1. a longitudinal section of the same bulb. a. the caudex ; b. c. d. the sheathing layers, which terminate in leaves. LECT. IV.] THE ROOT. BULBS. 187 the odorous, acrid, secreted juice which character- izes it, is found in greatest quantity in the fleshy internal layers of the young bulbs * ; and conse- quently these are the parts employed both medi- cinally and as a condiment ; whilst, in the mem- branous layers of the old bulb, it is scarcely per- ceptible either to smell or to taste. Such are the characteristic distinctions of those appendages of roots, which are denominated bulbs. They have been properly regarded by Dr. Darwin -f and others as subterraneous buds ; and as having a close affinity with the buds on the stem and branches of perennial plants : for, like these, they continue rather than reproduce the individual, and are liable to all the hereditary im- perfections and diseases of the parent. In some other particulars also this resemblance holds good; for, as in all plants which are furnished with buds, those produced in the first years of the plant are leaf buds only, and a succession of these occur be- fore flower buds are formed, so no flower bulb is ever produced from seed ; the first formed being always a leaf bulb, which, in many instances (as the Tulip for example), produces another leaf bulb, and so on for several successive generations, until at length a flower bulb is formed ; after which, one * In common language these young bulbs are termed cloves. f Phytologia passim. 188 CONSERVATIVE ORGANS. [LECT. IV. flower bulb at least is produced every season. Many circumstances connected with the physi- ology of bulbs might be treated of in this place ; but, as I shall have again to recur to them in no- ticing those bulbs which are formed on the stem, and among the flowers of many plants, I shall defer at present the further consideration of this part of our subject. And being now acquainted with the forms which characterize roots and their appendages, we shall, in our next Lecture, finish the consideration of these vegetable organs. LECT. V.] THE ROOT. 189 LECTURE V. THE SUBJECT OF THE FORMER LECTURE CONTI- NUED. OF SOILS AND MANURES. OF THE MEDI- CINAL AND DIETETICAL PROPERTIES OF ROOTS. \ I HE usual situation of roots is in the ground ; but many plants, although their seeds be sown in the earth, yet, will not vegetate in it, their proper soil being the bark of other living plants. Such are named parasitical, owing to their nou- rishment being obtained from those plants on which they fix, and which they rob of a part of their juices, often injuring them to a very consi- derable degree. The Misletoe (Viscum album) ; the Broom Rape^ Orobanche ; the majority of Lichens ; the Mosses ; some of the Ferns ; many of the Orchis tribe ; those minute fungi, which produce the diseases of corn and of some grasses, known by the names of rust, blight, and mildew ; the Sclerotium crocorum, a sort of tuber which attacks the bulb of the Saffron ; . and the Dry Rot, to the destructive powers of which the noblest specimens of architecture occasionally fall sacri- fices ; are parasitic plants. Some of this descrip- tion of plants, however, originally grow in the earth, and do not lose their attachment to it 190 CONSERVATIVE ORGANS. [LECT. V. until they find another plant to lay hold of, and into which they can dip their caulinar roots, or rootlike absorbents, which are protruded from the stem, in order to share its nutriment, and on which they are afterwards supported ; as, for ex- ample, the Cuscuta, or Dodder *, which may be regarded as the natural parasite of our indigenous Heaths and Hops. Some plants, after they have arrived at a cer- tain age, do not even require that their roots should be fixed to any spot ; but maintain life on what they can procure by absorption from the at- mosphere. Such are the Cacti, a curious tropical tribe of succulent plants ; on which account one of the species, the Indian Fig, Cactus opuntia, was recommended to the notice of seafaring people, by the late Dr. Anderson of Madras, for the purpose of supplying vegetable food on long voyages ; and as a preventive of scurvy. But the most curious instance of this kind is the aerial flower, Epidendrum flos ae'ra-f-, an East Indian parasitical plant, which continues to grow, blos- soms, and even perfects its seed, when it is torn * The Dodder germinates in the earth, and rising above it, shoots out filiform stems, which twine around the neighbouring plants. Its original root now decays, and a kind of warty roots are formed in the stem at every point where it touches the sup- porting plant by which it is nourished. Aerides matutinum of Willdenow. LECT. V.] THE ROOT. 191 from the tree on which it originally grew, and is suspended in the ceiling of an apartment *. Many aquatic plants, also, have roots which serve no other purpose than to fix them for a short time to the spot where they have germinated, from which they afterwards separate and float upon the surface of the water; thus the common Duck Weed, Lemna minor, which rises to the surface almost as soon as it has germinated, has filiform roots from three to six inches in length, which hang perpen- dicularly in the water, and having no attachment to any body, allow the plant to float freely in every direction. * Mr. Macnab, superintendent of the Botanic Garden at Edinburgh, has published an account of a suspended plant of Ficus Australis (Ferrugineous Fig), which had grown for eight months, up to the time of publishing his paper, February 7th, 1820, without earth, in the stove of that garden. The plant, which was originally growing in a pot in the greenhouse, on being removed into the stove, and treated in a peculiar manner, threw out several roots from the stem ; after which the earth was gradually removed from the original roots, and the plant left suspended in the air, affixed to the frame of the stove. Water was, however, sprinkled over the whole plant every day. " What," adds Mr. Macnab, "may appear rather remarkable is, that though this Ficus is a plant by no means free in pro- ducing fruit in the usual way of cultivating it, this specimen, quite suspended, without a particle of earth, was loaded with Figs during the months of September, October, and part of No- vember. 1 * Edinburgh Philosophical Journal, vol. iii. p. 77. 192 CONSERVATIVE ORGANS. [LECT. V. All subterranean roots assume a particular di- rection ; which is constant in every individual of the same species of plant. They are said to be perpendicular (perpendicular is) when the caudex, or main body of the root,, extends perpendicularly into the ground, as exemplified to the majority of fusiform roots (p. 130), and in the main or top root of most trees : and horizontal (horizontalis) when the extension is nearly parallel to the plane of the horizon, so that the root forms nearly a right angle with the stem or herbaceous part of the plant ; as, for instance, in Winter Green, Pyrola umbellata, Sweet Flag, Acorus calamus, and in the majority of the articulated roots. Many of the horizontal roots, whilst they run under the surface of the ground, push up stems at intervals so as to multiply the plant ; tnence the appellation creeping roots (R. repens) : as in common Spear Mint, Mentha viridis ; Couch Grass, Triticum x. repens ; Bulbife- rous Coral Wort, Dentaria bulbife- ra(fig.x) 9 and ma- ny other plants. And this proper- ty renders some weeds extremely obnoxious to the farmer : for, if any portion of a creeping root be LECT. V.] THE ROOT. 193 left in the soil, it will throw up a new stem, which in its turn produces an extension of the root. But if it be injurious in some instances, it also serves very important purposes in the economy of nature. Thus the sand Reed-Grass, Carex arenaria, by means of its creeping roots, binds together the dry sand of the flats on the sea-shore where it grows ; and assists not only in forming a fertile soil where sterility would otherwise reign ; but by preventing the sand from drifting, preserves from destruction the neighbouring fields, which already repay the labours of the agriculturist. We must not, however, confound the creeping with the progressive root (R. progrediens) , which, in extending itself, likewise shoots up stems and herbage at intervals ; since it differs in this cir- cumstance, that as it advances anteriorly it decays posteriorly; thus causing an obvious progression of the plant ; and at the same time limiting in a considerable degree its multiplication. Its direc- tion also is not necessarily horizontal, like that of the creeping root, but may be perpendicular ; the cause of the progression in that case being the production of a new root laterally : and, in this respect, these roots, although the caudexes are not tubers, yet have a close affinity with the attached tuberiferous roots. Thus the root of officinal VOL. i. o 194 CONSERVATIVE ORGANS. [LECT. V. Monkshood, Aconitum neomontanum (fig. ?/), af- fords an excellent example of the pro- gressive conical root; a. the old root supporting b. the lower portion of the stem ; c. the new root attached by the lateral offset d. to the basis of the stem b. : so that the new stem, which should have arisen from the bud e. would have been about one inch from the old plant, had it been left in the ground. The moniliform roots (p. 142), also, are progressive ; as those of Paeony, Pseonia officinalis, and night smell- ing Geranium, Pelargonium triste ; but in these the progression is made by suckers thrown up from the nodules ; such roots in this respect re- sembling the tubers on the pendulous tuberiferous roots. The difference, however, between the progressive roots and the tuberiferous, is very ob- vious; the caudexes and nodules in the former being real roots, furnished with lateral rootlets issuing from their surface; whilst the tubers, in LECT. V.] THE ROOT. 195 the latter, are appendages only to the real roots, which are fibrous arid issue from the basis of the stem, or the herbaceous part of the plant, and not from the surface of the tubers. Besides these roots which take the perpendi- cular, or the horizontal direction, there are some that assume an intermediate one, in which case the root is said to be oblique (R. obliqua) ; but, as in general, there is always an approximation either to the perpendicular or the horizontal, the truly oblique root is .very uncommon. In observing the position of roots in the ground, it should be recol- lected that it is the main root or caudex only which determines this point; and that, whatever its di- rection may be, that of the rootlets is always dif- ferent: thus, in perpendicular roots, the rootlets spread either horizontally or obliquely ; whilst in the horizontal and the oblique they descend per- pendicularly ; or, in every instance, they form an angle more or less acute with the surface of the caudex. The main root should be entire, also, be- fore its direction or character can be accurately ascertained ; for, if the apex be destroyed by any means, the caudex no longer elongates ; but sends off lateral shoots, which necessarily take an op- posite direction from that which is natural to the root. The situation of the root, in regard to the stem or the herbaceous part of the plant, is ge- o2 196 CONSERVATIVE ORGANS. [LECT. V. nerally at the base ; but roots originate from that part also of the stem which is above ground. Thus the Strawberry throws out lateral shoots, which are termed wires, from which roots descend at in- tervals into the ground ; and from the stems of the Ivy, the Jasmine, the Ash-leaved Trumpet- flower, Bignonia radicans, and other plants which fix themselves to walls and rocks, roots are pro- truded,, serving equally to imbibe nourishment and to give support to those climbing plants. Whenever a stem is surrounded with earth, roots are protruded from it. Du Hamel filled a cask with earth, and boring holes through the bottom of it, supported it on stakes three feet from the ground. He then pushed slips of plants through the earth in the barrel, and planted their ends, which passed out at the holes in the bottom of the cask, in the ground below it. These ends took root ; the parts between the ground and the cask put forth branches and leaves ; those surrounded by the earth in the cask protruded roots ; and those, again, above it became clothed with foliage. It is, also, well known that, if a twig of a Willow be bent and each end of it stuck into the ground, roots will be protruded from both extremities, whilst the middle or arched portion will be covered with leaves and branches ; and the extremities of the branches of all ligneous plants throw out roots, LECT. V.J THE ROOT. 197 if they be bent down and laid under the earth. In- deed it is not absolutely necessary to lay them un- der the ground to produce this effect ; for a ligne- ous plant growing in a sterile soil, provided the situation be shady and the air damp, will throw out roots from its branches, resembling those protruded by Ivy. I have in my possession a por- tion of the branch of a Bay, Laurus Indica, in the whole length on one side of which the bark is rup- tured, owing to the protrusion of numerous short rootlets. The plant from which this branch was cut, was growing (in 1814) in a kind of cave, formed by the intertwining roots of a noble Beech, on the summit of a chalky bank, in Lord Derby's grounds, near Epsom. The chalk had mouldered away from under the roots of the Beech, which, projecting forward as a roof, shaded the Bay that, perhaps, had been originally planted close at the foot of the bank : but all the soil had been long since washed away from its root, which adhering to the bare chalk, served no longer as an ab- sorbing organ, but merely to sustain the plant in its upright , position ; and as there were very few leaves on the branches, the rootlets, which were protruded from them, evidently maintained the life of the plant, by absorbing the water held in solution in the air of the shady spot where it grew. On the knowledge of facts such as these, plants are propagated by what are called layers, an opera- o3 198 CONSERVATIVE ORGANS. [LECT. V. tion, which consists in bending down a branch, or the s.tem of the plant, so that the knee or bent portion of it can be preserved, by means of pegs, under the surface of the soil. The part so treated is nourished by the parent plant, until roots are sent off from the part under the ground, when it is cut away, and thus becomes an independent plant. Some plants naturally propagate, or rather extend themselves in a similar manner. Thus, from the branches of the Banyan or Indian Fig- tree (Ficus indica), fibres are thrown out, which hang suspended like icicles, and grow thicker as they reach the surface of the ground, into which they strike root and become trunks, the branches of which root again in the same manner : and this progression of increase is continued until the ground is covered to a prodigious extent with an umbrageous labyrinth or grove, formed from one original trunk, impenetrable to the sunbeams *. One of these trees, called Cubber Burr, situated on an island in the river Nerbedda, exceeded 2000 feet in the circumference of its shade ; and in 1787, had 350 trunks. Religious festivals were held under its luxuriant canopy, which was ca- pable of affording shelter from the solar heat to * Pagodas are generally built in the neighbourhood of these trees; and under their friendly shade the Brahmins and de- votees perform their religious rites. LECT. V.] THE ROOT. 199 7000 persons. MILTON has immortalized the Banyan, by describing it as the tree under which our first parents retired to hide themselves after their fall : They chose The Fig-tree ; not that kind for fruit renown'd, But such as, at this day to Indians known In Malabar or Decan, spreads her arms Branching so broad and long, that in the ground The bended twigs take root, and daughters grow About the mother tree, a pillar's shade High over-arch'd, and echoing walks between : &c. Paradise Lost. The structure of roots does not differ materially from that of the trunk and branches ; we may, therefore, reserve the minute examination of that part of our subject until we come to treat of these organs ; and, at present, notice only their general structure *. All roots are either ligneous or fleshy. The ligneous belong to trees and shrubs, and are composed of an epidermis or scarf-skin, a cutis or bark, a vascular system, woody matter, and pith. The fleshy, which belong to herbaceous * The analogy, indeed, is so close, that a tree may be in- verted so as to change the roots into branches, bearing leaves and flowers, and the branches into roots producing radicles. This fact has been frequently proved, by repeating the experi- ment (first tried by Du Hamel) of planting a tree with the branches in the ground, and leaving the roots in the air ; after a season the buried branches produce radicles, and the roots raised in the air give out buds, stems, and leaves. o 4 -00 CONSERVATIVE ORGANS. [LECT. V. plants, consist chiefly of cellular and vascular textures, interspersed with slender bundles of woody fibre. In both the epidermis or exterior covering is the same, and seemingly destitute of vessels : it is present in every stage of the growth of the root from its origin, on the radicle of the embryon, to the tangled rugged arms which rivet down the venerable monarch of the wood, the pride of centuries : it serves the office of a filter, and allows nothing to pass through its pores that cannot enter the minute mouths of the absorbent vessels which open on the surface of the bark beneath it. The epidermis of roots is thus adapted for its situation in the moist soil; but, in other re- spects it does not differ from that which is spread over the stem, branches, leaves, and every other part of the plant, thejibfik, which are attached to all roots, whatever may be their figure, are, as has been already stated, mere bundles of vessels enclosed in a cuticular membrane ; and, as they take up from the earth the food of the plant, they are, in fact, the real roots ; whilst the caudex may be regarded as a magazine, in which the food that has been elaborated into the proper juice is de- posited for the particular uses of the vegetable economy ; and it is this deposition which affords the colour and odour which distinguish different roots. With regard to the mode of growth of the LECT. V.] THE ROOT. 201 root, Mr. Knight has advanced sufficient reasons for believing that this organ is not an elongation of the radicle; but is formed upon it after the germination of the seed by successive increments of new matter deposited at its apex ; for, whilst the root never elongates by the extension of al- ready organized parts, the radicle does elongate throughout all its parts in the germinating seed. The root, at first, consists of cellular matter only, contained in an epidermis, within which the cor- tical vessels are afterwards generated ; the circle of vessels, says Mr. Knight, " enclosing within it a small portion of the cellular substance, which forms the pith, or medulla of the root ; and these vessels gradually generate alburnum, which, in a transverse section of the root, appears arranged in wedges round the medulla." The same able phy- tologist further maintains that root shoots differ from stem shoots in not being emitted from the alburnum ; and that the presence of the alburnum is not essential is evident from the fact, that leaf- stalks, which contain no alburnum, nevertheless emit roots ; and these derive their existence, as in all other cases, from the proper juice conveyed in the returning vessels *. All roots have originally more or less of a tapering form ; which is partly the consequence * Phil. Transactions, 1809. 202 CONSERVATIVE ORGANS. [LECT. V. of their mode of growth ; for, as has been just stated, a root, in extending, is not lengthened through all its parts, but by additions made to the extremity, and as these, in the early stages of its growth, are made more rapidly than the alburnous depositions, which add to its diametrical dimen- sions, its shape must necessarily be tapering. That the longitudinal extension of the root is ef- fected by the deposition of new matter at its ex- tremity, was first ascertained by Du Ham el *, who having passed small pieces of silver thread trans- versely through a vegetating root, at distances which were accurately measured, found that the upper threads, those nearest to the stem, or herb- age, retained their original and relative situation, whilst that one only which was very near to the end of the root was carried down ; and this is the case both in succulent and ligneous roots. The lateral extension, on the contrary, in succulent roots, appears to depend on the deposition of ad- ditional particles, throughout the whole of their substance ; and in perennial and ligneous roots, by the formation of an annual new alburnum, in the same manner, as I shall explain to you in its proper place, occurs in the stems and branches of trees and shrubs. When the extremity of a root is destroyed, or when the root has attained the * Phys. des Arbres, liv. I. c, v. LECT. V.] THE ROOT. 203 natural limit of its longitudinal extension, it throws out lateral branches ; and these extend with most vigour in that direction in which the most abundant supply of nourishment is to be found ; a circumstance which has been very unaccountably attributed by some to a sentient, or an instinctive principle in plants. The fact is, that the influence of a quantity of manure, or of richer earth, is not confined to the immediate spot where it is de- posited ; but extends to a certain distance in every direction, diluted, as it were, by the poorer soil, until its power is altogether dissipated : as soon, therefore, as any branch of a root impinges upon the limits of this circle, it obtains a supply of nu- triment capable of exciting its vital energy in an increased degree : and as this augments in a direct ratio, as it extends, this branch in length and vigour necessarily far exceeds those on the opposite side of the main root, which have had a more scanty supply of nourishment. Thus, also, if a seed of a tree, conveyed by the wind, or other- wise, as may accidentally happen, be planted on the top of a wall, its roots will gradually extend until they reach the ground ; whilst the upright growth of the tree will be very scanty previous to the root establishing itself in the soil. Sir J. E. Smith, in referring to a fact of this kind, communicated to him by the Rev. Dr. Walker of Edinburgh, regard- ing an Ash, which grew on a wall in the Canon- 204 CONSERVATIVE ORGANS. [LECT. V. gate in that city, and which I have seen, explains it in the following manner: " Here the vital powers of the tree not being adequate, from scanty nourishment, to the usual annual degree of increase in the branches, were accumulated in the root, which, therefore, was excited to an ex- traordinary exertion, in its own natural direction downwards." I would, however, say that it was owing to the moisture of the earth soaking into and ascending the wall, a fact which, from every day's experience, we know takes place, that the root extended in the direction of its natural stimu- lus, the moisture ; the real exciting cause of its increased exertion : and the accuracy of this opinion is placed almost beyond controversy, by the experiment of Mr. Knight, which I am about to relate. Some Beans were placed in pots filled with earth, but were half covered only with the mould. The pots were then inverted on a grating of wood, so as to support the earth and the Beans, in such a manner, that the earth was above and the air beneath each radicle as it was emitted. Water was next introduced through the bottom of the inverted pots : the radicles extended horizontally along the surface of the mould, and in contact with it ; and in a few days emitted many fibrous roots upwards into it, which passed through one half of the mould. We may, therefore, venture to assert, Jhat although the natural direction of the radicle LECT. V.] THE ROOT. 205 is, in every instance, downwards, depending on causes which have not yet been ascertained *; and although roots, during their growth, assume va- rious directions, which invariably occur in the same kinds of roots, yet, that circumstances con- nected with their mode of growth, produce devia- tions, and affect the uniformity which might otherwise be expected. But many roots not only vary from their natural direction, but even change their characteristic figure, owing to circumstances connected with the state of the soil : thus, insu- lated trees, that are much agitated by the wind, have strong lateral roots, forming a kind of net- work that adheres firmly to the earth ; whilst the same description of trees, growing in a forest, has long slender tapering roots. This diversity may be thus explained : when a tree stands alone, the soil near the surface is partially loosened by the agitation of the tree by the wind, and conse- quently less opposition is afforded to the extension of the lateral branches of the root, than the un- moved ground offers to that of its perpendicular caudex ; whereas, when a tree grows in a forest, the mechanical obstacles offered to the extension of the lateral roots by the neighbouring roots, ex- * The different opinion^ that have been hazarded on this subject shall be noticed, when we take under our consideration the germination of seeds. 206 CONSERVATIVE ORGANS. [LECT. V. ceed those which the ground offers to the per- pendicular. Some Grasses, also, as the common Gat's Tail, Phleum pratense, floating Fox Tail, Alopecurus geniculatus, and others, which in a wet situation have fibrous roots, became nodose, when planted in a dry sterile soil : the cause of which cannot be better explained than in the words of Sir J. E. Smith. Presuming the herb to be starved, he adds, " by a failure of the nutri- mental fluids hitherto conveyed by the water of the soil, its growth would be checked ; and when checked, the same growth could not, as we know by observation on vegetation in general, be in- stantaneously renewed. A sudden fresh supply of food would, therefore, cause an accumulation of vital energy in the root, which would consequently assume a degree of vigour and a luxuriant mode of growth not natural to it, and become bulbous. Thus, it acquires a resource against such checks in future, and the herb is preserved alive, though in a very far less luxuriant state than when regularly and uniformly supplied with its requisite nourish- ment *." The nature of the Jibrils has been already mentioned. They may be regarded as the mouths of the plant : for, the extremities of their vessels being open, these suck up such fluid nutriment * Introd. to physioL and systematical Botany, p. 115. LBCT. V.] THE ROOT. 207 from the soil as can pass through the pores of the epidermis, and conduct it to the caudex, or main root, through the vessels of which it ascends to the leaves : there it is as it were digested ; and, being changed in its properties, is again conveyed by another set of vessels to the caudex,, in which it is deposited for the future exigencies of the plant. As a proof that the fibrils are the only parts of the root which take up the nutriment of the plant from the soil, Mirbel remarks, that " herbs perish at the foot of young trees, because the fibrils issuing from the collet (the point of connexion of the stem and the root) exhaust the ground ; but old trees extending their vigorous roots to a distance, allow the plants, which are close 'to them, to subsist and destroy those which are more distant." It has not yet been ascertained whether the fibrils are strictly annual productions, an opinion which was maintained by Du Hamel *, Mirbel -f^, and Sir J. E. Smith;}:, and adopted by Wildenow ; but which is doubted by Mr. Knight, who, although he admits that in roots of trees, or ligneous plants, crowded together in a garden- pot, the fibrils are often found lifeless in the suc- ceeding spring ; yet, remarks that he has "not * Phys. des Arores. j- Traite d'Anatomie. J Introd. to pkysiol.and system. Botany. Principles of Botany , Eng. trans. 260. 208 CONSERVATIVE ORGANS. [LECT. V. observed the same mortality to occur, in any degree, in the roots of trees when growing, under favourable circumstances, in their natural situa- tion *." My own experience does not authorize me to decide with confidence on this question; but it rather induces me to incline to the opinion that they are annual productions. In duration, roots are either annual, biennial, or perennial. Annual roots belong to those plants which are produced from the seed, grow to their full extent, and die in one year or season ; as, for ex- ample, Barley, Hordeum secale; the White Poppy, Papaver somniferum ; the common Pea, Pis urn sativum ; the garden Bean, Vicmfaba, &c. : bien- nial to those that live through the winter of the year in which they are produced, and after flower- ing and yielding seed, die in the following year, as the Carrot, Daucus car of a ; the genus Teasel, Dipsacus ; the Canterbury Bell Flower, Campa- nula medium, &c. : and perennial to those that blossom and produce seeds through many suc- cessive seasons, as the majority of herbaceous plants and all shrubs and trees. But the life of annual and biennial roots may occasionally be protracted much beyond its natural period, when any circumstance occurs that can prevent the plant from flowering, or, even when it does flower, * Phil. Trans. Part 1. 1809. LECT. Y.] THE ROOT. 209 from perfecting its seed. Thus I have preserved the life of a Sweet Pea until after Christmas, when it was destroyed by the severity of the weather, by nipping off the flowers as soon as they were fully blown ; and plants of tropical climates, which are naturally biennial, sometimes live for many years in our hothouses ; but they all invariably die after they have produced fruit. The cause of this is, that the vitality of the plants of this de- scription seems to be sufficient for continuing their life only till after the formation of the seed, the natural means of perpetuating the species ; in per- fecting which, the irritability and life of the in- dividual are completely exhausted; and, with the plant, the root perishes. In perennial plants, the fibrils only annually perish and are renewed ; they decay before the leaves fall in autumn ; and are again formed in the early part of spring. At least this is the opinion commonly received. The root, however, enjoys more vitality than any other part of the plant, and can reproduce all the other parts, when the tree is cut down or otherwise destroyed; except in the Pine tribe and some other dry resinous plants. Such is the ROOT an organ of the greatest importance, whether we consider it simply as fixing the plant in the ground, and enabling it to elevate its leaves and flowers in the air ; or, in a more important point of view, as selecting and taking up VOL. I. P 210 CONSERVATIVE ORGANS. [LECT. V. from the soil the materials fitted for the nourish- ment and support of the vegetable body. It is a wise provision of nature, that as plants are not endued with volition and extensive loco- motion, nor guided by instinct nor reason, they are subject to more regular and unalterable laws than the animal creation, at least than that portion of it which possesses those functions which have just been enumerated : their food is always placed within their reach, and they enjoy good health and arrive at perfection in their growth, in- dependent of external accidents to which animals are equally liable, when they are situated where the soil contains those principles which are best adapted for the various purposes of their economy. The consideration of this fact suggests the ques- tions, What is the composition of soils ? What part of soils is taken up as food by the roots of plants ? To answer them has long employed the attention of the philosophical observer, and many and very various opinions have been given to the public ; but it is only since modern chymistry made those discoveries which may justly be re- garded as the most splendid triumphs of experi- mental Science, that any thing rational and satis- factory has been advanced. I will endeavour to lay before you as clear a view as I am able of the most probable conclusions which may be drawn from these opinions ; and, in raising one corner of ECT. V.j THE ROOT. 211 the veil, remove much of the mystery with which, to ordinary observers, the subject appears to be enveloped. An investigation of this kind is as useful to the physician as to the botanist and agri- culturist; for, many of the exciting causes of diseases, particularly of epidemics, are to be looked for in the nature of the soil and other local circumstances connected with the situations where they originate. The fact cannot be too often repeated and im- pressed on your minds, that plants are living beings, possessed of powers which enable them to convert into their own material substance, matters of a nature apparently very different from it. Without keeping this in view, we should be forced to look for all the different productions of plants ready formed in the soil where they grow, and to suppose that these are simply taken up by their roots, and deposited in the different parts of the plant : an idea too incongruous to be admitted. On the contrary, they do not even take up those principles which are most abundant in the soil where they grow; bat select peculiar parts of them, although these are not found, in general, forming in their uncombined state any part of the vegetable frame. Linnaeus himself, however, I believe, and many others, have imagined, that every soil held in it something which is peculiarly the proper food of every kind of plant that can be p2 212 CONSERVATIVE ORGANS. [LECT. V. cultivated on it ; and thus that poisonous plants extract something, on which their hurtful proper- ties depend, which is not taken up by wholesome plants ; or, that the secretions of plants do not vary in their qualities on account of the difference in the action of the vessels which secrete them ; but owing to their components being already pre- sent in the soil. That this is not the case, how- ever, has been clearly proved by the discoveries of modern chymistry, which have enabled us to analyse both the soil and the vegetables that grow upon it. By its assistance the mode of investigat- ing the subject has been simplified, and more sa- tisfactory results obtained. The ultimate components of all the various substances produced by vegetables have been found the same, differing only in the quantity and the mode of their combination ; and the parts of a soil which supply these have been found to be much fewer than was previously supposed. As we formerly asserted, when noticing the nature of sap, if this juice could be obtained near enough to the extremities of roots, or in the fibrils by which the soluble part of the soil is absorbed, then we should be able, by a careful analysis, to ascertain the real nature of the substances ab- sorbed; and, by looking for these in soils, know how to supply their deficiency, or to diminish their superabundance. But as an accurate knowledge 4 LECT. V.] THE ROOT. SOILS. 213 of the components of sap cannot be obtained, owing- to that fluid, as it advances, even in the vessels of the root, dissolving some already di- gested vegetable matter, which had been depo- sited there in the preceding autumn, we are obliged to form conclusions certainly not free from error ; and to content ourselves with an approxi- mation only to the truth. From the knowledge, however, which we do possess I will endeavour first to point out to you the known general com- ponents of natural soils ; secondly, what part of these is taken up as food by plants ; and thirdly, in what manner, and by what means, soils are improved and rendered more productive ; or to in- vestigate the general nature of manures. Every soil fit for yielding nutriment to vege- tables may be supposed to consist of earth, water, air, a small proportion of metallic oxyds, and de- composing vegetable or animal matters, in which are included salts, gases, and vegetable extracts. Earth, which is the essential basis of all soils, is, as it is commonly spoken of, a compound of different earths ; the most general of which are Calcareous earth, Argillaceous earth, Siliceous earth, Magnesian earth, and Ferruginous earth. 1. CALCAREOUS EARTH comprehends lime, usu- ally combined with carbonic acid, in the state of limestone, chalk, shells, and marl which is a mixture of 'carbonate of lime with clayey and p3 214 CONSERVATIVE ORGANS. [LECT. V. sandy matters ; but lime is sometimes, also, found in combination with sulphuric acid, forming a substance called gypsum ; and more rarely with phosphoric acid. When too much calcareous mat- ter is contained in a soil, it is unfertile, owing to its absorbing moisture, and consequently remain- ing too dry. But the case is different when the calcareous matter is mixed with silica, for then the moisture absorbed remains in a free state, and not so united with the chalky matter as to disappear and be useless to plants. But the absorbing pro- perties of all calcareous soils are not alike ; and a great difference depends on the degree of com- minution of the calcareous matter. Thus, 100 parts of calcareous sand retain, according to Pro- fessor Schiibler's experiments, 29 parts only of water, whilst 100 parts of the same matter in the state of fine powder retain 85 per cent. In the first case, when calcareous earth and silica predominate in an arable field, they produce a hot and dry soil; when in the second, a moist and cold soil. 2. ARGILLACEOUS EARTH comprehends clay, which is generally mixed with siliceous sand and mineral substances, and is very retentive of mois- ture. 3. SILICEOUS EARTH is almost entirely com- posed of sand. The water passes so readily- through it, that very little is retained for the pur- LECT. V.] THE ROOT. SOILS. 215 poses of vegetation ; and soils which contain much of this earth are, therefore, barren and unpro- fitable. In the form of sand it retains 25 per cent, only of water ; while 100 parts of it, as it occurs with clay in an arable field, retain 280 per cent, of water. 4. MAGNESIAN EARTH is not so commonly found as the earths we have already noticed. The magnesia it contains is combined with carbonic acid, and mixed with siliceous particles. It ap- proaches nearest to the nature of the clayey earths in its power of retaining moisture ; that power enabling it to retain 4^ times its own weight of water. This renders it, when it predominates, very prejudicial to vegetation ; while it increases, when added in moderate proportion, the fertility of a dry sandy soil. 5. FERRUGINOUS EARTH consists of those oxyds of iron known by the names of ochres and pyrites mixed with siliceous matter. These oxyds, in parti- cular the pyrites, when in any considerable quantity in a soil, if it contains little calcareous matter, are extremely injurious to vegetation. The pyrites is a compound of sulphur and iron, and is converted by exposure to air and moisture into sulphate of iron, which destroys plants by over-stimulating them. Vegetable earths have the least specific gravity, and sandy soils the greatest, whether they be dry or moist: the vegetable earths contain, besides p4 216 CONSERVATIVE ORGANS. [LECT. V. vegetables in a state of decay, animal matter and a large proportion of salts, which are chiefly com- mon salt, sulphates of magnesia and of potash, nitrates of lime, and carbonates of potash and of soda. Such are the earths generally contained in soils : when any one of them abounds, the com- pound earth is named after this component ; as for instance, a calcareous soil, an argillaceous soil, &c. The principal difference which characterizes these various kinds of earths, is their power of retaining the next component of soils, WATER. Water, as forming a part of soils, is either che- mically combined with the earth, or merely me- chanically mixed with it, and retained in combi- nation by cohesive attraction. In the former state it is of no use to vegetables, in the latter it is essentially necessary for their support. If the soil be not sufficiently retentive, the plant is starved, for nothing can be taken up from the earth which is insoluble ; and, as we shall show afterwards, water itself is a principal part of the food of plants. If the soil be too stiff and re- tentive, the water remains upon its surface, and does not percolate to a sufficient depth to be ap- plied to the roots : and if the vegetable be of a succulent kind, the herbaceous part remaining constantly surrounded with moisture has its vege- LECT. V.] THE ROOT. SOILS. 217 tative powers weakened, and rots. This is par- ticularly the case in winter ; for, as the vital energy of the plant is then much lowered by cold, a dis- ease of the vegetable takes place, similar to what happens in a leucophlegmatic state of the animal body, from which the plant rarely recovers. The most efficient soil, as far as water is concerned, is that which contains a due mixture of carbonate of lime, sand, and pulverized clay, with some ve- getable or animal matters ; and in which the ma- terials are so mingled as to remain loose and per- meable to the air. This soil is calculated not only to retain the water in proper quantity ; but also to absorb it from the atmosphere, which is one great source of the supply that vegetables require : for water, as has been already remarked, is requisite for rendering the other matters in soils sufficiently soluble to be taken up by the roots of plants. All the earths are more or less soluble in water : thus lime is taken up readily in its pure state ; and also if the water contains much carbonic acid in solu- tion, when the lime is in the form of chalk, or a carbonate, in the proportion of about 82, p. 7. LECT. VI.] THE STEM. ITS DIRECTION. 253 that the stem extends horizontally, or on the surface of the ground, and sends out roots from below; as in lesser periwinkle, Vinca minor (h), and Ground Ivy, Glechoma hede- racea. Linnaeus, Sir J. E. Smith, and some other authors, describe another variety of the procumbent stem under the term trailing or sarmentose (sarmentosus *) : but the sarment is, properly speaking, not a stem, but merely a runner which performs the same functions as the underground runners, to which pendu- lous tubers are attached. It is, in fact, a vas- cular cord, intended to place the lateral pro- geny of the plant at a convenient distance from the parent, and to convey nourishment to the offset until such time as it takes root and is capable of supporting itself. It is at first sent off from the neck or collet of the root ; but, afterwards, it gives off itself roots at the points where the offsets spring; and, as the manner in which this is effected va- ries, it is of different kinds ; but these shall be demonstrated when we investigate the ap- pendages of the stem. The swimming stem * " Sarmentosus, repens, subundus est." Phil Bot. 82. 9. " Sarmentosus, trailing. A creeping stem, barren of flowers, " thrown out from the roots for the purpose of increase, is " called sarmentum or flagellum, a runner.'* Smith's Intro* duction, &c. p. 120. 4 254 CONSERVATIVE ORGANS. [LECT. VI. (C. natans) has a much better claim than the sarmentose to be ranked as a variety of the procumbent stem, as it is a real stem, and lies floating on the surface of the water, throwing out radicles from the under side : the Floating Club-rush, Scirpus fluitans, is an example. The term natans for denoting this position of the stems of aquatics is of importance in a systematic point of view, and is used in con- tradistinction to sunk (demersus) ; which im- plies that the stem lies below the surface, as in common Horn wort, Ceratophyllum demer- sum. Besides the diversity which prevails in the position or direction of the stems, we perceive, in glancing the eye over the vegetable kingdom, that they present much variety, also, with respect to general form. Thus, some are simple or undi- vided, while others are divided and subdivided into very minute ramifications. 1. The UNDIVIDED or SIMPLE stem (C. sim- plex) consists of one piece only, without any branches bearing leaves; although the flower- stalk may be divided ; as exemplified in knotty- rooted Figwort, Scrophularia nodosa. When, however, there is no division even of the flower- stalk, but the whole stem is one rod or column from the base to the summit, and is terminated by a single flower, or a simple spike, it is LECT. VI.] THE STEM. ITS DIVISIONS. 255 then termed very simple (simplidssimus) ; as in Bistort, Polygonum bistort a, the Date Palm, Phoenix tkctylifera, &c. 2. The DIVIDED stem, owing to the parts into which it divides being termed branches (rami), is denominated branched (ramosus *) ; and the manner in which this is effected produces diver- sities in the general aspect of plants of great importance to the systematic Botanist. It af- fords, also, a kind of physiognomical character to each species, by which it is immediately re- cognizable to the eye of observation ; and the perceiving this character, and retaining a clear impression of the perception in the memory, ought to be strictly attended to by the student. It af- fords him a sure guide to the more particular ex- amination of vegetable nature; associating the minute characteristics of each species so closely with its general features, as to call those up ra- pidly in the memory, whenever these present them- selves either to the eye or the imagination ; and, in Botany, you will soon be convinced how es- sential it is to fix, in as practical a manner as possible, every impression which the examination of plants can make on the mind. The greater masters of the old schools of painting were fully aware of the importance of studying the physio- * " Ramosus est ramis lateralibus instructus." Phil. Bot. 82, 21 . 256 CONSERVATIVE ORGANS. [LECT. VI. gnomy of plants, especially of trees ; and they carefully transferred it to their canvass. Thus you can recognize in their landscapes, as in na- ture, a Poplar, a Willow, the Cedar, an Elm, an Oak, or any other tree ; whereas, the general term trees is sufficient to describe the foliage, which crowds in confused masses the pictures of inferior artists. The branches are evident divisions of the principal trunk ; yet, in the majority of in- stances, the stem can be traced, as it rises amidst these divisions, from the base to the apex ; but as it is, also, in some plants entirely lost, Bo- tanists employ distinct terms to designate these opposite states : thus, continuus is used for the former, and for the latter decompositus. The branching of a stem admits of several varieties, each of which requires to be noticed. It is said to be a. Slightly branched (subramosus) , when the number of divisions are comparatively few. b. Much branched (ramosissimus) , when not only the greater divisions are numerous, but these are again divided and subdivided with- out order * ; as in the Elm, Ulmus campestris ; the Gooseberry-bush, Ribes Grossularia, &c. * " Ramosissimus ramis multis absque ordine gravidus." Pkit. Bot. 82. 2. LECT. VI.] THE STEM. ITS DIVISIONS. 257 Linnaeus and some others employ the term pro- liferous (prolifer) to denote a modification of the much-branched stem, in which the new branches shoot out only from the summits of the former ones *, as in the Scotch Fir, Pinus sylvestris : but, as Sir J. E. Smith remarks, the term is obsolete, and seldom used. c. " Abruptly branched (determinate ra- " mosus), when each branch, after terminating " in flowers, produces a number of fresh shoots, " in a circular order, from " just below the origin of " those flowers ; as in na- " ked- flowered Azalea, Aza- " lea nudiflora, and many " of the Cape Heaths f." d. Forked or dichotomous (dichotomus) (fig. i), when the divisions and subdivisions are, throughout, in alternate bifurcations ; as exemplified in Corn Salad, Valeriana locusta, Petty Spurge, Euphorbia peplus, and * when there are three flat sides A, and three rounded angles. 2. Four-cornered (tetragonus), (fig. #, 2), when there are four rounded angles. Sometimes the corners are bordered (fig. *, 2). 3. Five-cornered (pentagonus), (fig. t, 3), when there are five obtuse angles. In this instance, also, the corners are some- times bordered (fig. *, 5), and the in- tervals occasionally furrowed. 4. Six-cornered (hexagonus), fig. t, 4), when there are six obtuse angles. 5. Many-cor- nered (polygo- when the angles are numerous and obtuse: /3. Acute (acut& angulatus), when the angles are sharp, and the sides hollowed. t " Digonus, trigonus, tetragonus, pentagonus, polygonus, " precedents (anceps) species sunt." Phil. Bot. 82. 13. LRCT. VI.] THE STEM. FIGURE. 275 Of this variety of the angled stem there are, also, five sub-varieties : 1. Triangular (triangular Is), (fig. u, i). 2. Four-angled (quadrangularis) , (fig. u, 2) ; 3. Five-angled (quinquangularis) , (fig. U) 3). 4. Six-angled (soxangularis) , (fig. u, 4). 5. Many-angled (multangularis) *f~. In many-angled stems the angles are not always so acute as in the other sub-varieties. 7. Three-sided (triqueter*), (fig. *, i), when tliere are three flat sides forming acute angles ; as in Carex acuta. e. Angular (angulosus), implies that the angles are either very obscure, and the stem, con- sequently, can scarcely be placed in either of the foregoing arrangements; or that the angles are variable in number. There are two varieties of the angular stem, which have pe- culiar appellations ; viz. a. Furrowed (sulcatus) (fig. *, 5), when the stem is longitudinally in- dented with deep and rather broad hollows, like those of a fluted column || ; as in Common t " Triangularis, quadrangularis, quinquangularis, multangu- " tarts, ex numero angulorum prominentium." Phil. Bot. 82. 15. J " Triquetrus, latera tria plana obtinet." Ibid. 82. 14. || " Sulcatus sulcis excavatis latis profundis exaratus." Ibid. 82. 16. 'T 2 276 CONSERVATIVE ORGANS. [LECT. VI. Alexanders, Smyrnium olustratum : /3. Stri- ated (striatus), when it is longitudinally in- dented with fine parallel lines * ; as in Sorrel, Rutnex acetosa. d. Knotted (nodosus), (fig. w, 1), implying that the stem is divided, at intervals, by swellings or knots ; as in Knotty Cranes'-bill, Geranium nodosmn, &c. e. Articulated (articu- latus), that is, composed of joints, or apparently distinct pieces, united at their ends-j~. It is re- markable that Linnaeus, in his definition of this form of stem, should have confounded it with the kneed stem, as he must have been well aware that many articulated stems are per- fectly straight or erect. In some instances the articulated stems are knotted at the joints, but in others they are plain. Many of the Grasses afford instances of the former ; while the Cactus tribe sufficiently exemplify the latter. f. Kneed (geniculatus), (fig. w, 2), when an ar- * " Striatus lineis tenuissimis excavatis inscriptus." Phil. Bot. 82. 17. f " Articulatus, internodiis geniculatus : Piper." Phil. Bot. 82.31. LECT. VI.J THE STEM. FIGURE. 277 ticulated stem is more or less bent at each joint ; as in Floating Fox-tail Grass, Alopecurus geni- culatus; Three-flower Fescue Grass, Festuca triflora, &c. Such are the diversities which stems externally present to the eye. But these, although in their natural state they are sufficiently fixed to enable systematic Botanists to employ them as distinc- tive characteristics of species, which could not otherwise be distinguished * ; yet, are apt to vary, owing to peculiar circumstances connected with situation, climate, and soil. Thus we find a stem which, in its natural state, is round, assume a flattened appearance as if two or more stems or branches were united longitudinally side by side : an anomaly which is not uncommon in the Ash, several species of Daphne, the greater Nasturtium, Tropaeolum majus, and many other herbaceous plants ; and is denominated by authors f the clus- tered stem, caulisfasciculatus. It is not easy to account for this effect, since it is evidently not the simple lateral union of one or more stems or branches, a circumstance which might be pro- duced by pressure abrading the epidermis and cel- lular layer of the bark, and bringing the liber or * " Caulis in multis plantis ita essentiales praebet differentias, " ut eo demto, nulla certitude speciei." Phil. Bot. 276. f Smith's Introduction, p. 127. Keith's System of physiolo- gical Botany, vol. ii. p. 277. r3 278 CONSERVATIVE ORGANS. [LECT. VI. innermost layer of each of the united stems into contact ; for, were this the case, the pith of each of these conjoined stems would be still entire, and the aggregate would appear as distinct cylinders, according to the number of the stems ; whereas in the fasciculated stem there is one flat pith only, and the other parts of the stem are, in ar- rangement and number, the same as if the ano- maly did not exist. The idea, therefore, of the fasciculated stem arising from the pressure of one or more contiguous stems, and a natural graft being thus formed, as has been suggested by Mr. Keith, cannot be admitted. Several other ano- malies occur in the configuration of stems ; but as these are generally either the consequence of disease, or of some obstacle to the natural de- velopment in the individual plant occasioned by insects, the consideration of them may be de- ferred until we come to treat of the diseases in- cidental to the vegetable system. Independent of the diversities which we have already examined, STEMS have been properly distri- buted into distinct species. In this classification, Linnaeus and others who have copied after him, enumerate six species of stems ; but among these the footstalk of the leaf and the frond are impro- perly included ; for, as the former is decidedly a part of the leaf, and separates with it when it falls ; and the latter is a peculiarity connected with the foliage of distinct tribes of plants, the Palms, the LECT. VI.] THE STEM. CLASSIFICATION. 279 Algae, and the Ferns ; and is itself supported on a species of real stem (which I shall soon have occa- sion to describe to you), the stipe*, neither of them can with propriety be classed with stems. Mirbel-f- and Mr. Keith J omit the flower-stalk in their enu- meration of the species of the stem, and certainly with some propriety ; for, although it closely re- sembles the stem in its structure, yet, in an ele- mentary work, it is undoubtedly more intelligible to the student to describe it as a part only of it bearing the fructification, except when it proceeds immediately from the root. Contemplating the subject nearly in a similar point of view, I think we are authorized in distributing stems into Jive distinct species : the trunk, the stalk, t\\Q straw, the scape, and the stipe. 1. TRUNK (Truncus) is the appellation given to the stems of trees and shrubs . The trunk is * " Frons est dilatatio vegetabilis herbacea quae arete " cum cormi specie, qua sustentatur cohaeret, Palmis, Filici- " bus, Algisque propria." Willdenou, Spec. Plant, tomus v. Introd. p. xxi. f Siemens de Phys. vg. p. 99 and 622. J System of physiological Botany, vol. i. p. 43. Linnaeus uses the term trunk in the generic sense in which I have used the word stem ; and thus defines it : " TRUNCUS " folia et fructificationem profert ; species ejus sunt vii. Caidis, " Gulmus, Scapus, Pedunculus, Petiolus, Frans, Stipes; at " ttamus pars est." Phil. Bot. 82. T 4 280 CONSERVATIVE ORGANS. [LECT. VI. characterized by its ligneous structure, by being always perennial ; generally naked at the lower part; and divided and subdivided towards the summit into branches and twigs, bearing leaves and the fructification. It is thickest at the base, and gradually diminishes to the apex ; is covered with a thick, often dry and cracked bark ; and internally is composed of a central pith, sur- rounded by ligneous layers, the number of which varies according to the age of the plant. When it rises to a moderate height, like a simple column, before it divides into, or gives off branches, it is said to be arboreous, as exemplified in the Oak, the Elm, and the majority of trees ; but when the divisions or branches occur near to the soil, it is termed shrubby, fruticosus ; as in the Lilac, the Rosemary, and such-like. These terms, however, are always to be understood as having a relative signification only; for various circumstances, such as change of soil, climate, and the efforts of art, may metamorphose shrubs into trees, or reduce these again into shrubs; transitions which are by no means uncommon. 2. STALK (Caulis) is properly applicable to the stems of herbaceous plants only ; although the term is frequently used in a sense synonimous with trunk. The specific characteristics of the stalk are, that it is rarely ligneous, and lives but 4 LECT. VI.] THE STEM. CLASSIFICATION. 281 one or two years in the natural state of the plant. It may be divided and subdivided into branches, like the trunk ; and a greater number of these diversities which have been described belong to the stalk rather than to the trunk. 3. STRAW (Culmus) is a name strictly con- fined to the stems of the Grasses, Rushes, and the gramineous cerealiae *. It is either hollow, or partially filled with pith., and generally knotted, articulated, and kneed ; but very rarely branched. The knots are solid, confined to the articulations, and give origin to the leaves, which are sheathing at their base. It increases in length, tapering gradually to the apex, but not in diameter, is round, compressed, or triangular; and is fre- quently hairy; but, as Sir J. E. Smith properly observes, there is " no instance of such a scaly " culm as Linnaeus has figured in his Philosophia " Botanica, t. iv. f. 3f." 4. SCAPE (Scapus), strictly speaking> is a flower-stalk, as it bears the parts of fructification only, and is entirely devoid of leaves ; but, never- theless, as it proceeds immediately from the root, it may be properly classed as a stem ^. It is al- * " Culmus truncus proprius Gramini, elevat folia fructi- " ficationemque." Phil. Bot. 82. B. f Smith's Introduction, p. 128. f " Scapus truncus universalis elevans fructificationem nee " folia : Narcissus, Pyrola, Convallaria, Hyacinthus." Phil. Bot. 82. C. 282 CONSERVATIVE ORGANS. [LECT. VI. ways herbaceous; ,and is found either simple, and bearing one flower only, as in common Dan- delion, Leontodon taraxacum; or divided, and many-flowered, as in Cowslip, Primula veris. 5. STIPE (Stipes) is the term used to express the stem of Palms, Ferns, Fuci, and Fungi. It is generally cylindrical ; but sometimes swollen in the middle, and bears a frond, or the foliage which is peculiar to it, at its summit. In the Palms the stipe is in general a simple column bearing a spreading plume of leaves ; and of the same diameter at the summit as at the base, ex- cept in a very few genera *, which throw out branches. It is marked at regular distances by the cicatrices of the fallen leaves ; and increases in height by additions made to its summit by the development of a central gem, which throws out annually a new circle of leaves. In the Ferns it varies considerably in form, being round, chan- nelled, triangular, and quadrangular ; and is either devoid of vestiture, or is chaffy, scaly, spiny, or muricated, that is, covered with sharp hard tubercles : and the same diversities occur in the stipes of the Fuci. In the Fungi it is generally fleshy or leathery ; but it varies both in substance and form ; and although, in the greater number of instances, it is affixed to the centre of the cap, * Draccena, &c. LECT. Vil] THK STEM. 283 orpileus,as in the Mushroom, or nearly so ; yet in others its attachment is to the side of that body. Such are the peculiarities, connected with the exterior of stems, necessary to be noticed in this stage of our inquiries : in closing our examination of them, this question spontaneously presents it- self: why is there so great a diversity of form, vestiture, and mode of branching in the vege- table organs ? No satisfactory reply can be ad- vanced ; and, therefore, we are left to imagine that, as nature appears to delight in variety, the diversified and graceful forms and appearances of plants may be one source of pleasure prepared by Divine Benevolence for mortals. Be this as it may, man has not failed to render them accessaries to his comfort, and subservient to his necessities. In the cool shade of the branching arms of the Beech, or under the pillared canopy of the Banyan, he shuns the ardour of the meridian blaze ; with the shrubby and spiny Hawthorn, or the prickly Cac- tus, he encloses his fields ; while the pliant Osier is woven into baskets to transport their produce to the crowded city : the tall and straight Pine rises a mast, on which he spreads the sail that enables him to transport the riches of distant climes to his native shores ; and the incurved ribs of the venerable Oak, launched into the main, float, the protectors of his maritime rights, and the bul- warks pf his national independence. 284 CONSERVATIVE ORGANS. JJLECT. VII. LECTURE VII. SUBSTANCE AND ORGANIZATION OF THE STEM AND BRANCHES : ANATOMICAL DEMONSTRATION OF THE COMPONENT PARTS OF THESE ORGANS THEIR FORMATION, INCREASE, AND REPRODUCTION. THE examination of the external aspects of stems and branches, which we have just concluded, will readily enable you to form an estimate of the im- portance of the correct knowledge of terminology to the systematic Botanist ; for, without a definition and name for each diversity of feature, which the vegetable organs present to the sight or the touch, it would be impossible for him to fix these specific distinctions, by which alone the different members of the same family of plants can be readily recognised. This diversity, as you have seen, is very considerable in the external charac- ter of the stem and branches ; but when we ex- tend our examination to their interior structure, or organization, we find it confined within very narrow limits : nor will this excite our astonish- ment when we reflect, that nearly the same end is to be accomplished by these organs, whatever may be the habits or the physiognomy of the plant. LECT. VII.] THE STEM. 285 It is now my object to direct your attention to these differences, and to point out to you the cir- cumstances in which they consist. If we take a number of stems and branches of different kinds of plants, and cut, or break, or tear them transversely and longitudinally; we shall find that some of them are easily divided, whilst others resist almost all our efforts ; that some are moist, succulent, and fleshy; others fibrous, spongy, and dry ; and others again formed of both succulent and dry parts ; the latter pos- sessing very different degrees of compactness, in- duration, and tenacity. But, however considerable these differences may appear, the more we investi- gate the subject the more we shall be convinced, that, as far as respects substance, the stems and branches of the whole of the vegetable kingdom may be arranged under two classes, the Woody and the Herbaceous. A. WOODY stems (Caules lignosi) are those which contain a very large proportion of ligneous fibre ; or in which wood forms comparatively the greater part of their bulk. It is found either in threads or bundles which run longitudinally from the base to the summit of the stem ; or is united in one mass, formed of concentric circles, which cohere and compress each other nearly from the centre to the surface of the stem : the tribe of 286 CONSERVATIVE ORGANS. [LECT. VII. Palms affords examples of the first description of ligneous stems, and all other trees of the second. As far as respects substance, however, ligneous stems are not arranged according to the diversity of structure I have just hinted at ; but according to the degree of cohesion which binds together their ligneous fibres. Botanists have, therefore, divided woody stems into two genera, the solid and the fibrous. 1 . The solid woody stem (C. solidus) is that in which the cohesion is uniform, and the wood, consequently, compact and indurated; sometimes to a degree on which the knife will make scarcely any impression. 2. The fibrous (C.fibrosus) is that in which, the cohesion being irregular, the wood is con- stituted of fibrous bundles ; which, although adhering to each other, can be easily separated, either by tearing or by maceration. Ligneous stems of every description are peren- nial. B. HERBACEOUS stems (Caules herbacei) are those which contain a small proportion of woody matter ; but are composed chiefly of cellular sub- stance, and consequently can be easily cut or divided. There are three distinct kinds of her- baceous stems: the fleshy^ the spongy, and the hollow. LECT. VII.] SUBSTANCE OF STEMS, &C. 287 1. The fleshy stem (C. carnosus) is that in which the cellular substance, of which it is composed, is turgid with fluids; and when cut, presents a smooth, uniform moist surface, similar to that of a young Cucumber when transversely sliced. The common House-leek, Sempervivum tectorum ; and most of the sea- weeds, Algse, afford examples of the fleshy stem. 2. The spongy stem (C. spongiosus) is com- posed of a compressible, elastic, cellular sub- stance, contained within an epidermis ; either dry, the cells being filled with air only; or moist, the cells being partially filled with fluid, as in a sponge. The Mushroom tribe, Fungi ; Indian Corn, Zea mays ; and Great Cat's-tail, Typha latifolia, have spongy stems. 3. The hollow stem (C.Jistulosus) is a suc- culent or fleshy hollow cylinder, surrounded by a circle of vessels and ligneous threads, imme- diately under the epidermis ; and generally lined with a dry white spongy layer of pith. There are two varieties of the hollow stem. a. The uninterrupted (sine septis transver- sisj; when the hollow, or cavity, extends from the base to the apex of the stem, as in the genus Coreopsis (Coreopsis). b. The interrupted (septis transversis inter- tinctus) ; when either the longitudinal cavity is divided, at regular distances, by parti- 288 CONSERVATIVE ORGANS. [LECT. vn. tions or diaphragms (fig. a, 1) ; or the stem is knotted, and apparently made up of separate hollow por- tions, shut at each ex- tremity, and united at the knots. The different spe- cies of Horse-tail, Equi- setum ; Manured Reed Arundo donax; Common Water Drop wort, CEnathe Jistulosa ; the Castor oil plant, Ri- cinus officinalis ; and almost all the Grasses, and the Cerealia, afford examples of this variety (fig. a, 2). Such is the diversity of substance of sterns and branches ; we have now to examine the ana- tomical construction of these organs. To the stu- dent, this inquiry is not only important, as dis- playing the structure of the stem itself, and thereby enabling him to comprehend more per- fectly its- functions and mode of growth ; but, as all the parts concerned in the vegetable functions, as far as the preservation of the individual is con- cerned, are seen in the stem ; by a correct know- ledge of its organization he becomes acquainted with that also of the root, the leaves, and even, in some respects, of the flowers and the fruit. In this investigation it is essential to adopt some ar- rangement; and although none, which has yet been attempted, is free from just causes of cri- LECT. VII.] ANATOMY OP STEMS. 289 ticism, and probably none can be devised alto- gether unobjectionable; yet, if by any the examina- tion can be facilitated, the attempt to form one, as nearly perfect as the nature of the subject will admit of, is at least praiseworthy. The only arrangements which I shall notice, are those of Desfontaines and Mr. Keith. Des- fontaines' method is founded on the fact, that there are two grand divisions of plants, the Mo- nocotyledonous * and Dicotyledonous -f~ ; each of which displays a distinct and specific mode in the distribution of the parts of the stem. In the in- dividuals belonging to the first, the stem consists of bundles of woody fibres and vessels, inter- spersed throughout a cellular substance, and de- creasing in solidity from the circumference to the centre ; in those of the second it is composed of concentric and divergent woody layers, decreas- ing in solidity in the opposite ratio, or from the centre to the circumference, and containing a pith in a central canal J. But, as Mr. Keith has justly observed, this arrangement does not ex- haust the subject : as, independent of the Poly- cotyledonous plants which may be classed with the Dicotyledonous, there is no account taken of * Plants, the seeds of which have one lobe only, f Plants, the seeds of which have two Lobes, t Vide Mem. de V Instil. Nat. tome i. VOL. I. U 290 CONSERVATIVE ORGANS. [LECT. VII. those tribes which are altogether destitute of seed- lobes ; the stems of which, in their organization, exhibit neither the woody bundles of the one, nor the layers of the other. Mr. Keith, in order to ob- viate this objection, has arranged stems, in re- ference to their internal structure, into three classes : 1 . The caudex an homogeneous mass ; 2. The caudex an heterogeneous mass; and, 3. The mass consisting of bark, wood, and pith*. This arrangement is undoubtedly less exceptionable and more comprehensive than that of Desfon- taines ; but Mr. Keith has erred in arranging, in his second division, the herbaceous stems of dico- tyledonous and polycotyledonous plants ; for, al- though these have no concentrical circles of formed wood, such as characterize the ligneous stems of those tribes, yet the distribution of the ligneous matter and of the pith coincides sufficiently with that of these parts in the real woody stems to allow them to be classed together ; whilst it has little or no affinity with that of the ligneous bundles in the monocotyledonous stems with which they are classed in Mr. Keith's arrangement. A more perfect classification, perhaps, may be formed, by adopting the first division of Mr. Keith's arrangement and adding to it those of Desfon- taines ; as, besides embracing the whole of the subject, it leads us first to the examination of * System of physiological Botany, vol. i. p. 287 & seq. LECT. VII.] ANATOMY OF STEMS. 291 those stems which are the simplest, because com- posed of the fewest parts ; and progressively pre- pares us to understand the organization of those of a more compound nature. I shall, therefore, attempt such an arrangement, and class stems, in reference to their anatomical structure, into the three following divisions : i. Stems which display, internally, an appa- rently homogeneous mass. ii. Stems which proceed from monocotyledonous seeds. iii. Stems which proceed from dicotyledonous and polycotyledonous seeds. i. Stems which display, internally, an apparently homogeneous mass, when examined by the unassisted eye, seem to consist simply of an epidermis enclos- ing a parenchyma, composed either of cellular substance, of very different degrees of succulency, sponginess, dryness, and density ; or of inter- woven fibres, forming a leathery, or a felt-like tex- ture, or one not a little resembling that of washed animal muscle which has been macerated in spi- rits. When examined, however, by the aid of a good microscope, these different appearances of the internal mass are all found to consist of cel- lular substance, with vessels running through it, and anastomosing in a variety of directions. Mirbel erroneously asserts that this description of 292 CONSERVATIVE ORGANS. [LECT. VII. stems is destitute of vessels * ; for their presence, at least in the stipes of the Mushroom tribe (Fungi), and the stalks of many of the Algse, is demonstrable by placing a small transverse slice of any of them under a powerful microscope. They are not, indeed, so readily distinguished in a longitudinal slice, a circumstance which I am inclined to ascribe to the transparency of their coats confounding them with the cellular sub- stance in which they are imbedded ; and which consists of continuous, oblong cells, the membrane forming the sides of which is of very different degrees of thickness ; but, nevertheless, they may be made out by any one accustomed to the use of the microscope. I have never been able to satisfy myself of what description these vessels are, although it is evident, as Mirbel remarks, that they are neither the spiral, nor the annular ; but I suspect them to be the moniliform. The epidermis, which cannot be separated from the cellular mass it covers, is pierced with imperceptible pores, but, according to Mirbel, it has no miliary glands. In the different tribes of plants which afford examples of this description of stems, the reproductive or- gans are either altogether deficient, or so ob- scure as to have eluded the researches of phyto- * Vide Element de Phys. veget. l ere Partie, p. 37 ; and Journ. de Botanique, torn. iii. p. 36* LECT. VII.] ANATOMY OP STEMS. 293 legists; and thence they have been regarded as agamic. Many of them have no stem ; but among those which possess it, in some it is solid, or rather entire, in others hollow; and in the lat- ter case, the cavity is often partially lined with a very lax, dry, cellular web *. A conspicuous root is rare; and, when it exists, consists of a few small radical fibres only, composed of a single thread, covered with a cribriform epider- mis. Scarcely any facts are yet known respect- ing the development and growth of this descrip- tion of stem. ii. Stems which proceed from monocotyledonous seeds are more complex in their structure than the preceding. They are composed of two dis- tinct parts, ligneous and cellular, which assuming a determinate character, enable these stems to be readily distinguished, even by the naked eye. They comprehend both solid and tubular, or entire and hollow stems ; and as there is some difference in the arrangement of the parts in these varieties, we shall examine them separately. * See Plate 5, fig. 1, which represents a longitudinal sec- tion of the stem of Tall Agaric, Agaricus procerus ; a. the stem ; b. its lax cellular lining, or pith, which in this example nearly divides the hollow of the stem into two unequal cavi- ties ; c. a fragment of the hat or pileus, to show the simplicity of its union with the stem. u3 294 CONSERVATIVE ORGANS. [LECT. VII. a. If a solid rnonocotyledonous stern, that of a Palm for example, be cut, either longitudinally or transversely, it is seen to consist of an epider- mis enclosing ligneous bundles or cords, more or less symmetrically distributed in a parenchyma or medullary substance. If the section be longi- tudinal, these ligneous cords are observed to run longitudinally, and extend from the base to the apex of the stem, sometimes in straight lines ; but occasionally assuming a zigzag direction, so as to touch each other at different distances : closer together and firmer towards the circum- ference of the stem, and more apart and softer as they approach its centre *. If the section be transverse, the divided extremities of the ligne- * Vide Plate 5, fig. 2, 3, 4. Fig. 2 represents the section of a transverse cutting of a Palm, the Ptychosperma gracilis (copied from a plate of Mirbel) ; a. the ligneous bundles closer together, more indurated, and in greater numbers than in the space marked b, in which also, however, they are less distant and more compact in that portion which is farther from the apex of the section, than in that which is nearer to the centre of the stem. Fig. 3. A. 'a transverse slice of the scape of the Tiger- flower, Tigridia pavonia, slightly magnified ; the ends of the ligneous bundles being shown by the white dots, which in the natural stem appear on a green ground. Fig. 4-. A longitudinal section of the scape of Great Reed- mace, Typha latifolia, showing the ligneous bundles, which in this instance do not inosculate. LECT. VII.] ANATOMY OF STEMS. 295 ous bundles appear like spots, which are in some instances of a dark colour, and in others white, dispersed over a white or a green ground, in the order just described. The epidermis adheres closely to the parenchyma beneath it ; and in some plants of this class, the greater density of the cellular substance at the circumference gives the appearance of a bark, which is never, how- ever, present in this description of stem. Such is the general character, and the distribution of the parts, in what may be termed the lig- neous solid monocotyledonous stems* ; but when they have more of an herbaceous character, such, for example, as the scape of Great yellow Gar- lick, Allium Moly, there are no indurated lig- neous cords ; but the vessels run in the midst of longitudinal layers of condensed cellular matter, and in a transverse section appear as white dots forming a circle round the central cells, which are generally much larger than those of the circumfe- rence, and assume in some degree the aspect of a pith *}- ; so that in the longitudinal section, the * The student, who cannot procure the stem of a Palm, may attain an excellent idea of the internal structure of the solid monocotyledonous stem in the flower-stalk of the Tigri- dia pavonia, which is now very common in the stoves of the London florists and nurserymen; or in that of the Typha latifolia, not unfrequent in ditches, flowering in July. f Vide Plate 5, fig. 5. A. a transverse slice of the scape of Allium Moly, nearly of the natural size. u4 296 CONSERVATIVE ORGANS. [LECT. VII. diameter of the stem appears divided by two seem- ingly solid cords, into three nearly equal compart- ments. Mirbel, in treating 1 of this description of steins, says, " their pith, instead of being en- " closed in a canal, in the centre of the stem, ex- " tends almost to the circumference * ;" but there are exceptions to this remark, as, for instance, in the Rush, Juncus, which has a perfect pith, sur- rounded, however, by a cellular tube, in the sub- stance of which the vascular cords characteristic of the monocotyledonous stems are perfectly ap- parent, arranged in beautiful order; and dis- tinguish it from an herbaceous dicotyledonous stem, the vessels of which, as I shall afterwards demonstrate to you, are arranged in a very dif- ferent manner. Such are the appearances which, to the naked eye, or to the eye aided by a common lens, the solid monocotyledonous stems present. Under the microscope, we perceive that each ligneous cord is composed of very narrow oblong cells, and of vessels which are either spiral, or annular, or porous, those in the centre being always spiral ; that, in the cellular substance of the more solid stems, the cells are chiefly oblong, whilst in that v " Et leur moelle, au lieu d'etre resseree dans un canal, " au centre de la tige, s'etend presque jusqu'a la circonference. 1 ' Element de Phys. veg. Partie 1. p. 117. LECT. VII.] ANATOMY OF STEMS. 297 of the herbaceous they form irregular hexagons, except towards the circumference, and in the immediate vicinity of the vascular cords*; and that the membrane forming them is perforated with minute pores, surrounded by a glandular border. In the common Rush, Juncus conglo- meratus, and some other monocotyledons, the cells of the pith are of a very curious structure : appearing, in a transverse section, like minute heptagonal wheels or circles divided by rays passing from the centre ; and these are evidently filled with air *f-. In common MareVtail, Hip- * Vide Plate 5. Fig. 3, B. represents a magnified section of fig. 3, A. to demonstrate the greater density of the cellular substance towards the circumference. Fig. 5, B. a highly magnified section of fig. 5, A. a. the cellular substance more condensed close to the epidermis than in the heart of the stem ; b. the vascular bundles. Fig. 5, C. a highly magnified longitudinal section of the semi- diameter of the slice A. which has the appearance of fig. 5, d. to the naked eye: a. the cellular matter, close to the epidermis, appearing as oblong cells ; I. one of the seemingly solid cords, which form the circle of vascular bundles, con- sisting of two spiral vessels, surrounded by oblong cells; e. the cellular substance which constitutes the mass of the stem. In this view the membrane forming the walls of the cells is seen studded with points, which, under the microscope, appear to be either amylaceous granules or circular apertures surrounded by a border. Mirbel supposes they are pores, and that the border is glandular. f Vide Plate 5, fig. 6, which represents a magnified section of a transverse slice of the common Rush : a. the pith com- 298 CONSERVATIVE ORGANS. [LECT. VII. puris vulgaris, the air-cells, which are on the out- side of a central column, composed of vascular and ligneous bundles, are divided from each other by smaller cells, which are filled with aqueous fluid : whilst, at every whorl of leaves, bundles of vessels are given off from the central column, and these are surrounded by condensed cellular matter, which forms a kind of diaphragm ; by the repetition of which the whole stem is divided into compartments at regular intervals *. posed of circular or heptagonal cells, divided by septa passing from the centre of each, like rays; b. the cylinder which surrounds the pith ; composed of green parenchymatous mat- ter, in which bundles of spiral vessels (c. d.) run, surrounded by condensed cellular matter ; e. departments consisting of air- cells dividing the vascular bundles ; f. bundles, apparently of entire vessels, forming the striae on the surface of the stem ; g. lacunae or open spaces between the pith and its enclosing tube. If I might be permitted to hazard an opinion regarding the use of this organization, it would be this : that, as the Rush has no leaves, the green parenchymatous tube is intended to perform a function analogous to that of leaves, these organs consisting chiefly of a similar substance. * Plate 5, fig. 7, a section of a transverse slice of the stem of common Mare's-tail, Hippuris vtdgaris, an aquatic plant : a. the exterior, or cellular part (enclosed merely by the epi- dermis), consisting of large air-cells, each surrounded by smaller cells, which are generally found turgid with aqueous fluid ; b. the central column, which consists chiefly of spiral vessels, one of which is drawn out at c. All aquatic plants contain very large air-cells ; which are most abundant in their stems, if their leaves be few or comparatively small, or the greater number of them is above the surface of the water ; and LECT. VII.] ANATOMY OF STEMS. 299 It is necessary to notice, particularly, the structure of one variety of the solid, herbaceous, monocotyledonous stems ; as it constitutes the link betwixt the solid and the fistular monoco- tyledons. It is articulated, giving off leaves at the joints only ; and these are sheathing or em- brace the stem. On dissection, we find the vas- cular bundles, which can scarcely be called lig- neous, enclosed in a cellular parenchyma, and running in straight lines between the joints ; but at these they inosculate in arches, and send off horizontal branches, some of which terminate in the leaves and others in the lateral shoots that originate at their basis. This structure is beau- tifully displayed in the stem of Spiderwort, Trade- scantia Virginica, in which the circles forming the annular vessels being nearly opaque, the course of the vascular bundles is readily traced ; and the manner in which the vessels are given off, and inosculate, distinctly seen by the aid of the microscope *. in the leaves, if these be large or mostly immersed, as for in- stance the Common Reed-mace. * Vide fig. 9 and 10, Plate 5. Fig. 9, a. a. a. a. a. a. the vascular cords, as seen by means of a good lens by reflected light, forming arches at the joint ; b. 6. the base of the leaf, with the vessels which supply it proceeding nearly from the centre of the joint ; d. the fragment of a lateral shoot, the vascular bundles connected with which are seen originating from the same spot as those of the leaf. Fig. 10, a Small por- tion of the joint highly magnified, to display the structure of 300 CONSERVATIVE ORGANS. [LECT. VII. b. The hollow or fistular monocotyledonous stems are composed of distinct portions, united by knots ; at each of which the cavity is divided by a diaphragm : or, rather, each portion may be re- garded as a distinct individual, which takes its origin from one knot, and terminates in another, out of which again a new individual arises, and so on in succession, as I shall more particularly de- scribe in explaining the mode of growth of these stems. The general structure of this description of stems is best exemplified in the Grasses. Thus, in Wheat we perceive the upper articulation rising within the knot, in which the lower has termi- nated ; with the leaf which infolds it crowning the embracing knot*. The organization of this variety of the monocotyledonous stem cannot be readily distinguished without the aid of the mi- croscope. It is seen, in a longitudinal section, to consist of several layers of narrow oblong cells, which constitute its exterior and more solid part ; and of an interior more open cellular substance, the vascular cords and the inosculation of the vessels, a. The bed or sheath in which the vessels run ; b. the vessels themselves apparently composed of separate rings, held together by small spicular bodies, which appear as lines on the surface of the vessels, within the transparent sheaths ; when detached, they appear like acicular crystals. The cells are studded with amy- laceous granules. * Vide Plate 5, fig. 1 1. a. the knot of the straw bearing a fragment of its leaf; b. the new articulation rising within the knot. LECT. VII.] ANATOMY OP STEMS. 301 enclosing vascular, ligneous cords, composed of oblong cells like those on the circumference, sur- rounding spiral and annular vessels. In the trans- verse section the divided extremities of these cords appear as clustered vascular spots in the cellular substance *." The bark, if the surface of the stem can be so named, of the more solid monocotyledons is formed of the footstalks of the leaves; but the real epidermis of both the ligneous and herbaceous stems of this tribe, is always, as has been already stated, so closely applied to the part which it covers, as to be inseparable from it by any means. Owing to this circumstance it appears of a cellular structure, and its cha- racter is regulated by the nature of the parts it immediately encloses. In those plants, in which it can be readily examined, it displays, under the microscope, a regular series of organic exhaling pores, each apparently surrounded by a glandu- * Vide Plate 5, fig. 8. A. B. C. A. represents the transverse slice of a stem of Wheat, as seen through a good lens ; B. a longitudinal section of the slice highly magnified ; a. the outer cellular substance, composed of oblong cells and entire vessels ; b. the interior cellular lining of the cylinder of the stem ; c. a vascular bundle, consisting of a spiral vessel surrounded by oblong cells ; C. a transverse section of the slice ; a. the tu- bular cells immediately under the epidermis ; b. the common cellular substance ; c. the vascular bundles. 3 302 CONSERVATIVE ORGANS. |~LECT. VII. lar border ; as is beautifully demonstrated in the culm of Wheat * : but in some plants, as, for in- stance, the Common Rush, these apertures are perceptible in the furrows only between the striae, the elevations being apparently free from any exhaling pores-f~. In some of the herbaceous dico- tyledons, silex is found deposited in, or rather immediately undeiv the epidermis. Monocotyledonous stems, those even of the largest diameter, display no medullary rays, such, as I shall soon have occasion to demonstrate, as characterize the dicotyledonous ; nor do such ap- pear to be necessary, owing to the extensive dis- tribution of the cellular matter throughout the substance of these stems. The woody bundles, however, become indurated by age, and the more external being enlarged by the deposition of new ligneous matter, they at length occasionally touch each other, and form a circle of continuous wood ; but the interior bundles never attain this state, and are always sufficient to distinguish the stem as a monocotyledon. Monocotyledonous stems increase in length or height ; but, with very few exceptions, not in diameter. As I have had no opportunity of tracing the manner in which this is effected, in the ligneous * Vide Plate 5, fig. 12. t Vide Plate 5, fig. 13. a. the appearance of the cuticle over the ridges ; b. its cribriform appearance in the furrows. LECT. VII.] ANATOMY OF STEMS. 303 monocotyledons, I shall follow the description of Mirbel, in explaining to you the growth of a Palm. When the young plant, rising from the seed, ap- pears at the surface of the ground, the leaves, originally folded up and sheathed within each other, separate themselves, increase in number, and form a sheaf-like group. Those on the cir- cumference now spread out, perform their func- tions, and are detached ; but their bases re- maining, form a solid or ligneous ring, which is the origin of the stem. Within this circle the sheaf of leaves rises vertically ; owing, perhaps, to the resistance at the circumference ; and the exterior ones having spread out on every side in the same manner as the former, drop after a time, also, and leave their bases to form another circle ; within which the sheaf still rising, again spreads out another range of leaves ; and in this way the stem is gradually formed by the evolution of the terminal leaf-bud, and the induration of the footstalks of the fallen leaves. The whole stem displays the cicatrices of the successive circles of detached leaves, and these becoming hardened by their exposure to the air, and the ligneous bundles within them being older, in a direct ratio as they are nearer to the surface (the development of parts always taking place in the centre), the substance of the stem is necessarily softer within, and harder as it approaches the circumference. 304 CONSERVATIVE ORGANS. [LECT. VII. Owing to the mode of growth, also, which has just been described, the stem is always naked, columnar, and terminated with leaves and fruc- tification in the form of a magnificent crown, as exemplified in the Palms. The stipe, therefore, or this kind of monocotyledonous stem, may be regarded as a fasces of ligneous vascular rods imbedded in cellular substance, and terminating in leaves * : and its vitality being, in a great de- gree, dependent on the herbaceous part, if the central bud, or cabbage-}-, as it is commonly called, be cut off, the whole plant immediately dies. In tropical climates, some kinds of Ferns rise with a stipe resembling that of the Palms ; but this appears to be, according to Mirbel, " a * The height to which some Palms arise, without increasing in diameter, is truly astonishing. Thus the Ptychosperma gracilis rises more than sixty feet above the surface of the ground, with a stem not four inches in thickness. The eleva- tion of the Areca oleracea is often not less than one hundred and eighty feet ; and " although," says Mirbel, " its diameter " is greater than that of the Ptychosperma, yet, it is certain " that it never increases in thickness." Elem. de Phys. veg. t. i. p. 12. Its stem is, nevertheless, thicker in the middle than either at the base or the summit, when the Palm has attained to a certain age ; which is justly ascribed to its vegeta- tive powers being more vigorous at the middle period of its ex- istence. f The terminal bud of the Areca oleracea is boiled and eaten as a delicacy, under the name of Cabbage ; and the plant is called the Cabbage Palm. LECT. VII.] ANATOMY OF STEMS. 305 " simple fasces of petioles, or leaf-stalks * ;" al- though circumstances occasion these to unite in the interior of the stipe, and form masses of com- pact wood. This variety of stipe does not increase in diameter. The Aloes, the Yucas, and the Dracaena differ in their mode of growth from the Palms, inas- much as they give off branches and increase in the diameter of their stems. From the observa- tions of M. Aubert du Petit-Thouars -f- , who traced the mode of branching, and the consequent in- crease of diameter in the stem of the Dracaena, it appears that the branch originates in a small protuberance under the epidermis, which it soon ruptures ; and extending, first unfolds some scales and then leaves, the result of the successive de- velopment of which is a cylindrical branch com- posed of ligneous cords, similar, in every respect, to the parent stem, and seated upon it like a graft. The ligneous cords, at the point of their junction with the adult stem, spread out at first like rays, and then extend, in an opposite direction to the growth of the branch ; the exterior descending in straight lines towards the earth, whilst the others, after ascending a little at first, soon, also, bend down and take the same course, which the whole * ElSmens de Phys. vSget. 1. p. 121 . f Essais sur la Vegetation, &c. Paris, 1809, p. 1. VOL. I. X 306 CONSERVATIVE ORGANS. [LECT. VII. preserve until they are lost in a mucilaginous sub- stance, secreted under the detached epidermis. A layer of new ligneous cords is thus applied over those of the old stem, and the development of other branches still adding fresh layers, the stem is gradually increased in diameter. M. Aubert du Petit-Thouars thinks that, for the formation of branches in these plants, a vital point (un point vital) exists at the axilla of each leaf; but re- mains latent and inactive, unless peculiar circum- stances occur to call it into activity ; and this, he conceives, constitutes the difference between these gems and the buds which appear in the axillae of the leaves in the great majority of dicotyledonous plants. I shall, however, soon have an oppor- tunity of demonstrating to you that the same circumstance occurs in dicotyledonous stems ; on which buds sometimes appear, that have existed in a latent state for many years; and can be traced back to their origin in the change of or- ganization, occasioned by the frustrated effort to develop them, in the successive layers of wood which have annually added to the diameter of the stems, on which they are ultimately developed. The development and growth of the herba- ceous, solid, monocotyledonous stem is nearly the same as that of the ligneous, except that the parts are more rapidly evolved. If we trace the growth of the stem of the White Lily, for example, LECT. VII.] ANATOMY OF STEMS. 307 springing from a full-grown adult bulb, we per- ceive that it first appears on the surface of the ground like a large naked leaf-bud, which, when dissected and minutely examined, consists, not only of leaves overlapping each other, but also of the rudiments of the flower and fructification. Before this has risen much above the earth, the exterior leaves separate at their apex from the others and spread themselves out to the air and light, to form those secretions which are partly deposited in the portion of the stem below them, for the purpose of affording it firmness and solidity; whilst the more succulent portion above them extends, carrying with it the bud, until the leaves, next in succession, spread out and har- den it in its turn. As the stem continues to ex- tend and the leaves alternately to expand, it thus attains the summit of its height. In this pro- gress, the stem, as it advances, gradually loses a portion of its diameter ; and a transverse section , of it, near the summit, displays very few ligneous vascular cords, compared with those of one near the base ; and, consequently, contains less lig- neous matter. At the point, however, where the flower-stalks spring, it again thickens ; and the attachment of these closely resembles that of the branches of the Dracaena. In our demonstration of the anatomical struc- ture of the stems of this tribe of plants, we stated 308 CONSERVATIVE ORGANS. [LECT. VII. to you that those which are articulated, or knotted and nevertheless solid, form the link between the more common solid herbaceous, and the hollow or fistular monocotyledonous stems. As they display something in common with each, and yet differ from both, in some degree, in structure, so, also, they differ from both in the development of their parts. The leaves are given off at the joints, or knots only, and in general embrace the stem, or are sheathing to a certain extent: and when they protrude branches, these originate in gems formed at the joints, between the stem and the leaf, in a manner very closely resembling that in which young lateral bulbs are formed on the ex- terior of the laminated bulbs. These productions, indeed, may be almost regarded as lateral pro- geny, rather than real branches ; for they shoot out radical fibrils at the lower part of a knee which they often form with the main stem ; and, taking root in the earth, perpetuate the existence of the plant after the decay of the original stem. The parts of the stem below these lateral shoots increase, in a small degree, in diameter, from causes similar to those which have been already detailed as pro- ducing the increased diameter of the stem of the Dracsena; and it is in this respect chiefly that this stem differs in the manner of its growth from the culm or fistular monocotyledonous stem. This description will be better understood by refer- LECT. VII.] ANATOMY OP STEMS. 309 ing to the marginal cut, in which 1. represents J 2 a portion of the stem of Trad escan ti a Virginica, with two lateral shoots or branches : a. a young shoot, partially envelop- ed in its sheath b. and its origin fully displayed by the removal of the leaf in the axilla of which it is seated ; c. another shoot, but covered by the base of the leaf, the upper part of which is cut away at d. In 2, which represents the same portion of stem divided longitudinally, the manner in which the buds a. b. c. are given off is rendered more obvious. The white longitudinal lines are the vascular cords, which always appear white amidst the green parenchyma; whilst the trans- verse septa between each articulation, being formed by the branches of these vessels assum- ing an oblique direction, appear, also, of a white colour. The vascular cords, from which the ves- sels of the shoots originate, are easily distin- guished by their greater* size, and by passing di- rectly to the base of each shoot. The Culm, as has been described, consists of x3 310 CONSERVATIVE ORGANS. [LECT. VII. hollow articulations connected by knots; which in their growth appear to rise from within each other ; the new one always carrying with it the central bud, which in the last of the series termi- nates in the inflorescence. The leaves are sheath- ing and given off only at the knots, which are solid, or rather are filled up with a dense cellular substance, containing at first a saccharine nutri- tious juice, which Darwin asserts to be essential to the growth of the next joint, for it is absorbed or at least disappears as that is perfected. The vascular cords, which run in straight lines throughout the length of each joint, divide as they approach the knot, forming an inosculated plexus, and send off branches to the leaf and to the new articulation, to which the cellular substance, that forms the diaphragm interposed between the cavity of each articulation, actually belongs ; giving to the basis of the new joint the appearance as if it rose from a bulb. It was this appearance, as seen by the naked eye, which suggested the theory advanced by Dr. Darwin, that each joint of these stems is a distinct individual; and that the whole of a stem of any grass, Wheat for ex- ample, is a successive series of leaf-bulbs and leaf-buds produced in one year, and terminating in a flowering bulb. " At the first joint of the " stem of Wheat," says the Doctor, " on or LECT. VII.J ANATOMY OF STEMS. 31 L " within the surface of the earth, a leaf is pro- " duced ; from which rises the principal or central " bud, and around it many new buds, which " strike their roots into the soil. After this cen- " tral bud, and those around it, have risen six or " eight inches, a new leaf and a new leaf-bud " rises on each of them, producing a second joint of " the stem ; and lastly a flower-bud is generated " at the summit, which are all evidently distinct " vegetable beings, as there is a division across " the stem at each joint, which shows there is no " connexion of the pith or brain, or spinal mar- " row, between the lower and upper joints. That " a new bud thus constitutes each joint of the " stem of Wheat and other grasses is further " evinced ; first, by the existence of a leaf at each " joint without a lateral bud in its axilla, as oc- " curs in other vegetables : secondly, because for " the nourishment of this new leaf-bud a reservoir " of sweet juice is prepared in the new joint ; as " in the bulbs of many plants : and thirdly, be- " cause the lower leaf dies, and the sweet juice is " absorbed, as the upper leaf becomes vegete *." Setting aside the fanciful allusions to the brain and spinal marrow, there is much plausibility in this theory ; and it even appears supported by the ana- tomy of the knots of these stems, as far as that can * Phytologia, sect. ix. 3, 1. x4 312 CONSERVATIVE ORGANS, [LECT. vn. be perceived by the naked eye, or by the aid of an ordinary lens ; but when the microscope is em- ployed, the vessels can be traced from one joint to the other, passing through the spongy cellular diaphragm, and pursuing their course to the sum- mit of the stem. To illustrate this, I have ad- joined (see marginal cut) a plan of the vessels in the knot of the Wheat stem, as dis- played by the microscope, in a longitudinal slice of a straw. In this plan the white lines a. . represent the course of the vessels in the cylindrical part of the old joint running in a perpendicular direction, which they preserve up to the point b. b. where the leaf se- parates from the new articu- lation c. which is sheathed within it. At d. d. and between that point and b. b., branches are given off, which meet in the centre of the cellular matter of the knot, and again join the vessels a. a. at . b. b. where they enter the leaf f.f. in which they terminate. A little lower, however, opposite g. g. vessels are sent off, which at first are curved in- wards, but soon acquire a perpendicular direction and constitute the vessels of the new joint as seen LECT. VII.] A N ATOM Y OF STE M S . 313 at i. 2. The density of the cellular substance around the inosculating vessels constitutes the diaphragm, which is interposed between the arti- culations of this description of stem : and the new joint does not appear hollow at first, the cellular substance filling up its cavity as at k. for a short space above the knot. There can be, therefore, little doubt that the articulated culm is as much one individual as the stipe ; and the truth of this opinion is not shaken by our inability to explain the object of its articulated structure. But if it be allowable to hazard a conjecture, I would say, that as the cords of vessels run in straight lines, and are comparatively remote from the surface in these stems, the enlargement at the knots is partly intended to afford space for permitting branches from these vessels to be given off to supply the new joint which originates there: accord- ingly we see branches added to the vascular cords as they arrive at the knots; and these, after passing through it, approach each other and dis- appear in the vascular cords again pf the new joint ; the necessity for the branches no longer ex- isting. The original cords of the old stem ac- tually terminate in the leaf, which is the limit of the growth of the joint ; and in this respect gives some colour of truth to Darwin's theory of the individuality of the articulations. The cellular substance itself, in the knot, is probably intended 2 314 CONSERVATIVE ORGANS. [LECT. VII. partly for supporting these additions to the vessels, and partly for the development of the new bud in the axilla of the leaf ; which, notwithstanding the assertion of Dr. Darwin, occurs in the grasses in the same manner as in other plants, although in our climate it is not always evolved *. The saccharine juice, also, secreted in the knot, is more likely to be required as the first nutriment of the embryon bud (in some degree a new being -f-), which may be developed at this point, than to forward the growth and extension of the next joint ; which is already so constituted as to be able to make use of the nutriment with which it is supplied from the soil, through the medium of the roots. Such is the structure and the mode of growth of monocotyledonous stems. The positive features which chiefly characterize them in point of struc- ture, are the separate vascular ligneous cords, and intermixed cellular parenchyma; but they are distinguished more strikingly by negative qualities ; as, for example, those of having no proper bark, on * In tropical climates almost all the grasses give off branches from buds formed in the axilla of the leaves ; and even in this climate this occurs in Nodose Canary Grass, Phalaris Nodosa, and several other perennial grasses. f I have said, " in some degree," because, even allowing the bud to be lateral progeny, yet it is, strictly speaking, an extension only of the parent, and not a new being, in the sense in which this term is properly applied to seminal progeny. LBCT. VII.] ANATOMY OF STEMS. 315 liber, no alburnum, and no medullary rays; parts which, as I shall soon have occasion to demon- strate to you, belong, exclusively, to the dicoty- ledonous and polycotyledonous stems. I do not, at present, attempt to detail to you either the opinions of others, or my own conjectures, re- lative to the manner in which the various parts of these stems are formed; nor to trace their par- ticular functions ; as we shall enter fully into this part of our subject, when we arrive at the proper moment for taking into consideration the combined functions of the root, stem, and leaves. iii. Stems belonging to plants, which are pro- duced from dicotyledonous and polycotyledonous seeds, are in every respect alike in point of struc- ture ; and, therefore, for the sake of brevity, I shall describe both under the single appellation of Dicotyledonous stems. In treating of these, their natural division into woody and herbaceous imme- diately presents itself to our attention. A. WOODY DICOTYLEDONOUS STEMS consist of three distinct parts, the bark, the wood, and the pith. They are best exemplified in trees and shrubs ; but as the structure of each of these parts differs according to the age of the plant, it is re- quisite to examine them, both as they appear in the young plant or the shoot one year old only, and in the trunk and branches of older subjects. If the young shoot of any tree or shrub, the 316 CONSERVATIVE ORGANS. [LECT. VII. Horse Chesnut for example, be cut either trans- versely or longitudinally, the parts which have been enumerated, are rendered evident to the na- ked eye. If the section be transverse, it is seen to consist of a central spongy or cellular portion, which is the pith, enclosed within a ring of more solid consistence, which is the wood; and this, again, is environed by another circle of an in- termediate degree of firmness, which is the bark. If the section be longitudinal these parts are seen, in the same order (vide fig. 1, Plate 6), extending the whole length of the shoot ; the pith a. appears like a central column, guarded on every side by the wood b. y and this bounded by the bark c., which forms the exterior envelope of the whole. Running the eye, however, along the section, we perceive that the bark is not continuous ; but where the buds d. e. project, it appears as if re- flected over them, while the exterior fibres of the wood enter into their substance. At the bases of the leaf-stalks,/*, g. this is not the case ; for these appear as if seated upon the bark, connected with the shoot merely by the cords of vessels h. i. which penetrate the wood, and are apparently lost on the surface of the pith. At the summit of the shoot the pith appears to terminate, enclosed by the wood as if by an arch ; whilst the bark still covering the wood mingles with the substance of the leaves which form the terminal bud k. In LECT. VII.] ANATOMY OF STEMS. 317 pursuing our inquiries, we find that the bark is readily detached from the wood, b. and is sepa- rable into three layers ; that the wood is fibrous and more compact and harder within than in its exterior part ; and that the pith is evidently com- posed of cells, which, ,in the more succulent parts of the shoot, are filled with an aqueous fluid, and in the drier with air. Such is nearly the sum of the information we can obtain from an examina- tion with the naked eye ; to secure, therefore, an accurate knowledge of these parts, we must call in the aid of the microscope ; and, with its as- sistance, let us examine each of them in the order in which they present themselves in the shoot under our inspection, beginning with the bark. In taking this course, we shall confine our actual demonstrations to this shoot : pointing out, how- ever, and illustrating as far as possible, the va- rieties which have been remarked in each part, in different ligneous stems, both in their first year's growth, and in the after-periods of their ex- istence. a. The BARK. In the shoot we are now ex- amining, which has been cut in the autumn, the bark when separated from the wood is about the sixteenth part of an inch in thickness, and ap- pears, to the naked eye, to be composed of four very distinct parts. 1. A dry, leathery, fawn- coloured, semi-transparent, tough membrane, 318 CONSERVATIVE ORGANS. [LECT. VII. which is the cuticle ; 2. a cellular layer which adheres, although not very firmly, to the cuticle, and is named the cellular integument ; 3. a vas- cular layer ; and 4. a whitish layer, apparently of a fibrous texture, which is the inner bark ; and which, as we shall afterwards find, is of a more complicated structure than the other layers. We shall now view these parts separately under the microscope. 1. The Cuticle. Before demonstrating the structure of this part, it is necessary to remark, that I prefer the term cuticle to that of epider- mis, in reference to the exterior covering of stems and branches, in order to distinguish it from the thin unorganized pellicle which has already been described (page 93) under the name Epidermis, as one of the general components of the vege- table structure; and which is, in fact, the ex- terior part of the cuticle. The cuticle may be raised from the cellular integument by the point of a knife, and this is the best method to obtain it for minute examina- tion *. When thus separated and placed under the microscope, it appears, to consist of two layers ; * Some authors recommend boiling the shoot or cutting, in order to separate the cuticle ; but although it is thus readily separated, yet, the boiling coagulates and thickens its sub- stance, rendering it opaque and destroying its natural structure. LECT. VII.] ANATOMY OF 8TEMS. 319 the outer being the unorganized pellicle of true epidermis, and the inner a vascular texture, com- posed of minute vessels which terminate externally at the surface of the stem, and internally in the cellular integument *. These are, apparently, an- nular vessels with oblong pores ; and, although I have never been able completely to satisfy myself that they penetrate the real epidermis, yet, they probably do so to perform the office of exhalants or of absorbents. The abrupt manner in which these vessels terminate in the cellular integument, readily accounts for the facility with which the cuticle separates from that portion of the bark -f-. Such is the cuticular portion of the bark of the Horse Chesnut ; but the structure of this part is not the same in all ligneous dicotyledonous stems. In that of the Pear, Pyrus communis, it consists rather of transverse cells than of vessels, the outer series of which is covered by the real epidermis: this is the case also in the lesser Pe- riwinkle, Vinca minor, in which there are three series of such cells ; in the Laburnum, Cytisus * Vide Plate 6, fig. 2, a. f It was probably this vascular part of the cuticle, which led Du Hamel (Phys. des Arbres, liv. 1. c. ii.) to describe the vegetable epidermis as a tissue of delicate parallel fibres inos- culating at regular intervals, or united by lateral fibres, so as to constitute a network, the meshes of which are filled up with a thin, transparent pellicle. 320 CONSERVATIVE ORGANS. JJLECT. VII. Laburnum, it is composed of the epidermis simply covering a layer of an irregularly cellular or spongy character ; in the Laurustine, Viburnum Tinus, of one layer of cells covered by the epider- mis ; and the same is the case in the Vine, except that the cells are extremely minute, and oblong in the length of the stem, having the appearance of vessels in the transverse section. These and similar varieties in the structure of the cuticle account for the want of coincidence in the descriptions of authors. The true epidermis or exterior layer of the cuticle is necessarily cribriform, whether it act as an exhaling or an absorbing surface ; and the man- ner in which the pores are arranged, does not differ less, in different plants, than the structure of the interior layer. It is frequently studded with hairs, glands, and prickles ; but, as these are not peculiar to stems, their particular struc- ture shall be demonstrated, when we treat of the general vegetable appendages. In young and succulent shoots, the cuticle is generally almost colourless, and semi-transparent, transmitting the green colour of the exterior part of the cel- lular integument over which it lies; but it becomes opaque and coloured by age, or rather on losing its vitality ; for, as it is annually reproduced, on the ligneous stems under consideration, the old layer, if it does not fall off, cracks and is pushed LECT. VII.] ANATOMY OF STEMS. 321 outwards by the increase of the diameter of the stem ; and the accumulation of such layers forms the rugged surfaces of stems, as for example of the Elm, the Oak, and the majority of trees *. In the greater number of instances it cracks verti- cally, and is pushed outwards with a portion of the cellular integument by the new epidermis, which can be brought into view by removing these rugged portions. In others it splits horizontally, and the new cuticle is formed immediately under the old, which, after a time, detaches itself in fragments ; or, there is a succession of cuticles, which, although one is formed every year, yet do not separate annually, but occasionally only, in multiplied layers, that can, however, be readily detached from each other, as in the Currant and the Paper Birch. Some trees, the Plane for ex- ample, annually throw off the cuticle at once, in large flakes ; and in this respect, such plants re- semble those reptiles that cast their skins or their crusts, as the snake, the spider, and the lobster. 2. The Cellular integument. On carefully raising the cuticle of the young shoot of the Horse Chesnut, we find under it a cellular layer ; which, in a transverse section of the stem placed under * In Plate 6, fig. 10. a. represents a microscopic view of the various layers which form the rough cuticle of an old stem of the Lilac, Syringa imlgaris ; the innermost only of which retains its vitality. VOL. I. Y 32*2 CONSERVATIVE ORGANS. [LECT. VII. the microscope, is seen to consist of two distinct parts, both cellular, but nevertheless different. The exterior, or that on which the cuticle imme- diately reposes, appears to be composed of a dark green, semiorganized pulp, in which the cells are irregular both in their dimensions and form (vide Plate 6, fig. 2, 6.), and has somewhat of the as- pect, as Mr. Keith aptly expresses himself, of " a " distinct and separate epidermis in an incipient " state, rather than a true and proper pulp :" while the interior is less coloured and composed of regular hexagonal cells (Plate 6, fig. 2, c.), the sides of which are perforated and frequently studded with small granular bodies. It is the ex- terior layer of the cellular integument, which is the seat of colour of the young twig, and the green hue of which is transmitted through the yet semitransparent cuticle : its appearance, and the fact that it is annually reproduced, led Mr. Keith to believe that it is really the next year's cuticle in an incipient stage of organization. But the vertical direction of the cells, while those of the cuticle are horizontal, is sufficient to overturn this opinion, (Plate 6, fig. 7, a. I. 2.) These two portions of the cellular integument are particu- larly noticed by Mirbel, who denominates the exterior the herbaceous tissue, and the interior the parenchyma ; and conceives, with much pro- bability, that the deeper green colour of the latter 2 LECT. VII.] ANATOMY OF STKMS. 323 depends on the exposure of its juices to the light and on the resinous nature of these juices. He re- gards the whole of the cellular integument, also, as a glandular body serving to separate the trans- pirable matter from the other fluids * ; an opinion which I shall have occasion to notice more par- ticularly when we investigate the functions of the stem. The cells of the interior portion, in the young shoot of the Horse Chesnut, are very regular hexagons, except in those places where there is any pressure, or where the adjoining parts require a variation of form, when a change takes place; but, independent of these circum- stances, the pure hexagonal form does not prevail in the stems of every species of the natural tribe of plants under examination. Thus in Privet, Ligustrum vulgare, the cells are variously formed, some being nearly circular, others rudely ellip- tical, and some very obscurely heptagonal : in the Elder, Sambucus nigra, they are equally irregu- lar ; in the Common Lilac, Syringa vulgaris, the proportion of real cellular matter compared with that of the semi-organized pulp is small, and the cells, which are of an oblong figure, are com- pletely filled with minute amylaceous granules ; and this is the case, also, in the Laburnum, Cytisus Laburnum, in which, however, they * Elemens de Physiologic vegetate, l ere par tie, p. 103. Y-2 324 CONSERVATIVE ORGANS. [LECT. VII. are regular hexagons : in the Pear, Pyrus com- munis, they resemble globular utricles : and in the Rock Rose, Cistus Ledon, they are irregu- lar oblong hexagons. Such are the diversities of figure of these cells ; but it is still a ques- tion whether the membrane of which they are composed be single or double, as I formerly re- marked (page 75), in describing the cellular tex- ture among the general vegetable components? Senebier and Link are both of opinion that each cell is a separate utricle, completely distinct from those which are in contact with it, and conse- quently that the partitions are double on every side. Link further contends * that there are no visible organic pores in these partitions, the fluids passing from the one to the other by a double fil- tration ; and that the appearance of pores is occa- sioned by small amylaceous granules scattered over their surfaces. It is certainly not easy to deter- mine this question as far as regards the double or single nature of the cellular membrane, although I am disposed to believe it is double ; both from the appearance which it presents under the micro- scope, and also from the greater facility which such a supposition affords of explaining the origin, and the hexagonal figure of the cells, a point which shall be discussed in its proper place ; but, if * Vide ROMER, Collect. Botanic, facs. 1. p. 163. LECT. VII.] ANATOMY OF STEMS. 325 any confidence is to be placed in the microscope, there can be no doubt of the existence of the cel- lular pores. That many of the cells, however, are filled with minute particles, is perfectly evident ; and the number of these is always greater where a branch is given off. Their use is yet unascertained, but it is not improbable that they are of a nutri- tive nature, and deposited in the cells to be dis- solved by the ascending sap for the evolution of new parts. Independent of these particles, the cellular integument is filled both with coloured and colourless secreted juices ; and it is very pro- bable that this part performs some changes on the sap thrown into its cells, similar to those effected in the leaf. The cellular integument is partially destroyed, and reproduced, a great part of the old portion being pushed outwards with the cuticle which is annually detached ; while new cells are added to that which remains at the time the new cuticle is produced. 3. Vascular layer. Imbedded in the cellular integument and impinging on the internal sur- face of the bark, are distinct bundles of entire vessels, each of which is so arranged as to pre- sent, in the transverse section of the stem under consideration, a semilunar aspect * ; and, in the * Vide Plate 6, fig. 2, d. Y3 326 CONSERVATIVE ORGANS. [LECT. VII, longitudinal section, that of a fascis of flexible cords, readily separable from each other, and from the surrounding cellular substance * ; which is condensed where it comes in contact with these bundles. These vessels are supposed to convey downwards the proper juice of the plant, ela- borated from the sap, by the action of the light and air in the leaf; and this opinion is supported by the fact, that it is from them the milky juice of the Fig-tree and the coloured juices of other plants exude, when the stem is trans- versely divided. In some stems, as, for example, that of Laburnum, Cytisus Laburnum, the vascu- lar bundles coalesce, and form nearly one conti- nuous layer or circle around the wood ; and in others, although they do not actually coalesce, yet, they approach so close as almost to assume the same character. As the stem increases, these vascular bundles become impervious, and are pushed outward with the cellular integument, giv- ing place to a new layer which is annually pro- duced. 4. Inner bark. Immediately under the vascu- lar bundles, we find another layer -f- which consti- tutes the internal boundary of the bark. In the transverse section of the stem of the Horse Ches- nut now before us, it appears under the microscope *Vide Plate 6, fig. 7, a. 3. t Plate 6, fig. 2, e. and fig. 7, a. 4. LBCT. VII.] ANATOMY OF STEMS. 327 to consist of the extremities of longitudinal fibres closely united together ; and, in the tangental section, these fibres are seen running in a waving direction and touching each other at certain points only so as to form oblong meshes, which are filled with cellular matter. The nature of this structure will be better understood by referring to the mag- nified marginal plan, in which a. a. a. represent the reticular ar- rangement of the longitudinal fi- bres, and b. b. b. the cellular meshes. This layer is denominated LIBER, a name imposed from its having been employed to write on before the invention of paper. As the network formed by the dividing threads of the meshes is not readily dissolved in water, whilst the cellular matter which fills them up is remarkably soluble, the liber of some plants, for example the Daphne lagetto, when soaked in water and afterwards beaten, forms a very beautiful vegetable gauze ; which may be used as an article of dress. A coarser specimen of this gauze, or lace, is seen in the bark of many of our indigenous trees, par- ticularly the Oak, when it has been long exposed to the weather, after being separated from the trunk. This regular arrangement, however, of the longitudinal texture of the liber, is not found in every instance ; for in the Fir and some other v4 CONSERVATIVE ORGANS. [LECT. VII. trees the longitudinal threads are seen lying nearly parallel to one another, without any meshes or intervening cellular matter. Like the other parts of the bark, the liber is annually reproduced. The old layer loses its vitality, and is pushed out- wards by the new ; the accumulation thus formed constituting what botanical writers have called the cortical layers, which Malpighi supposed de- rived their origin from the older bark. The vitality of the stem of dicotyledonous plants is more conspicuous in the liber than in any other part. If the bark be wounded, or a portion of it be removed, layers gradually extend themselves from the liber on each side of the wound until it is closed up ; but, as this is not ef- fected in one year when the wound is extensive, and as the new layers are thrown out by the liber only which is annually renewed, the cicatrix, if the healed portion can be so named, always re- sembles a hollow cone, the base of which is the exterior of the trunk. The union of a graft, or of a bud taken from one tree and implanted on ano- ther, succeeds only when the liber of the bud, or the graft and that of the stock is placed in immediate contact ; the union in these instances closely resembling that which occurs when two raw surfaces of a living animal body, or of two distinct animals, are retained for some time in con- tact. Grew, Malpighi, Du Hamel, and others LRCT. VII.] ANATOMY OF STEMS. 329 supposed that the liber annually changes, by hard- ening, into the alburnum or young wood, an opinion which is still maintained by some of the ablest phytologists * ; but which I shall afterwards prove to you is founded upon mistaken principles. It is through the liber, however, that the matter in which the new wood is formed, which annually augments the diameter of the trunk and branches, is secreted; and hence the importance of this portion of the bark. Such is the structure of the bark of the stems of woody dicotyledons ; and that of the root does not materially differ from it; any difference de- pending, perhaps, altogether on the medium in which these two parts are situated. In the bark the secreted juices of plants, and consequently their medicinal qualities, are chiefly deposited; but the consideration of the functions of this part and its properties must be deferred, until the whole of the structure of the stem has been de- scribed. b. The WOOD. Pursuing our investigation in the young stem of the Horse Chesnut ; when the whole of the bark is removed, we find, imme- diately under and slightly adhering to it, a firmer and more compact substance, which, both in a longitudinal and a transverse section, ap- * " Le liber endurcie, de verdatre qu'il etait, devient blanch- " atre, et prend le nom d'aubier." MIRBEL, Element de Phys. vcg. t. i. p. 106. 330 CONSERVATIVE ORGANS. [LET. Vlt, pears to constitute a cylinder, enclosing a co- lumn of spongy cellular matter or pith. This is the wood. It has been regarded 1 , in reference to the vegetable, as answering the same end as bone in the animal body ; but, except in its property of giving firmness and support to the plant, the ana- logy does not hold good. It is at first soft and vascular, and is then called Alburnum ; but it af- terwards becomes hard, and, in some trees, is of a density almost equal to that of iron. In a trans- verse section of our stem of Horse Chesnut, it ap- pears, to the unassisted eye, a continuous circle of a homogeneous structure, of a very light straw colour exteriorly or near the bark, and greenish interiorly, or where it is in contact with the pith ; but in some other trees, as for example the La- burnum and the Elder *, this circle appears tra- versed, at nearly regular distances, by rays of an evidently different structure. These are found, however, to exist also in the stem of the Horse Chesnut, and in every other woody dicotyledon when it is examined by a magnifying glass; and they are observed in the soft wood, or alburnum, as well as in the hard and most perfect wood. These two distinct parts, which constitute the wood, may be described under the names Concen- tric and Divergent layers ^. * Vide Plate 6, fig. 3. 4. 5. f These names are adopted by Mr. Keith (System of physio- logical Botany ) y and are more expressive of the parts they are in- tended to designate, than any others which have been employed. LECT. VII.] ANATOMY OF STEMS. 331 Placing a very thin transverse section of the stem of Horse Chesnut under the microscope, the wood no longer appears solid and compact ; but of an irregular reticulated texture -f-. In this state, however, the concentric and divergent layers are readily distinguished; the open spaces in the former being evidently the transverse orifices of divided longitudinal or vertical cells and vessels, whilst those in the latter are the lateral openings of horizontal cells ^. Let us now examine se- parately the minute structure of each of these parts as they appear in the stem of one year's growth. 1. The Concentric layers consist apparently of longitudinal fibres, which are, however, appa- rently not solid, but narrow tubes or oblong cells, the sides of which are thick and nearly opaque, and of vessels of different kinds. These are ar- ranged parallel to each other, except where they are separated by the divergent layers, as may be seen in a thin tangental section of any stem placed under the microscope ; and is rudely dis- played in the marginal plan, in which a. a. a. a. a. represents t Vide Plate 6. Fig. 2. * represents the first state of the soft wood or alburnum,/, the more perfect wood, and h. the orifices of the large vessels of the wood. J Vide Plate 6, fig. 2.g. 332 CONSERVATIVE ORGANS. [LECT. VII. the longitudinal fibres or oblong cells ; c. c. c. the vessels which in the Horse Chesnut are porous, and b. b. b. the exterior ends of divergent layers. In the alburnum, the walls of the concentric tubes are tender and transparent ; but by the deposition of ligneous matter in the membrane of which they consist, and in the tubes themselves, they become opaque and firm ; and according to the degree of this, the wood is more or less dense, hard, and tenacious. Other matters, also, are deposited in this part of the woody texture ; such for example as Guiaic in that of the Guiacum officinale, colouring matter in the Logwood, Haematoxylon Campechianum, and even silex, which has been extracted from the Teak wood, Tectona grandis, by Dr. Wollaston. The vessels of the concentric layers are chiefly porous and an- nular, and their section produces the openings ob- served in the transverse section of any stem* ; but besides these, in the circle of the wood of the first year's growth, a circle of spiral vessels surrounds the pith *f~. These are, however, justly regarded by * Vide Plate 6, fig. 2. h. and fig. 8. a. a. a. The tubular nature of the oblong cells, forming the concentric layers, is rendered evident in this section of the common Elder, fig. 8. at b. b.; and the distinct character of the divergent layers at c. c. Leuwenhock, in 1680, first delineated the porous vessels of the wood. Vide his Epist. Physiolog. p. 14-. 19. f Vide Plate 6, fig. 7. d. e. 7, 7, 7. In this figure, also, the first, or half-organized state of the alburnum, is represented between b. c. ; and the vascular structure of the perfect wood LECT. VII.] ANATOMY OF STEMS. 333 Mirbel not as vessels of the wood ; but of a dis- tinct sheath lining the wood, which he has deno- minated F6tui me*dullaire ; and there is un- doubtedly some reason for this distinction, inas- much as these vessels have not been detected in the wood, but always in immediate contact with the pith. I shall more particularly examine this opinion before describing the structure of the pith. 2. The Divergent layers * consist of flattened masses of cellular substance, which cross the con- centric layers at different parts, and, separating the bundles of longitudinal tubes of which they consist from each other, produce the reticulated arrangement seen in the tangental section of any steal ; the oblong tubes and vessels forming the tissue of the network, the meshes of which are filled up by the cells of the divergent layers. The individual cells, which are narrow and horizontal in their length, extend in series from the centre to the circumference of the wood; and conse- quently form nearly right angles with the tubes of the concentric layers^. They communicate with each other by pores ; so that fluids may readily pass through the whole series, and of between c. d., the smaller vessels being marked by the figure 5, and the larger by 6, showing distinctly their porous and an- nular structure. * Dr. GREW was the first author who described these layers under the name of insertions. f Vide Plate 6. fig. 7. *. *. 334 CONSERVATIVE OKGANS. [LECT. VII. course transversely through the wood ; and Mirbei remarks that, " in many coniferous trees the di- " vergent rays are not cellular ; but consist of ho- " rizontal tubes which extend from the medulla or " pith to the bark*." Whether they are cellular or tubular, the layers, or masses, are flat, or in plates, with the edges placed vertically, and thicker in the centre than either above or below, appearing therefore of a lozenge shape (see marginal plan, p. 331) when vertically divided ; whilst in their transverse section they display a slight inclination to the wedge form -J-. They are much more deli- cate in their structure than the concentric layers ; and readily dissolve, like the common cellular texture, so that when a thin tangental slice of wood is macerated in water, the divergent layers are decomposed and leave the meshes of the con- centric layers empty, displaying the appearance of a network or lace similar to that formed by the macerated liber. From the cellular texture of the divergent lay- ers, they are regarded by some authors as pro- cesses of the pith or medulla ; and hence have been named medullary rays ; and Mr. Keith ob- serves, they are " apparently nothing more than " the vesicles or cellular tissue of the pulp that " originally existed in the alburnum now deprived * Elemens de Phys. veg. l re partie, p. 110. t Vide Plate 6, fig. 2. g. LECT. VII.] ANATOMY OF STEMS. 335 " of its parenchyma ; but still filling up the in- " terstices of the concentric layers, and binding u them together like a cement.** But there is little difficulty in demonstrating the error of both these opinions ; for, examining the alburnum in a very early stage of its formation at the moment it is passing from the gelatinous state in which it is first deposited, we find the rudiments of both the concentric and divergent layers already assuming the form which they afterwards maintain. Were any further reason required to prove that the diver- gent layers do not originate in the pith, it would be found in the fact, that many of them cannot be traced to the pith ; although the more conspicuous of them traverse the whole of the wood, from the pith to the bark. Such is the structure of wood in the stems of one year's growth of ligneous dicotyledons ; and it is found nearly of the same structure in the root : a fact, which is rendered evident, not only by the microscope, but also to the unassisted eye, by the decomposition of the divergent layers in the ligne- ous part of roots which have been dug up and long exposed to the action of the atmosphere. The concentric layers, longer resisting the action of the weather, remain after the divergent have dis- appeared, and display a beautiful network, more or less open in its meshes, according to the den- sity or sponginess of the wood. 336 CONSERVATIVE ORGANS. [LECT. VII. Wood in its soft state, or that in which it forms the outer circle in every ligneous dicotyle- donous stem and branch, is, as has been already mentioned, named alburnum. While it con- tinues so, it is endowed with nearly as much ir- ritability as the liber; and, as shall be after- wards fully described, performs functions of great importance in the vegetable system ; but when it becomes hard these functions cease, and in time it loses even its vitality ; not unfrequently decaying in the centre of the trunk of trees ; which, nevertheless, still flourish and put forth new shoots as if no such decay existed. To carry on, therefore, the functions of the wood, a new circle of it is annually formed over the old ; and thus, also, the diameter of the trunk and branches present, by the number of these an- nual zones, a pretty correct register of their age, each zone marking one year in the life of the part*. There are, however, exceptions to the criterion thus afforded of the age of the plant, for circumstances may occur to prevent the zone from being formed of a thickness which will be per- ceptible after a few years have passed over, and it is pressed between other zones. If the summer * Vide Plate 6, figures 3, 4, 5, which represent sections of the same stem, in the first, second, and third year of its growth ; and fig. 6. which displays two zones, as they are seen in a longitudinal section of the stem of the Elder. LECT. VII.] ANATOMY OF STEMS. 337 be unusually cold, or if the leaves of the tree or the shrub happen to be much devoured by caterpil- lars, it gains very little that season in diameter. From the same cause the zones are also of un- equal degrees of hardness : but, independent of the comparative density of each, the hardness of the whole increases with the age of the tree, so that they are hardest in the centre, and less and less hard as they approach the circumference. The outermost layer, being alburnum, is always soft, and continues so until another layer is formed over it ; but if the tree be barked the alburnum assumes the apparent character of wood in the same year ; and hence it has been recommended to bark trees the year before they are intended to be cut down. "The German foresters," how- ever, " have proved that wood treated in this " manner is less elastic, and is more easily in- " jured by humidity and insects * ;" which I conceive is owing to the natural change of al- burnum into wood not depending on a simple hardening or condensation ; but on such a de- position of ligneous and other particles in its tex- ture, as tends to increase the cohesive attraction of all its parts, and consequently to augment both its hardness and its elasticity ; while the ex- posure of the alburnum, by stripping off the bark, * Mirbel, Elemens de Phys. veg, l cfe partie, p. 110. VOL. 1. Z 338 CONSERVATIVE ORGANS. [LECT. VII. produces merely a simple condensation of the solid matter, a hurried crystallization of the salts, and a hasty consolidation of the other secretions. In- deed, when wood acquires its firmness by the na- tural means connected with its growth, it is a well- known fact that the hardest is always of the slow- est growth; as exemplified in the comparative hardness of the wood of the Oak, which is of very slow growth, compared with that of the Willow or Horse Chesnut, which are trees of rapid growth ; and even in that of the wood of the same tree when growing in a dry and in a moist situation. Much difference of opinion has existed among Phytologists regarding the origin of each suc- cessive layer of wood. Linneus conceived that it is formed from the pith, and added internally ; but the absurdity of this opinion must be imme- diately obvious to any one who examines the trans- verse section of any stem or branch more than two years old. Dr. Hales supposed that it is formed from the zone of the prior year, by the horizontal dilatation of the vessels, and "the " shooting of the longitudinal fibres lengthways, " under the bark, as young fibrous shoots of roots " do in the solid earth * ;" an hypothesis which has had almost as few followers as that of Linneus. Malpighi taught that the liber is annually trans- * Vegetable Staticks, p. 340. LRCT. VII.] ANATOMY OF STEMS. 339 muted into alburnum * ; an opinion which was af- terwards supposed to be fully established by the experiments of Du Hamel and of Dr. Hope, and is still maintained by Mirbel^. The first ob- ject of Du Hamel was to ascertain whether the new layer of wood was formed by the bark, or by the former layer of wood. He raised a portion of the bark of a growing tree, and introduced under it a piece of tin foil, over which he carefully bound down the bark ; and, after the wound was healed, allowed the tree to remain in the ground for some years. He then cut it down, and found layers of wood on the outside of the tin foil ; but none had been formed between the foil and the wood with which it had been placed in contact ^. This experi- ment, although it was decisive of the fact, that the new layer of wood is not formed by the old layer which preceded it, yet has been justly objected to by Mr. Knight as by no means confirming the opi- nion of Malpighi regarding the transmutation of the liber into wood; and the same objection may be applied to the following experiment of Dr. Hope, detailed by Sir J. E. Smith, on the authority of his son, Dr. Thomas Hope. " A longitudinal incision * Anat. Plantarum. f " La transformation du liber en aubier est prouvee par " Tobservation microscopique et par 1'experience." Elemens de Phys. veg. p. 106. J Phys. de Arbres, 1. iv. c. 7. z 2 340 CONSERVATIVE ORGANS. [LECT. VII. " several inches in length was made in the bark of " a branch of Willow, three or four years old, and " the bark loosened, so that it might be slipt " aside from the wood in the form of a hollow " cylinder, the two ends being undisturbed. The " edges of the bark were then united as carefully " as possible, the wood covered from the air, and " the whole bound up to secure it from external " injury. After a few years, the branch was cut " through transversely. The cylinder of bark was " found lined with layers of new wood, whose num- " ber, added to those in the wood from which it " had been stripped, made up the number of rings " in the branch above and below the experi- " ment *." But if these experiments do not prove the truth of Malpighi's opinion, they completely disprove the hypothesis of Hales; and throw great light upon the fact, that the alburnum is actually formed from the secretion deposited by the vessels of the liber ; an opinion which has been fully es- tablished by the experiments of Mr. Knight*)-. * Introduction to physiological and systematical Botany , chap. iv. f It is but justice to say, that although Dr. Hales states it as his opinion that " the new zone of wood is formed by the " shooting of the longitudinal fibres lengthways under the bark, " as young fibrous shoots of roots do in the solid earth ;'* yet that he afterwards remarks, " we may observe that nature has " taken great care to keep the parts between the bark and wood LECT. VJI.J ANATOMY OF STEMS. 34l Before, however, entering into the details of Mr. Knight's doctrine, it will be proper to notice that of Dr. Grew, who conceived that a new ring of sap-vessels is first generated in the mucilage thrown out between the bark and the wood, to which he gave the name of Cambium; and this ring of vessels, lining the inner surface of the liber of the former year, is converted into a new layer of liber that ultimately splits into two por- tions, the outer of which forms the new layer of bark, and the inner the new layer of wood. Not- withstanding the inconsistency of this hypothesis, in supposing that the mere separation of the two portions of the liber could produce, in one of these, a new organization and properties, so dis- tinct as those which the wood possesses from the liber, yet it has been characterized, by a late able writer on vegetable physiology, as " perhaps more " conformable to fact" than any other *. Du Hamel made several experiments to ascertain its truth. He passed threads of fine silver wire through the bark of a tree, some near the outer part, or towards the epidermis; others near the liber, others through the liber itself; and others between the wood and the liber. After " always very supple with slimy moisture, from which ductile " matter the woody fibres, vesicles and buds are formed." Ve- getable Staticks, p. 340. * Mr. Keith. 342 CONSERVATIVE ORGANS. [LECT. VII. some years, when the tree was cut down, those that were placed towards the epidermis were found covered by a thin, decayed, and friable crust only ; those that were placed near the liber were now among the external cortical layers ; those that were passed through the liber were now im- bedded in the wood ; as were those, also, that had been placed between the wood and the liber *. At first sight this experiment appears perfectly con- clusive of the truth of Grew's opinion ; but, when we consider the probability that the wood and the liber are formed at the same time ; for the vi- tal action which is capable of forming the one is undoubtedly equal to the generation of both ; the difficulty of drawing an accurate inference from such an experiment is obvious : and if our reason- ings must be hypothetical, there is certainly more wisdom in deciding in favour of that conjecture, which explains the effect by simple and direct means, than of that which supposes two causes, the one consequent on the other ; and the second involving a difficulty as great as that which it is intended to explain. But the true explanation of the phytological fact under consideration, was reserved for Mr. Knight, whose experiments and observations have settled almost every doubt upon the subject. Mr. Knight could not avoid admitting that the ex- periments of Du Hamel and of Dr. Hope were * Physique des Arbres t 1. iv. c. 7. LECT. VII.] ANATOMY OF STEMS. 343 sufficiently satisfactory, so far as regarded the fact that the presence of the bark is necessary for the formation of the new zone of wood ; but he denied their conclusions ; and it was still essential to ascertain, whether the wood is transmuted liber ? or, if not, whether the matter of which it is formed be a secretion from the bark, or supplied by some other of the vegetable organs ? That it is not transmuted liber is evident from the dissimi- larity of the liber and the alburnum. Thus, accord- ing to Mr. Knight *, the commencement of the alburnous layers in the Oak is distinguished by a circular row of very large tubes, which appear in spring, arranged in ridges in a gelatinous mass, beneath the cortical vessels ; but such tubes are not found in the bark of the tree, which would be the case were the alburnum a transmutation of the liber. The bark of the Wych Elm (Ulmus monta- na) also is so fibrous and tough that it may be formed into cords, while that of the Ash (Faxinus excelsior) is very fragile and not at all fibrous; ne- vertheless the wood of both these trees> and conse- quently the alburnum, is nearly alike-f-. As con- vincing a proof also is the simple fact, that the layer of alburnum is often more than twice the thickness of the bark. But the question was set- tled by the following experiment of Mr. Knight. * Phi Intophical Transactions, 1805. f Ibid. 1808. z4 344 CONSERVATIVE ORGANS. [LECT. VII. He cut out a ring of bark from the stern of an Apple tree, and another from that of a Crab tree, which were particularly distinguished from each other by the colour of their wood. He then trans- posed these rings, applying and fixing, by means of a firm bandage, the bark of the Crab tree quite round the uncovered part of the stem of the Apple tree, and that of the Apple tree round the stem of the Crab tree. The air was excluded from both by a plaister of bees wax and turpentine, and covered with well-tempered clay. The in^ ner surface of the Crab tree bark had sinuosi- ties that corresponded with elevated parts of the alburnum of the tree from which it was taken, occasioned by the former extension of many branches ; but that of the Apple tree bark was smooth. In a short time a vital union took place between the applied pieces of bark of both trees, and the bark and alburnum of the trees on which they were bound ; and before the end of the ensu- ing autumn "it appeared evident that a layer of " alburnum had been, in every instance, formed " beneath the transposed pieces of bark, which " were then taken off." " Examining," conti- nues Mr. Knight, ".the organization of the albur- " num, which had been generated between the " transposed pieces of bark of the Crab tree, and " which had formed a perfect union with the al- LECT, VII.] ANATOMY OF STEMS* 345 " burnum of the Apple tree, I did not discover any " traces of the sinuosities I had noticed ; nor was " the uneven surface of the alburnum of the Crab " tree more changed by the smooth transposed " bark of the Apple tree* The newly generated " alburnum, beneath the transposed bark, ap- " peared perfectly similar to that of other parts of " the stock, and the direction of the fibres and " vessels did not in any degree correspond with " those of the transposed bark *." Nothing, in my opinion, can be more decisive of the ques- tion than this experiment ; for, although Mr. Knight himself modestly suggests that it is not " calculated to prove that the newly generated " bark was not converted into alburnum ;" yet it is not probable that the merely transposing the bark of one tree to another would alter the ori- ginal features of the liber of the transferred por- tion in so short a period as one season, if such a change should even afterwards occur ; of which, however, we have no evidence. The obvious con- clusion therefore to be drawn from this experiment is, that although the alburnum is generated through the medium of the bark, yet, it is deci- dedly not transmuted liber. The opinion of Mr. Knight is, that the bark deposits the alburnous matter ; but that the leaves * Phil. Trans. 1808, Part I. p. 104-5. 3 346 CONSERVATIVE ORGANS. [LKCT. V1L are the organs in which this matter is elaborated from the sap ; or, in other words, that the albur- num is generated from the cambium, which is part of the proper juice of the plant, formed by the exposure of the sap to the light and air in the leaf, and returned from it by the vessels that pass down from the leaf into the interior bark, by which it is deposited. To determine this point, he re- moved narrow circles of bark from shoots of Apple trees, " leaving a leaf between the places where " the bark was taken off; and on examining them " frequently during the autumn," he found that the diameter of the shoot between the insertion of the leaf-stalk and the lower incision was as much increased as in any other part of the tree ; but when no leaf was left " on similar portions of in- " sulated bark, on other branches of the same " age, no apparent increase in the size of the wood " was discoverable*." No other inference, than that the leaf is the essential agent in producing the increased diameter of the wood, could be drawn from this experiment: and Mr. Knight further found that where the deposition of the proper juice returned from the leaf is greatest, that is, at the points where the returning vessels enter the inner bark, there the formation of al- burnum is observed to take place in ridges cor- * Phil. Trans. 1801, P. I. p. 2, p. 335. LECT. VII. ] ANATOMY OF STEMS. 347 responding to the number of the vascular bundles. The fact, indeed, that the leaf is es- sential for the formation of wood, had been ob- served by Dr. Hales, who took off circles of bark half an inch in breadth, at several places, from two thriving shoots of a dwarf Pear tree, leaving on all the remaining intervening ringlets of the bark, except one, a leaf-bearing bud, which produced leaves the following summer. Each ringlet, on which a bud was left, grew and swelled below the bud, or at its lower edge ; but that one on which no bud was left " did not increase at " all * :" but he drew a very different and less pro- bable conclusion from his experiment, which it is unnecessary to mention in this place. The above-mentioned experiments of Mr. Knight readily explain why trees and shrubs, the leaves of which are destroyed by caterpil- lars, form scarcely any new wood in that sea- son ; and, indeed, every one who has ever pruned a tree, or shortened a growing twig, must have observed that the part above the last leaf al- ways shrivels and dies, while all below it con- tinues to live and increase in diameter: and we observe the same thing in the lower part of the stem of a young tree, when a portion of the bark has been gnawed off by sheep, or accidentally de- stroyed. The part above the wound continues to * Vegetable Staticks> p. 145, fig. 28, 29, 30. 348 CONSERVATIVE ORGANS. [LKCT. VII augment in diameter, as represented at a in the marginal cut, because it is supplied by the alburnous matter furnished from the leaves ; but that below it, b, ceases to grow, and continues in the same state, the communication by the interior bark between it and the foli- age being completely destroyed. From the same cause, also, the pith of a stem is thrown apparently out of its centre, for some distance below the point where a branch is given off; a circumstance which I shall very soon have occasion more par- ticularly to notice *. The only objection of any weight which has been advanced against Mr. Knight's theory, is founded on the fact, that when a stem is wholly, or in the greater part stripped of its bark, and the denuded surface excluded from the action of the air, a glary fluid is exuded from the albur- num, or soft wood, which gradually becomes or- ganized and cellular. Detached spots of bark are thus reproduced, and, gradually extending, coalesce, until the stem is again clothed with * Vide Plate 6, fig. 6. which displays, moderately mag- nified, a longitudinal section of an Elder (Sambucus nigra) of two years growth, with a luxuriant branch of the same age : a. the trunk has the pith c. in the centre, except below the branch b. where the additional wood e. [augments the diameter of the stem on that side, LECT. VII.] ANATOMY OF STEMS. 349 a new bark, capable of performing all the func- tions of the original. This fact was first observed by Du Hamel. But Dr. Hope's experiment proves that a similar matter is exuded from the liber, when the bark is detached from the wood ; and Mr. Knight observed that in this case it becomes sooner organized than when it is exuded from al- burnum ; and as we cannot conclude that the new bark is first generated from the former year's al- burnum, and then the new alburnum from this newly formed bark, the only mode of getting over the objection is, by supposing that, under that state of circumstances which as it were obliges the wood to form a new bark, the descending juice from the leaves being impeded in its course downwards through the bark, finds its way into the alburnum in much greater quantity than is required for the ordinary purposes of the plant in that part of its structure ; and the alburnous vessels taking on a retrograde action, it is thrown out in the manner described at different points over the denuded trunk. This opinion is sup- ported by analogy in the animal body, on which, when one organ is destroyed, its function is per- formed by another. We are, therefore, fully war- ranted in adopting Mr. Knight's theory, as far as it maintains that the wood is formed from the proper juice which descends from the leaf through the inner bark ; but, in doing so, there are some 350 CONSERVATIVE ORGANS. [LECT. VII. points which have been overlooked by Mr. Knight, and to which it is necessary to direct your at- tention. In explaining his theory, Mr. Knight seems to .imply that the whole of the change which the sap undergoes is effected in the leaf, and that the liber is the mere medium of transmission of the alburnous matter. But if, as we must admif is most probable, the whole of the secretions of the plant are produced from the same proper juice elaborated from the sap, by its exposure to light and air in the leaf; and consider the great diver- sity of these ; it is more likely that some altera- tion takes place in the bark, previously to the cambium being thrown out by its vessels ; and, con- sequently, we must admit the force of Mr. Keith's suggestion, that it is " possible the proper juice " may receive its final degree of modification in " the bark itself*." It is by such an admission only that we can satisfactorily explain the fact observed by Mr. Knight in one of his experiments, that a small quantity of wood was generated even at the lower lip of an insulated portion of bark, on which there was neither bud nor leaf -f~. It is also necessary for the sake of those unacquainted with physiological reasonings, to remark that it is not * System of physiological Botany -, vol.ii. p. 229. f Phil Trans. 1803. LECT. VII.] ANATOMY OF STEMS. 35 1 to be supposed, from the term deposition of albur- nous matter which Mr. Knight employs, that no- thing more is necessary for the formation of the al- burnum than the deposition of that matter. For although alburnous matter may be justly said to be generated from the sap after its elaboration in the leaf, whether we designate it by the term alburnous matter, or cambium, or proper juice, yet it is merely the pabulum ; the organization of the alburnum, or the transmutation of the cambium into its cel- lular and vascular texture, being the result of the vital principle operating upon it, in a manner which we do not understand, and not of any simple coa- gulation, or attraction, or chemical affinity of its parts, in any way similar to what would take place in the same matter, wherever deposited, if de- prived of vitality. The simple fact, therefore, is, that the sap is changed into proper juice in the leaf, and returned into the bark, where part of it being poured out in a gelatinous form between the liber and the wood, there becomes the raw material from which the new zone of wood, in its state of alburnum, and the new layer of liber, are ma- nufactured by the vital principle inherent in the living plant. Such is all that is necessary to be known, in the present stage of our investigation into the origin of the annual zones of wood, by which the diameter of ligneous dicotyledonous plants is 352 CONSERVATIVE ORGANS. [LECT. VII. augmented. Many important points, indeed, re- lating to the inquiry, and involving other hypo- theses, still remain to be examined ; but these must be deferred until I again bring the question, as it is connected with the general theory of the growth of the vegetable body, under your consi- deration *. The following simple facts, therefore, * Although it is unnecessary to load the text with any fur- ther details of the various hypotheses, which have been ad- vanced in explanation of the origin of the annual zones of wood, which increase the diameter of ligneous dicotyledons, yet it may not be improper to give here a slight sketch of that of M. Aubert du Petit Thouars, which has for some years past divided the opinions of the French Botanists. He regards each gem which exists at the axilla of a leaf, as an em- bryon resembling, in some respect, that contained in the seed ; and which, to effect its evolution, draws its nourishment from the succulent parenchyma on the bark on which it is seated, and with which he supposes it to have an immediate communi- cation. As soon as the gem begins to be evolved, it sends down fibres, which our author regards as the roots (veritables racines) of the gem ; and these growing and extending by the organic power (qui, comme I'electricite et la lumiere, semble ne point connoitre de distance), pass down between the wood and the bark from the gem whence they originate to the roots of the tree, taking up, as nutriment (la matiere de leur accroissement), in their passage the viscous fluid, or cambium, which at this season is found between the wood and the bark. The sum of these radical fibres constitutes the new layer of wood, which appears in a concentric circle, owing to the leaves, and consequently the gems, whether they are opposite or alternate, rising on every point of the circumference of the LECT. VII.] ANATOMY OF STEMS. 353 relative to the origin and increase of the concen- tric zones of wood, observable in the transverse section of any tree or shrub of more than one year's growth, and which has sprung from a seed stem or shoot. In the same manner, he supposes the liber is formed by an equal elongation of the fibres df the interior bark of the gem ; so that each gem has thus a double commu- nication with the root of the tree. The latter part of our au- thor's opinion closely resembles that promulgated by Dr. Dar- win, who supporting the theory of the individuality of buds, thus expresses himself: "The bark is only an intertexture " of the caudexes of the numerous buds, as they pass down " to shoot their radicles into the earth" (Phytologia, 4to. 1800, chap. i. 1, 2. 3) ; but he also so far maintained the opinion of Du Hamel, as to suppose that as these caudexes form a new bark over the former one, that of the last year is transmuted into alburnum. M. Aubert Du Petit-Thouars explains the formation of the medullary rays by supposing that the fibres which, according to his doctrine, constitute the new layers of wood and liber, determine the formation of a certain quantity of parenchyma, which is deposited interiorly by the ligneous fibre, and exteriorly by the fibres that form the new liber. (Essais sur la Vegetation consider ee dans le Developpement des Bourgeons, Paris, 1809, 2 e Essai.) I shall not endeavour to point out the very hypothetical nature of this doctrine ; but merely observe, that, with the exception of the remark that each gem is a distinct embryon, the whole is founded upon assumption ; and that it is utterly destroyed by the simple fact, that isolated spots of bark and alburnum are formed on de- corticated stems, which cannot be the roots descending from the gems, unless we suppose that these have penetrated the wood and again protruded at the points where the new patches of bark and alburnum appear. VOL. I. A A 354 CONSERVATIVE ORGANS. [LECT. VII. with two or more cotyledons, are to be regarded as fully established : 1. That the proper juice descending from the leaf through the vessels of the bark, and poured out between it and the wood of the preceding year, is the material from which the new wood or alburnum is formed. 2. That the organic power, or vital principle, inherent in the plant, transforms this viscous fluid into cells or vessels, or regularly organized alburnum ; the divergent layers being formed at the same time as the vertical or concentric layers. 3. That under certain circumstances, such as the entire decortication of a stem, the lateral com- munications, which exist throughout the vegetable structure, may conduct the descending juices through the wood, so as to be thrown out in detached spots on its surface ; and there be trans- formed into new bark and alburnum. The use of the wood to the plant, exclusive of its power of supporting and elevating the buds with their leaves and fructification in the atmo- sphere, is chiefly confined to its soft or alburnous state. In this state it is endowed with a high degree of irritability. But if it be freely exposed to the atmosphere during a few hours only, all vegetation for ever ceases on that surface ; and, although the bark may close above it, and new wood be formed over it, yet, no vital union LECT. VII.] ANATOMY OP STEMS. 355 takes place, and the new wood will always remain distinct*. In this state, also, through its tubes the sap is raised to the summit of the highest trees ; and, according to Mr. Knight, when this function ceases in winter, it becomes " a reservoir " of the sap or blood of the tree, as the bulbs of " the Hyacinth and Tulip, and the tuber of the " Potatoe certainly do of the sap or blood of these " plants ;" an analogy which is good ; but we must remark, in making this admission, that it is not the sap, but the proper juice (succus proprius), which is there laid up, and is dissolved by the first ascending sap in the spring. The alburnum does not attain its entire thick- ness at once ; but continues to increase during the greater part of the summer, at least till after the midsummer shoots are protruded ; and even, if circumstances occur to stop its progress, such for example as a week of very cold weather in the middle of the season, two layers may be formed in the same year ; an event not unlikely to happen in this variable climate, and which throws an uncertainty on the mode, which has been al- ready alluded to, of determining the age of trees by counting the number of the concentric zones of * In this new wood the divergent rays are distinctly seen ; a sufficient reason, as Mr. Knight remarks, for believing that they are not processes of the pith. AA2 356 CONSERVATIVE ORGANS. [LKCT. VII. wood displayed in the transverse section of a stem. As the alburnum changes into wood it loses its irritability; but an unexceptionable theory of the means by which this change is effected, is still a desideratum in vegetable physiology. The opinion that the lignification is the consequence of the mere loss of fluid parts, and the approximation of the solid, cannot be admitted as correct; because, in this case, the drying of the alburnum even after a tree is cut down, would give it the consistence and characteristics of perfect wood, which is not the fact. I am inclined to believe, that the ligneous matter is deposited in a manner somewhat ana- logous to the deposition of the phosphate of lime in bones ; and that this deposition continues to be effected long after the alburnum assumes the cha- racter of wood. The heart wood, even of an old tree which is sound, contains some moisture as long as the tree continues to grow, which can only be accounted for by supposing that it still lives *, and, consequently, is in some degree under the control of the organic power ; and, as a lateral communication is preserved between the exterior and the interior of the stem by means of the diver- gent rays, there is no improbability in supposing that a sufficient supply of proper juice finds its * Darwin (Phytologia, xviii. 2, 12.) asserts that the heart or internal wood is not alive ; but this is not the case, for the whole is alive whilst the tree remains sound. LECT. VII.] ANATOMY OF STEMS. 357 way to the innermost layer of wood, until that has received its final degree of induration. I know of no observations which tend to prove that the concentric zones are diminished in thickness as they increase in solidity and density, which would be the case, did they shrink or suffer any com- pression ; and if it be true that no such change takes place, their progressive induration can be explained only by admitting that there is a con- tinual deposition of new matter. The truth of such an opinion is further confirmed by the change of colour, which the alburnum undergoes in pass- ing into the state of perfect wood. With regard to the period at which the wood attains its final degree of induration, I would say> with Mr. Keith, that perhaps no layer has acquired this state until " such time as the tree has arrived at " its full growth*." Returning to our shoot of Horse Chesnut, if we scoop out the pith from the ligneous cylinder that encloses it, we perceive this is lined with a thin green layer or coating; which, to the un- assisted eye, appears to resemble in its structure rather the cellular integument of the bark than any part of the surrounding wood. This is the MEDULLARY SHEATH (etui mtfdullaire) of Mirbel and the French Botanists : for, although it was * System of physiological Botany , vol. ii. p. 230. AA3 358 CONSERVATIVE ORGANS. [LECT. VII* first noticed by Dr. Hill, who named it Corona, yet, it has been overlooked or confounded with the wood, or the alburnum, by almost every suc- ceeding British Phytologist, until Mr. Knight's at- tention was directed to it in searching for his cen- tral vessels. It is readily distinguished, in either a transverse or a longitudinal section of many stems, by its green colour, which appears deeper as con- trasted with the dead white, the more usual hue of the pith which it surrounds ; but it is also easily traced in the succulent dicotyledonous stem as soon as it is evolved from the seed, separating the pith from its herbaceous investiture. When viewed under the microscope the me- dullary sheath appears to be composed of a cel- lular substance, in which are imbedded longi- tudinal layers of spiral tubes *. It is not easy to comprehend the meaning of Mr. Knight, when he speaks of another description of vessels as being found here, to which he says " the spiral " vessels are every where appendages," and which he names central vessels, " to distinguish them " from the spiral tubes and the common tubes of " the wood-f~." In the stems which I have exa- * Vide Plate 6, fig. 7. in which the space from d. to e. re- presents the appearance of the medullary sheath in a longitudi- nal section of an annual twig of Horse Chesnut ; 7. 7. 7. the bundles of spiral vessels ; and 8. the parenchyma, or green cel- lular lining of the sheath. f Phil. Trans. 1801. LBCT. VII.] ANATOiMY OF STEMS. 359 mined, the spiral vessels are seen in some instances detached, or not immediately accompanied by any other description of vessels, and in others they are accompanied by either cribriform or annular ves- sels according as the one or other of these are the common vessels of the alburnum and wood. Thus, in longitudinal and in tangental sections of the medullary sheath of the Horse Chesnut, the spi- ral vessels are accompanied with the cribriform vessels only of the wood ; and such is, also, the case in the Elder (Sambucus nigra), and in the Lilac (Syringa vulgaris). In the Elastic-gum Fig tree (Ficus elastica), the sheath of which is not dis- tinguishable by colour, the spiral vessels are seen close to that part of the first year ! s wood, which touches the pith, and consequently in the situation of the sheath ; but they are not immediately ac- companied by any other vessels : those nearest to them, however, are cribriform, like the larger vessels of the wood. In the Medlar (Mespilus), in which there is, likewise, no evident medullary sheath, the spiral tubes, which are very numerous, are accompanied by the common vessels only of the alburnum : and in the Cinnamon (Laurus Cinnamomi) the accompanying vessels are annular, such being the vessels of the alburnum and the wood. The cells of the medullary sheath (Plate 6, fig. 7, d. e. 8.) are narrow and oblong ; and, therefore, when it is not coloured it is scarcely A A4 360 CONSERVATIVE ORGANS. [LECT. VII. distinguishable from the wood, except by the spi- ral vessels ; which have not yet been discovered in any layer of formed wood subsequent to the first; for their apparent existence in stems and branches of several years' growth, is owing to the lignification of the medullary sheath. Grew and Hedwig, however, have represented them as existing in the wood ; but, although I have searched for them in every species of wood, yet I have not been able to detect them ; and as the same result has followed the investigations of Du Hamel, Mirbel, Knight, Mr. Keith, and others, I am inclined to regard my conclusion as correct. The cells which are between the layer of spiral vessels and the pith (Plate 6, fig. 7, d. e. 8.) ; and which are the site of the colouring matter, when this part of the stem is green, as it is in the example now before us, have a cribriform struc- ture. The spiral vessels of the medullary sheath vary in their arrangement, and thus the widely separated bundles they form in the Elder, give the canal of the pith a furrowed character; in the Pear tree their arrangement produces an irregular pentagon ; and this is, also, the case in the Oak ; while in Laurel- leaved Magnolia, M. grandiflora, the bundles are seen at the distance of four or five diameters from each other^ and projecting forwards, so as to seem imbedded as it were in the pith. But, whatever LECT. VIJ.] ANATOMY OF STEMS. 361 may be the arrangement, it appears to be in a great degree regulated by the disposition of the leaves, into which the spiral vessels in every in- stance direct their course, leaving for that pur- pose the medullary sheath, and traversing the wood, a little below the insertion of each leaf. As the medullary sheath forms the only parti- tion between the bark and the pith in the tender succulent shoot, before the ligneous matter is de- posited, and is in its texture lax, and incapable of affording sufficient support to the delicate coats of vessels, such as are found in the alburnum, if these were distended with ascending sap, the vessels that run through it are of a different structure from those of any other part of the ve- getable. The elastic thread of which these spiral vessels are formed is tough, and possesses irrita- bility ; and being stimulated to action by the effort of the sap to dilate the diameter of the vessel, contracts in its length in each coil alternately, and after each contraction again returns to its first state, producing a vermicular motion, which enables these vessels to conduct forward the sap. Thus : the contraction in length of the portion of the thread which forms the first coi}, lessens the diameter of that portion of the tube, and hence the fluid contained within in it will be displaced and moved either upwards or downwards ; but as the resistance opposed to its return, or movement 362 CONSERVATIVE ORGANS. [LBCT. VII. downwards, is the greater owing to the pressure of the ascending sap, it must necessarily advance ; and this contraction being repeated in every suc- cessive coil, the fluid is moved forward with a suf- ficient impetus ; while the new quantity of sap which supplies the place of that carried forward, and which rushes into the coil at the instant of its relaxation, forming the basis of resistance to the return of the portion before it, and at the same time exciting a renewal of the contraction, its pro- gression must be uninterrupted. I should be an- ticipating what I have to say on the general ascent of the sap (or, as it has been erroneously termed, its circulation), were I now to explain the means by which this fluid is raised through the ligneous parts of a tree, until it arrives at the succulent twigs, in which alone the spiral vessels are active ; and it is, therefore, here merely necessary to add, that I believe these to be the only vegetable vessels endowed with contractility, or which act in any manner analogous to the arteries of animals. If this hypothesis be maintainable, the spiral vessels are the sap vessels of the succulent stem and the an- nual shoot of dicotyledonous ligneous plants ; and their spiral structure is essential for the performance of their conducting function, in the spongy medul- lary sheath, or cellular parenchyma in which they are imbedded. How long they continue to act as sap vessels it is impossible to conjecture ; but they LKCT. VII.] ANATOMY OF STEMS. 363 may maintain their irritability, and consequently their contractility, for two or more seasons, or as long as the medullary sheath remains succulent ; although, as it is not necessary for the progression of the sap that they should act by alternate con- traction and dilatation, after the alburnous or lig- neous vessels are completed, it is probable that they lose these properties, after the first year of the life of the stem or of the twig. If we inquire what are the opinions of other phytologists respecting the functions of the spiral vessels, we find Malpighi regarding them as bron- chia, or air-vessels *, and the same opinion is supported by Grew*f~, Hales |, and Du Hamel. Grew, however, believed that they acted as sap- vessels in the wood ||, in which, as I have already stated, he fancied he had detected them ; and Du Hamel once suspected that they might contain highly rarefied sap. The supposition that they are air-vessels, probably originated from their always appearing empty when examined ; but on the same account the animal arteries were regarded as air- vessels by the ancients and their followers, until * Anal. Plantarum. f Anatomy of Plants, fol. edit. \ Vegetable Statics, chap. v. Physique des Arbres. || His words are, " in the wood the sap ascendeth only by 4< the air-vessels." Veg. of Trunks, chap. i. 364 CONSERVATIVE ORGANS. [LECT. VII. the discoveries of the immortal Harvey demon- strated them to be blood-vessels. The opinion that they contain air only, is still maintained by Mr. Knight, who supposes them to be appendages to another set of vessels, which he denominates central, and through which he supposes the sap ascends as soon as it arrives at the bud or suc- culent shoot, " aided by the spiral vessels *." I have however demonstrated to you that the spiral vessels are unaccompanied by any others, at least in all those trees which I have examined, and are even at some distance from the alburnous vessels in the annual shoot of many plants. Mr. Knight, it is true, in another essay-}-, en- deavours to explain the meaning of his term cen- tral Vessels, by saying that they are the same as the tubular tissue of M. Mirbel ; but, as the spiral vessels form part of Mirbel's tubular tissue, which besides comprehends all the other species of vegetable vessels, this attempt at explanation only renders the subject more confused. Mr. Keith, in criticising Mr. Knight's explanation, has fallen into a curious mistake regarding Mirbel's opinion of the vessels which carry the sap. He says, " If we regard their (the central vessels) re- " spective functions, they can correspond only to " the small tubes, as it is by them alone, accord- * Phil. Trans. 1801. f Ibid. 1807. LECT. VII.] ANATOMY OF STEMS. 365 4 " ing to M. Mirbel, that the sap ascends." (Syst. of phys. Bot. vol. ii. p. 119.) Now, Mirbel's words are, " la seve monte par les gros vaisseaux." (EUmens, t. i. p. 208.) The cells of the medul- lary sheath are, indeed, oblong", and, therefore, somewhat tubular, and to these probably Mr. Knight refers ; but, admitting this, we cannot see in what manner the ascent of the sap through these cells can be aided by the spiral vessels, the bundles of which are, in many instances, placed at the distance of three or four diameters from each other, and consequently the intermediate cells are beyond the sphere of their influence. Mr. Knight's opinions on this subject have certainly much less weight than they usually and most de- servedly possess. Grew, as I have already stated, first suggested the idea of the spiral vessels acting as sap vessels, and Du Hamel supposed he had detected them in the performance of this function. Hedwig * also asserts, that he observed the sap issuing from the orifices of the spiral vessels in a horizontal section of the stem of the Pumpkin, Cucurbita Pepo, and the squirting Cucumber, Momordica Elaterium ; and Senebier ^, more lately, remarked the same occurrence in a section of the stem of the Sago * Fundamentum Hist. Nat. Muscorwn Jrondosoruntj Lip. 1732, p. 55. f Phys. veg. t. i. p. 107. 4 366 CONSERVATIVE ORGANS. [LKCT. VII. Palm, Sagusfarinifera. Dr. Darwin may perhaps, however, be regarded as actually the first who taught that the spiral vessels convey fluids ; and he suggested the idea, which, with some modifi- cations, I have adopted, of the manner in which they carry forward their contents. " It is easy," says he, " to conceive how a vermicular or peri- " staltic motion of the vessel, beginning at the low- " est part of it, each spiral ring contracting it- " self, till it Jills up the tube, must forcibly push " forward its contents without the aid of valves *." He, however, considered them absorbent vessels ; and erroneously supposed that they pass down the trunks of trees from the caudex of each bud, to the roots. Finally, Mirbel-j- regards the spiral vessels as sap vessels ; whilst Mr. Keith considers the reformed opinion of Grew the most correct, namely, that they transmit not only air but sap \. c. The MEDULLA or PITH. Returning to our shoot of Horse Chesnut, we find the tube which is formed by the wood and lined with the medul- lary sheath, as has been already described, filled with a white, dry, very compressible, spongy sub- stance : this is the Medulla or Pith. In the suc- culent state of a stem or a twig, it is turgid with aqueous fluid ; but before the wood is perfected it *Phytologia, sect. 11, 8. t Eltmens de Phys. veg. l ere partie, p. 205. J System of phys. Botany, vol. ii. p. 120. LKCT. VII.] ANATOMY OF STEMS. 367 becomes dry and spongy ; except near the termi- nal bud, or where branches are given off, in which places it long retains its moisture. The form of the pith is regulated by that of the cavity it fills, which in the majority of instances is nearly circular ; but to this there are many ex- ceptions. Thus, in a horizontal section of a young stem or twig of the Elder, Sambucus nigra, and the Oriental Plane, Platanus orientate, we find it circular, but furrowed by the bundles of spiral vessels of the medullary sheath. It is oval in the Ivy, Hedera Helix, and the Ash, Fraxinus ex- celsior ; irregularly oval and furrowed in the Ori- ental Plane, Platanus orientalis ; triangular in the Oleander, Nerium Oleander; pentangular in the Oak, Quercus Robur ; four-sided, with the angles obtuse or tetragonal, in common Lilac, Sy- ringa vulgaris, and yellow flowering Horse Ches- nut, ^Esculus flava ; pentagonal in the Walnut, Juglans regia, and hexagonal in the common Dog- wood, Cornus sanguined. M. de Beauvois is of opinion that the situation of the leaves on the stem regulates the form of the tube which the pith fills, an opinion which M. Mirbel * regards as fallacious ; but it is nevertheless true, that in the horizontal section of many stems, the form of the medulla differs at, and immediately under the * Etemens de Phys. v(g. 1 partie, p. 111. 368 CONSERVATIVE ORGANS. [LRCT. VII. places where the leaves are seated, from what it is in the intermediate spaces. In the Lilac, for example, the obtuse tetragon becomes an ellipsis near the insertions of the leaves, which are op- posite; while each angle of the Oleander is length- ened into a horn or process. But besides these diversities of form, the pith varies in diameter in other respects. In the young tree, of a few inches in height, it is smallest at the basis of the stem, largest in the middle, and smaller again at the summit ; and in the growth of each future year, nearly the same variations in its diameter are observable. The pith, in the majority of ligneous dicotly- ledons, is longitudinally entire; but in some, the Walnut for instance, it consists of a succes- sion of transverse diaphragms intersecting the hollow cylinder of the wood, with the interven- ing spaces empty *. In others the continuity of the medullary column is broken by ligneous plates, which proceeding from the side of the central tube, either partially intersect it or completely partition off portions of it, as in several of the Magnolias ; while in others, again, it is merely a spongy sheath, lining the interior of the cavity, as in the stem and branches of Woodbine, Loni- cera Periclymenum. Where the branches are *Vide Plate6, fig. 11. LECT. VII.] ANATOMY OF STEMS. 369 given off from a stem, a thread of medulla, in some instances, separates from the central column, and entering the branch, is gradually augmented to a diameter proportionate to that of the branch *. Such a circumstance led M. Aubert Du Petit-Thouars f to describe the medulla at the base of every bud, as an inverted cone, the apex of which originates from the pith of the branch on which the bud appears ; but in general this is not the case, the pith commencing in the bud itself, which originates at the surface of the stem ; and hence no direct union exists between the pith of the branch and that of the stem (see marginal figure). In the annual shoot, the wood shuts up the canal of the pith at its extremity, as soon as it ceases to grow for the season, as is seen in the longitudinal section of our shoot of Horse Ches- nut, immediately under the terminal budj; and thus isolates it from the shoot of the next year. In many plants this forms a kind of woody partition, which marks the limit of the growth of each year in the length of the stem ; but in others it is ab- sorbed, the continuity of the pith being, appa- * Vide Plate 6, fig. 6. representing a cutting of the common Elder; in which d. the pith of the branch b. is united by a small thread to e. the pith of the main stem a. f Histoire d'un Morceau de Bois, p. 153. t Vide Plate 5, fig. 15. VOL. I. B B 370 CONSERVATIVE ORGANS. [LECT. VII. rently, uninterrupted from the root to the apex of such stems. Those partitions are almost always present when the pith is composed of distinct plates, as in the Walnut, or of a spongy sheathing membrane, as in Woodbine. The colour of the pith, in the succulent shoot or in the young plant, is green, which, as the cells empty, changes to white; but to this there are some exceptions. Thus it is yellow in the Bar- berry, Berberis vulgaris ; pale brown in the Wal- nut, Juglans regia ; fawn-coloured in the Sumach, Rhus Coriaria ; and pale orange in yellow-flow- ered Horse Chesnut, yEsculus flava; but it is more frequently coloured in the caudex of the root than in the stem. Placing a thin slice of pith, taken either from a vertical or a horizontal section of our shoot of Horse Chesnut, or of any other plant, under the microscope, it appears to consist of hexagonal cells *, which are larger and more regular in the centre than near the circumference -j~. In very young stems and succulent shoots, these cells are filled with an aqueous fluid, and closely resemble the cellular integument % ; but, in older stems and twigs, they are found empty, or, more accurately * The cellular structure of the pith was first pointed out by Grew in his work on the Anatomy of Plants, fol. 1682. t Vide Plate 6, fig. 2, k. J Vide Plate 6, fig. 2, c. LECT. VfJ.J ANATOMY OF STEMS. 371 speaking, filled with air. The cells retain the hex- agonal form in their empty state ; but in some, as the Walnut, this is destroyed in the lamellae, into which the pith then separates; and the same oc- curs in the interior of the medullary sheath of Woodbine, and similar hollow stems. In the' greater number of plants no vessels are percep- tible in the pith ; but in some, entire vessels con- veying proper juice are present, as in the Gum- elastic Fig tree, the proper juice of which is seen exuding from different points of the pith in a hori- zontal section of the stem : and, in all plants, the cells communicate with each other by means of or- ganized pores, which are visible under the mi- croscope. M. Aubert Du Petit-Thouars has lately affected to regard the medulla as deserving that name only after the cells become empty, naming it parenchyma in the early or succulent stage of its existence * : but this is at best a useless refinement ; for, although I am not prepared to admit, with Mr. Keith -f-, that there is an essential difference be- tween the membrane composing the cells of the parenchyma or pulp, and that forming those of the pith, yet the insulated and enclosed situation of the pith is sufficient to obtain for it the dignity of * " Ce n'est que par 1'extraction des sues qu'elle (la Mo- " elle) contient qu'elle est devenue Moelle." Essais stir la Vegetation, p. 205. f System of phys. Bot. vol. i. p. 324. B B 2 372 CONSERVATIVE ORGANS. [LECT. VII. being a distinct organ in every stage of its ex- istence. To enter at present upon the explanation of the formation of the pith, would be anticipating our inquiries into the general theory of vegetation, a part of our subject for the discussion of which we are not yet fully prepared. Regarding, how- ever, the mere mechanical causes which possibly operate in producing the hexagonal form of the cells, I may remark that, if the ,gelatinons pulp, which constitutes the earliest state of the pith, as- sumes the form of globules as the first effect which the operation of the vital organizing influence pro- duces on it ; it is easy to conceive that, from the individual inflation of these occasioning them to press in every direction upon one another, within a certain limit, each globule will necessarily acquire an hexagonal form. In this state the flat surfaces of the enclosing membrane of each globule which are in immediate contact, uniting and acquiring firmness, while the contained fluid is dissipated and air admitted, the cavity it occupied will re- main as an hexagonal cell ; and of such is the dry pith constituted. To produce this effect, how- ever, the following circumstances are necessary: 1. The membrane which divides each cell from those adjoining it, must be double, which Link * * Secunda Dissert, in Homer's Collect. Bot. fascic. i. p. 163. LECT. VII.] ANATOMY OF STEMS. 373 and Kieser * have demonstrated to be the case-f-. 2. This membrane must be cribriform, such as I have described it, when seen under the microscope, to permit the aqueous fluid to pass readily through it, and to admit the air into the cells to supply its place. But it is necessary to remark here, that this fact, although supported by the observations of Sprengel, Mirbel, and others, is positively de- nied both by Link and Kieser ; but, as it was re- quisite for those who denied the porosity of the membrane to state how fluids can be transmitted from one cell to another, Link has advanced the unphilosophical suggestion, that this is effected by a double filtration through invisible pores: and the ingenious Mr. Ellis, who gives credit to Kieser's observations, is forced to admit that this filtration can be accomplished, " consistently with the in- " tegrity of the cellular texture, only by the ex- " ercise of the alternate functions of secretion and " absorption J." 3. The mass must be free from any external pressure, which would, eventually, destroy the regular form of the cell. It is not easy to conjecture by what means * Mem. sur I' Organization des Plantes, p. 91. f We may also regard as an analogical proof a fact lately ascertained by Dr. Barclay, that each side of every cell in the honeycomb is double, or composed of two plates of wax. Vide Wernerian Trans, vol. ii. J Supplement to the Encyclop. Brit. vol. i. Part 2. BB3 374 CONSERVATIVE ORGANS. [jLECT. VII. air is introduced into hollow ligneous stems ; but, probably, in those in which the pith is sheathing, as in the Woodbine, the union between the utricles may be less intimate in the centre, and the air insinuating itself between them, while the cells are emptying themselves, compresses their sides together and separates them ; and extending itself in the length of the stem, forms it into a hollow tube. In others, as the Walnut, it may be intro- duced laterally, in separate quantities, at dif- ferent points at the same time, and these dilating, compress the horizontal portions of the emptying cells between them, so as to produce the medul- lary plates, which characterize that and similar stems. Phytologists have not differed more on any subject than in assigning the functions of the pith. When we reflect that Caesalpinus, who lived in the sixteenth century, taught that the pith is less essential to the life and growth of a tree than the bark *, it is astonishing that writers of so re- cent a period as Dr. Darwin and Sir J. E. Smith, should be found maintaining the accuracy of the ancient opinion, that the pith is to the vegetable what the brain and spinal marrow are to the animal body-}-. Darwin's imagination led him * De Plantis. Flor. 1583. f Such was also the opinion of Linnaeus, Amcen. dead. vol. iv. p. 372. LBCT. VI 1.] ANATOMY OF STEMS. 375 even so far as to believe that it furnishes "the " power of motion, as well as of sensation to the " various parts of the vegetable system *." Hales and Linnaeus also regarded it as the seat of the vital energy of the plant -f~ ; and Mr. Lindsay, of Jamaica, " thought he demonstrated the medulla " in the leaf-stalk of the Mimosa puclica, or Sen- " sitive Plant, to be the seat of irritability ;" nor " can I," says Sir J. E. Smith, " see any thing " to invalidate the opinion J." It would, never- theless, be no difficult task to prove the unte- nable nature of these doctrines, were they not com- pletely destroyed by the experiment of Mr. Knight, who abstracted more than an inch of the pith from the shoot of a vine, above and below a leaf and bud ; both of which, " with the lateral shoot an- " nexed, continued to live, and did not appear to " suffer much inconvenience ; but faded a little " when the sun shone strongly upon them ." Now the life of the shoot, even admitting with Darwin, for the sake of argument, that each bud has a sen- * Phytologia, xviii. 2, 13. f Linnaeus, like Darwin, compared it to the spinal marrow. J Introd. to phys. and syst. Bot. chap. vii. Phil. Trans. 1801, p. 338. Mr. Keith gives a quotation from Theophrastus to show " that this experiment had been performed, and the result as- " certained," even in the time of that naturalist. System of phys. Bot. vol. ii. p. 211. B B 4 376 CONSERVATIVE ORGANS. [LECT. VII. sorium of its own ||, could not be supported un- der such a circumstance, did any analogy exist between the medulla of plants and the spinal marrow of animals. Grew and Malpighi believed that the functions P a P aver somniferum, nearly the whole of the cells are oblong, or rather tubular ; and are ar- ranged with great symmetry, in the manner figured at a. (see marginal cut). A few cells very highly magnified are fi- gured at b., chiefly to show the appearance of the lateral fur- rows or transverse slits, which in the root of the Gourd, I am disposed to believe that, the simple spiral is rarely, if ever, found in the annual root. 3 LECT. V1IJ.] STRUCTURE OF HERBACEOUS ROOTS. 433 are very conspicuous in these cells. I have not been able to detect any fasciculi of proper vessels in roots of this description, and I am, therefore, disposed to believe that these tubular cells are not only the reservoirs, but the conductors of the proper juice. That they are endued with con- tractility, and communicate freely with each other, is evident: for, by making a horizontal section of the root, the exudation of juice is much greater than can be contained in the range of cells which is divided ; and, by placing a longitudinal slice of the root under the microscope, we find the cells of several successive ranges empty and shrunk. That this longitudinal communication is regulated by valves, or something of a similar nature, is probable; for, the exudation of the juice is much more considerable on the divided surface, and the shrinking of the cells extends to more distant ranges, in the portion of the divided root which remains attached to the plant, than in that which is separated from it. The disposition of the parts of the root is more varied in biennial than in annual roots. Taking the root of Burdock, Arctium lappa, as an ex- ample, we find that, in the first season of its growth, or in the seminal plant, it consists of a thick cellular bark, the cells of which are irre- gular hexagons arranged in concentric circles around a large central part, which is composed VOL. I. F F 434 CONSERVATIVE ORGANS. [LECT. VIII. chiefly of oblong cells, or rather short hexa- gonal tubes. These tubes in the transverse section appear arranged in beautiful rays proceeding from the centre to the circumference, and, in the lon- gitudinal section, show nearly the same symme- trical ranks that have been described as exist- ing in the Poppy root. The sap vessels are com- paratively few in number, and are arranged in rays through the central part. They are larger than the cellular short tubes, which are condensed in the line of each ray of sap vessels, so as to produce a very beautiful appearance in a transverse slice of the root examined under the microscope. All the sap vessels are punctuated, but I have not been able to satisfy myself that they are spiral. In the second year, a new circle of short tubes is formed with Some additional sap vessels interspersed through it, both preserving the radiated arrange- ment ; the old bark appears lacerated, shrivelled, and pushed outwards, whilst the space betwixt it and the new central matter, is filled up with fresh cortical cells. The short tubes appear to be the principal re- servoirs of the mucus, with which this root abounds ; but it is present in the cortical cells also, a lateral communication existing between these and the hexagonal tubes. It is apparently in- tended for advancing the fructification of the plant, being gradually absorbed as that process is LECT. VIII.] STRUCTURE OF HERBACEOUS ROOTS. 435 perfected; at which time the short tubes are emptied, and their sides gradually become ligne- ous and opaque. The root of the Carrot, Daucus carota, re- sembles that of the Burdock in general structure, but the sap vessels are comparatively more nu- merous, and the cellular rays more condensed. The cells and short tubes are rather four-sided than hexagonal; and there is no symmetrical ar- rangement of the latter. The sap vessels are all punctuated, even the smallest and most recently formed, which militates against Kieser's opinion of transformation. In the root of Hemlock, Conium maculatum, the sap vessels are situated chiefly in the centre, in fasciculi, interspersed with cellular matter, dis- posed in narrow wedgelike masses, divided by more condensed cellular matter, closely resem- bling the divergent rays in the roots of trees. The cortex is thick, and contains various fasciculi of proper vessels, disposed at regular distances, so as to form a kind of double circle. The sap vessels are punctuated spirals. Such is the structure of these three biennial roots. The diversity they display is sufficient to demonstrate the variations in the position of the parts, which occur in biennial herbaceous roots. In all of them the proper vessels constitute the greater part of the bulk of the root ; and appear FF2 436 CONSERVATIVE ORGANS. [LECT. VIU. to be chiefly reservoirs of the proper juice formed by the first year's foliage, which is expended in the formation and perfecting of the flower-stem and the fructification, the productions of the second year. As the growth of the flower-stem, therefore, and the evolution of the flower advance, the root, instead of increasing in bulk, gradually shrivels, and becomes of a more ligneous texture, owing to the absorption of the proper juice and to the emptied state of the short tubes. On this account we find that those biennial roots, the Carrot and Turnep for example, in which the art of cultivation has so much increased the deposition of nutritious matter, as to render them important as articles of food to man and other animals, cease to be fit for this purpose very soon after the flowers of the plants to which they belong make their appearance. The perennial herbaceous roots are still more varied in structure, as far as regards the proportion and arrangement of the vascular parts, than either the annual or the biennial. It would be impossible in this place to demonstrate even a very small pro- portion of those diversities : and these roots have been too little examined to admit of any classifi- cation founded on structure. I shall venture, how- ever, to arrange them into two classes, the first comprehending those which, besides sap and pro- per vessels, are composed chiefly of short tubular LECT. VIII.] STRUCTURE OF HERBACEOUS ROOTS. 437 cells, which appear symmetrically arranged in the longitudinal section of the root: the second, those which consist of sap and proper vessels, and com- mon cellular matter only, exhibiting no peculiar symmetry of arrangement in the longitudinal sec- tion. 1. Perennial herbaceous roots composed chiefly of tubular cells symmetrically arranged. As examples of this class, we may select the roots of Dandelion and of Marsh Mallow; because the principal secretion in the one is an opaque, white, glutinous fluid, and that in the other a transparent colourless mucus. If we place a transverse and a longitudinal slice of the root of Dandelion, Leontodon Taraxacum, in the first year of its growth, un- der the microscope, it appears composed of a cellular pith a. (see marginal cut), surrounded by ten fasciculi of sap vessels b. ; and a very thick cortex, which consists, interiorly, of a concentric layer of smaller cells arranged in rays, through which run numerous fasciculi of proper vessels, c. arranged so as to form three concentric circles ; and exteriorly of a mass of hexagonal cells, the same as the pith covered with a thick cuticle, d. The sap vessels are punctuated, the perforations being oblong trans- FF3 438 CONSERVATIVE ORGANS. [>ECT. VIII. verse slits; and the radiated cells are tubular and arranged in symmetrical order. The cha- racter of the proper vessels can scarcely be made out, owing to their transparency and the white juice with which they are filled; but they are evi- dently perforated and communicate laterally with the tubular cells, into which the juice they convey is filtered, to be preserved for the purposes of the plant. This juice probably undergoes some change in its passage, as it appears to be more pel- lucid in the tubular cells, than in the proper vessels. As the root advances in age, additional vessels are added to the fasciculi of sap vessels, until the whole of the central part of the stem is nearly oc- cupied with them ; and, the original cellular matter being closely compressed between the fasciculi, the vascular portion assumes the aspect of one large fasciculus. Vessels are added exteriorly also ; but these are much fewer in number, although they are larger and more distinct than those within the original circle. In the same manner a new layer of cellular tubes and of proper vessels is an- nually added to the bark, so that a transverse section of an old root appears to the naked eye to consist of an opaque woody central part sur- rounded with concentric circles, alternately opaque and transparent. The old cuticle, with a portion of cellular matter adhering to it, is annu- ally pushed outwards, as in the trunks of trees, LECT. VIII.] STRUCTURE OF HERBACEOUS ROOTS. 439 and its place is supplied by a new one ; but in this plant the cellular matter, which is found situated immediately under the cuticle in the young*, is deficient in the old root. The transverse section of the root of Marsh Mallow, Althaea officinalis, displays a kind of pith composed of one large fasciculus of sap vessels in the centre, surrounded by a mass of cells ar- ranged in rays, having a few small fasciculi of sap vessels dispersed through it; and several larger on its verge, forming an interrupted circle round it. The cortex is thick, cellular, and contains clus- ters of proper vessels arranged in a radiated form* In the longitudinal section we find that the sap vessels are punctuated, but not spiral; and the tubular cells are comparatively much shorter and wider than those in the root of Dandelion ; a cir- cumstance which appears almost to be essential, when we consider the nature of the mucilaginous secretion deposited in them. In both these roots the central part readily se- parates from the cortex; and, except in very young roots, it is from the latter only that we can extract the secretions on which their value as me- dicinal agents depends. 2. Perennial herbaceous roots composed chiefly of common cellular matter. The root of Deadly Nightshade, Atropa Belladonna, may be taken as FF4 440 CONSERVATIVE ORGANS. [LECT. VIII. an example of the general structure of this di- vision. It is composed of a central part and a very thick bark. Placing a transverse section of the young root under the microscope, we find that the central part consists chiefly of cellular matter postured in a radiated manner, with one large fasciculus of sap vessels in the centre, and several smaller fasciculi interspersed through it, forming nine or ten indistinct, interrupted rays. In the root of the second year, the additional sap vessels appear as an interrupted circle bounding the cen- tral part; and in older roots, as new circles are annually added, the transverse section resembles, in some degree, that of a ligneous stem ; or it ap- pears to consist of a pith, concentric circles of wood traversed by divergent rays, and a bark. In the longitudinal section we perceive that the sap vessels are punctuated, but certainly not spiral; and that the cells are oblong, but not arranged in the symmetrical manner, which characterizes the former division of the roots under examination. Decorticating the root, we perceive, also, that the fasciculi of sap vessels do not run in straight lines, but take a waving course; and by vessels separat- ing from one fasciculus coming in con- tact with those separated from another, the whole appears like a reticulated tex- ture on the surface of the central part (see marginal cut); a circumstance, however, which is not peculiar to this root, but is LECT. VIII.] STRUCTURE OF HERBACEOUS ROOTS. 44l general to all herbaceous roots in which the cen- tral part consists of rather more cellular than vascular matter. Such is the general structure of herbaceous roots. The main caudex in every instance is more or less of a spindle shape ; but it frequently be- comes forked near the apex, in which case there is a separation of the sap vessels, in the same man- ner as occurs in dividing a skein of threads into two or three parcels; each fork of the root con- taining a portion of the vessels belonging to the main body of the root. In the lateral branches or rootlets, however, this is not the case. These are generally given off at right angles with the main root; and each is composed of one large fasciculus of sap vessels enclosed in a cellular cortex ; but very few of these vessels are given off from those of the caudex; the majority being new vessels ge- nerated in the puncta vitalia; in which the rootlets originate. Whether these new vessels anastomose (using the expression as it is employed in speaking of animal vessels) with the vessels of the main root, I have not yet been able to satisfy myself; they are evidently closely applied to and lost on the surface of the fasciculus of the caudex whence they originate, and pour their contents into it; but this might be effected without an anastomosis, by the lateral transmission through the punctures in the coats of the vessels. 442 CONSERVATIVE ORGANS. [LECT, VIII. In general, the lateral rootlets extend in the direction of their axis, and display an uniform structure throughout their length ; but this is oc- casionally altered by the nature of the soil. Thus, on the root of the common Sweet-pea, Lathyrus odoratus, when cultivated in a dry soil, we fre- quently find that the rootlets, instead of extending as fibres, swell and assume all the external charac- ters of knobs; which, however, differ from tubers in being merely reservoirs of nutriment, without containing the germs of future buds. When one of these is dissected, the vessels are seen to origin- ate as in the other rootlets, but they soon divide, and embrace the cellular mass which contains the nutritious matter, of which the knob is the reservoir (see marginal cut, in which a. represents the knob, and b. b. b. the natural radicles). The explanation of the formation of these occasional knobs on herbaceous roots, advanced by Sir J. E. Smith, does not now appear to me so satisfactory as I formerly supposed it to be (page 206) ; for I find that they appear on the roots when " no sudden fresh * supply of food is furnished." It is probable, also, that such a supply, if made to a half-starved plant, the roots of which as yet display no knobs, would occasion the na- tural extension of the rootlets, instead of producing LECT. VIII.] STRUCTURE OF HERBACEOUS ROOTS. 443 knobs. It appears to me, that when an herba- ceous plant is partially stinted of moisture and nutriment in the soil, the functions of some of the rootlets are destroyed, owing to the sap vessels losing their irritability and becoming obstructed; while at the same time the absorbing orifices of the cuticle become also impermeable. But as the other rootlets still continue in a natural state, the plant is kept alive; and as the proper juices are se- creted, the usual supply is sent to the affected rootlets, which, however, not being able to assi- milate it, the cellular matter in which it is depo- sited swells, and the rootlets assume the cha- racter of the knobs in question *. From the examination of the structure of her- baceous roots, which we have just concluded, se- veral practical hints may be obtained. Thus it ap- pears probable, that those roots, containing bland saccharine or amylaceous secretions, which possess a moist cellular central part, are the best adapted for rewarding the skill of the horticulturist, in converting them into articles of food for man and other animals. We also discover the reason that * Mr. Keith says, " This anomaly seems to be merely the " result of a provision of Nature, by which the plant is en- " dowed with the capacity of collecting a supply of moisture " suited to existing circumstances, and hence of adapting itself to the soil in which it grows." Syst. of Phys. Bot. vol. ii. p. 271. 444 CONSERVATIVE ORGANS. [*,ECT. VIII. the central parts of some medicinal roots are as valuable as the cortical; although, in general, the bark is that part of the root which is most richly stored with the proper juices of the plant. In examining unknown roots, when we find the central part woody, we may always conclude that it consists entirely of sap vessels, and therefore is useless either as food or as medicine; but when it is composed chiefly of a moist, cellular substance, we may expect to find it useful, from having some of the proper juice and secretions of the plant de- posited in it. The ignorance of this fact, led Pharmacopolists, until very lately, to expend un- necessarily much labour and time in preparing several vegetable decoctions and extracts, as, for example, the decoction of sarsaparilla ; for the preparation of which the root underwent long macerations and much boiling, from an idea that its virtues were contained in the ligneous central part ; whereas the saponaceous mucus for which it is valued, is deposited solely in the cortical part, and can be entirely extracted by cold water. In selecting the proper period for digging up roots for medicinal purposes, it ought to be re- collected that, as the proper juices which are stored up in the roots of such plants as produce leaves only in one year, and then flower and die in the second, are expended in the process of fructi- fication ; biennial roots should be taken up at the LECT. VIII.] STRUCTURE OF HERBACEOUS ROOTS. 445 end of the first season of their growth, for then the cells are turgid with the secretions. Perennial roots, also, should be dug up before the central part becomes ligneous.; for, as these roots increase in diameter by annual additions to both their cen- tral and cortical parts, in the same manner as the stems of trees, the interior of the central part be- comes every year more and more inert, and ul- timately decays; so that, in employing such roots, when old, even before they decay, the active principle they contain becomes too largely diluted with the inert matter to answer the purposes expected from them as drugs. In closing our researches into this part of Phy- totomy, the anatomy of stems and roots, I may observe that the subject has, hitherto, been imperfectly examined; although the fieJd of in- vestigation is very extensive, and the harvest it contains calculated to repay amply the toil of the most assiduous labourer. The improvements which are daily making in the construction of the micro- scope, are likely soon to set aside all the obstacles depending on the minuteness of the parts; and a very little experience is sufficient to make the stu- dent 'expert in the use of this instrument. The best authors to be consulted on the subject are Grew, Anatomy of Plants; Malpighi, Anatome Plantarum; Rudolphi, Anatomic der Pflanzen; Kie&er, Me*moire sur FOrganisation des Plantes; 446 CONSERVATIVE ORGANS. [iJSCT. VIII. Mirbel, ElSmens de Physiologic v6g6tale; Corn- par etti, Prodromo de Fisica vegetabile; Du Hamel, La Physique des Arbres; Hill on the Construction of Timber; Krocker, Diss. de Plan- tarum Epidermide; Bauer, Tracts relative to Botany, London, 1809; Bolimer de Vegetabilium celluloso Contextu ; Reichel de Vasis Plantarum spiralibus ; Histoire d'un Morceau de Bois, &c. par A. A. du Petit Thouars; Keith's System of phy- siological Botany ; Supplement to the Encyclo- paedia Britannica ; and Mr. Knight's papers in the Philosophical Transactions. LECT. IX.] HYBERNACULA. 447 LECTURE IX. OF LEAVES IN THEIR UNEXPANDED STATE, OR AS THEY ARE CONTAINED IN THE GEM: IN THEIR EXPANDED STATE, OR AS CONSTITUTING FOLIAGE. HAVING concluded our examination of roots, stems, and branches, we are now prepared to in- vestigate the structure of leaves. In winter, while the power of vegetation is inactive, and the groves and forests present the desolate appearance of naked stems and branches, the majority of trees, shrubs, and many other plants may be regarded as existing in a state of torpor, similar to that which some animals expe- rience in the same season. During this period the leaves are enclosed in small pyramidal bodies, either projecting from the surface of the stem and branches, or seated upon the roots; and in this state they remain until the warmth of the vernal sun, again rousing into action the vegetable func- tions, enables them to burst open their coverings, and clothe the woods anew in all the luxuriance of foliage. In our examination of leaves, therefore, we must regard them both as they are shut up in these 448 CONSERVATIVE ORGANS. [LECT. IX. hybernacula, or winter habitations, and in their expanded state. The pyramidal bodies which I have just no- ticed are well known by the name of buds; and the appellation is so universally applied in this country in reference to their appearance on trees in early spring, that it would be pedantic to re- ject it ;. besides, I have already used it in speaking of the origin of branches. But as the term bud is also employed to denote the separate flower before it blows, and as the purposes of science are better attained by using a word applicable to the object only which it is intended to present to the mind, I shall employ the term hybernaculum as syno- nimous with bud, in treating of this part of our subject. A HYBERNACULUM may be defined : an organic body which sprouts from the surface of different parts of a plant, enclosing the rudiments of the new shoot ; and which is capable of evolving a new individual perfectly similar to the parent. This is a modification of the definition of Gsertner *, which * " An organic body generally sprouting from the surface " of a plant, without previous fecundation ; in the beginning " distinct from the peculiar and permanent membranes of the " plant ; but which, in a certain time, either becomes a part " of the parent, or separates from it, and by the increase of its " own substance becomes a new plant, closely resembling the " parent." Gartner de Fructibus Plant, p. 3. LECT. IX.] HYBERNACULA. 449 is objectionable only in expressing an opinion, as to the early state of the bud, that may be dis- puted*. Some hybernacula remain attached to the parent ; others detach themselves after a cer- tain period, but both kinds are to be regarded as the lateral progeny of the plant ; for even that which remains attached possesses, in a certain de- gree, a separate or distinct vitality, by which it is enabled to exist when forcibly detached from the parent. Thus a bud taken from a tree, and pro- perly planted in the ground, covered with a glass to prevent too great an exhalation of its natural moisture, will grow and become a tree resembling, in every respect, that from which it was taken. But still the plant, whether raised from a bud thus forcibly detached, or from one which na- turally detaches itself, is an extension only of the parent, displaying all its individual peculiarities the effects of soil or culture, and inheriting all its diseases; whereas a plant raised from a seed is a new individual, displaying the generic and specific characters only of the parent. From the same cause, also, plants which are natives of a warm climate, when taken to a colder, and pro- * " Hylernaculum est pars plantae includens herbam em- " bryonem ab externis injuriis." Phil. Bot. 85. The ad* mirable simplicity which characterizes all Linnaeus 's defini- tions is here conspicuous ; but the definition is objectionable in being equally applicable to the seed as to the hybernaculum. VOL. I. G G 450 CONSERVATIVE ORGANS. [t,ECT. IX. pagated by slips or buds, never become so com- pletely naturalized as to bear all the variations of the new climate with impunity; but plants which are propagated by seeds, although natives of very warm climates, yet, become perfectly na- turalized to colder in a certain number of genera- tions. Botanists enumerate three kinds of sponta- neously separating Hybernacula, the Propago, the Gongylus, and the Bulb; and one which does not spontaneously separate from the parent, the Gem. The PROPAGO * is thus denned by Gaertner : " A simple leafless, polymorphous, or variously " shaped germ, in some instances naked, in others " enclosed in a cortical sheath^ which sponta- " neously separates from the parent, and is scat- " tered in the manner of seed." It is a small pulpy or cellular body of no regular shape ; and is sometimes covered with an epidermis. It is readily found in dividing the tubercles and shields or saucerlike bodies, which appear on the surfaces of Lichens, in an early stage of their * This term was used by the ancients chiefly to denote a cutting of the Vine, when buried in the ground to throw up new shoots ; but it was applied also to cuttings of other plants. Arbores aut semine proveniunt, aut plantata radice, aut propa- gine, aut avulsione, aut surculo, aut insito et consecto arboris trunco. Pliny, 1. 17. c. 10. LECT. IX.] HYBERNACULA. 451 growth. According to Gaertner all the Algee are propagated by the propago, and not by seeds as Hedwig and other phytologists have asserted. With such names as contending authorities on this subject, may we venture to suggest, that al- though the sexual organs of these plants have not yet been discovered, and although they throw off the propago as lateral progeny, yet they may, also, produce real seeds ? Many of the more per- fect plants are propagated in both ways ; and we know that this occurs even in the animal kingdom ; for the aphis which is first propagated by sexual intercourse continues its species, through several successive generations, by lateral offsets. " The GONGYLUS," according to Gaertner, " is " a simple, leafless, somewhat globular, solid " germ attached to the parent under the bark, " and separating spontaneously from it." He ob- serves that it has a close affinity to the tubers found on roots; but differs inasmuch as the tu- ber possesses as it were a multiplied life, so that it may be divided into as many pieces as there are foliaceous gems on its surface, from each of which a new plant will arise. The gongylus consists of cellular matter like the propago, but of a much firmer and more solid consistence, and is always covered with an epidermis. Gaertner supposes that the Fungi, or Mushroom tribes, are altogether pro- pagated by gongyli : but Michelius, Hedwig, and G G 2 452 CONSERVATIVE ORGANS. [LECT. IX. Bulliard detected their seeds ; and there is every reason for believing that these and many of the other tribes of lower plants produce both seminal and lateral progeny. The BULB having been already denned (p. 164), and also described as it appears attached to the roots of plants ; we have now to examine it only as it appears upon the stem *. It is found on the stems of several species of the Lily, on that of bul- biferous Coralwort, Dentaria bulbifera, and of drooping Saxifrage, Saxifraga cernua, &c.*f- seated in .the axilla of the leaves (Plate 4, fig. 9, a. a.). If we take the Tiger Lily, Lilium tigrinum, as an example, we shall find the bulbs appearing like a white speck in the axilla of the leaves long- before these expand. A few days, however, after the expansion of the leaf the bulb assumes a py- ramidal form, which gradually enlarging and swelling in the centre, at length appears of an ovate or nearly globular shape, with a keel ter- minating in a point. During this transformation * Mirbel denominates the caulinar bulb, Bulbille, Bulbillus, and thus defines it : " Petite bulbe qui nait sur differentes par- " ties de la plante hors de terre, se detache et prend racine." Element de Phys. veg. Partie 2, p. 634. I Sir J. E. Smith says he has seen bulbs form on the flower- stalk of three-coloured Lachenalia, Lachenalia tricolor, whilst lying for many weeks between paper to dry ; and these on being put into the ground have become perfect plants, though of slow growth. Infrocl, to Botany, p. 112, note. LECT. IX.] HYBERNACULA CAULINAR BULBS. 453 it also acquires colour on that part of the surface exposed to the light; passing first from white to green,, then to light brown, and lastly to a very deep shining jet brown. On attaining this degree of maturity it separates from the stem on the slightest touch, or it spontaneously loses its hold ; and, dropping to the ground and vegetating, throws out roots and acquires all the characters of a root bulb. Linnaeus, nevertheless, regarded the caulinar bulb as a gem, and denominated it Gem- ma decidua* ; but, independent of its spontane- ously separating from the parent, which the real gem never does, it differs in other essential cha- racters from the gem. Examining one of these bulbs in a mature state, we find at its lower part a depression re- sembling the hilum or scar, which on a seed points out the place of its attachment in the seed- vessel. It consists, in the bulb, of a depression enclosing three elevated points, which indicate the place where the vessels connecting it with the parent entered. The bulb itself consists of two outer scales, the uppermost large and embracing the lowermost, which projecting forms the keel of the bulb, and embraces another scale within it, which in its turn embraces a fourth, and so on to * Species Gemmarum variae sunt, Deciduce in Dentaria, Ornithogalo, Lilio, Saxifraga. Phil. Bot. 85. G G3 454 CONSERVATIVE ORGANS. [LECT. IX. the germ or embryon. This struc- ture is illustrated in the marginal cut, which displays a transverse section of the bulb very consider- ably magnified : a. is the upper- most or largest scale., composed of a mass of cellular matter enclosed in a cuticle, with seven fasciculi of vessels, as marked by the dots where they are divided, running through it ; b. the second or lowermost, or keel scale, with its vessels; and within it are the third, fourth, and fifth scales, each embracing the one within it. The whole of the cells are filled with minute amylace- ous granules, mingled in a clear, viscid mucilage ; the opacity of the fluid being greater in the outer scales and diminishing in a direct ratio as these approach the centre of the bulb. Each scale is covered with a beautiful, readily separating epider- mis. Making a longitudinal section of the stem of the Lily, so as to divide a bulb, in situ, directly through its axis, we find that the fasciculus of vessels a. (see marginal cut) which nourishes it, and which may be regarded as its um- bilical cord, is a portion of the bundle b. which is given off to supply the leaf. As it ap- LECT. IX.] HYBERNACULA CA.ULINAR BULBS. 455 preaches the bulb, it divides into three fasciculi, one of which enters the upper scale c. where it splits to form the seven fasciculi that run through it, as has been already demonstrated ; another passes into the keel scale d. and the third subdi- vides to supply the interior scales and the em- bryon. The conducting vessels are all simple spirals; but I have not been able to determine the character of the returning vessels. When the bulb is perfected it separates from the stem, as has been already mentioned, in the same manner as ripe fruit falls, the umbilical vessels dividing at the point of attachment. Caulinar bulbs when planted produce leaves only, like those formed on the separated scales, or raised from the seeds of the Lily; and like these, also, they exhibit all the peculiarities and diseases of the parent. From the analogy which we recognise between the sexual progeny of animals, and the seminal progeny of vegetables, we are accustomed to re- gard the distinct vitality of seeds and their reten- tion of 1 life as a matter of course; and even per- suade ourselves that we comprehend the manner in which the principle of vitality is conveyed to the embryon ; but, although bulbs resemble seeds in many respects, yet as they are not the result of the sexual functions, we find much difficulty in comprehending how they acquire and maintain G o 4 45 6 CONSERVATIVE ORGANS. [LECT. IX. that individual vitality, which is necessary for pre- serving them in a state fit for vegetating, after their separation from the parent, and before they are planted in the ground. It would be anticipating the arguments I have to advance on the causes of vegetable reproduction, were I to enter largely into the consideration of this subject at present; but I may observe that, whether the progeny be direct or lateral, a certain organization, whatever that may be, peculiar to the species to which the indi- vidual belongs, is requisite for retaining the vital principle in conjunction with matter, and this is found in the bulb as well as in the seed. In both, the embryon is to be regarded not as a part only, but as a compendium of the whole of the parent; and the organization is so complete in every part, that the separation from the parent effects no change in it; and, consequently, as long as no change occurs, both the bulb and the seed continue fitted for commencing the vegetating process, when placed under circumstances favourable for that event. But it may be argued that a small portion of a polypus will increase and become a perfect animal, and twigs of the Willow, the Vine, and of some other plants, after being separated for a considerable time from the parent, will vegetate if thrust into the ground and left there. We know too little of the nature of polypi, to explain the multiplication of the animal from cuttings, but it is probable that the vegetation of the twigs re- LECT. IX.] HYBERNACULA CAULINAR BULBS. 457 ferred to, depends on the organization of the germs contained in the buds of these plants approaching to that degree of perfection which is found in bulbs and seeds. It may, however, still be demanded how is this perfect organization accomplished in bulbs ? In seeds, while in the state of ovula, the peculiar stimulus of the pollen may produce a specific action capable of evolving all the parts of that peculiar structure, with "which we find vitality connected; but in bulbs we can scarcely sup- pose that the vital action which completes their or- ganization differs from that by which a gem, or branch bud, is formed ; and yet buds, when sepa- rated from the parent stem, will not live, unless they are either immediately planted, or inserted into another stock of a structure resembling that of the parent. A question thence occurs (admit- ting that all plants which throw off lateral pro- geny as bulbs, propagines, and gongyli, possess sexual organs, and, therefore, are capable of being also propagated in a direct way), namely, can any of the impulse communicated by the sexual func- tions influence the lateral progeny? That such an influence exists is probable, if it be allowable to reason analogically and refer to the animal kingdom; for we find that from the egg of the aphis, which is laid in the autumn, and is the re- sult of the sexual intercourse of males and fe- males, a young insect is produced in the spring ; which after casting its skin once or twice, pro- I 458 CONSERVATIVE ORGANS. [LECT. IX. duces a living progeny without sexual intercourse ; and " this offspring produces others by this solitary "propagation, till the tenth generation; then a " sexual progeny of males and females is produced," and eggs are laid from their copulation*. Now, although no experiments have been made to ascer- tain how long a bulbiferous plant may be propa- gated by bulbs only, checking every effort for the production of seed, yet, we know that in bul- biferous plants, when the production of bulbs is considerable, the seeds are seldom ripened, and even the sexual organs are often defective; and the reverse happens when the production of bulbs is either scanty or defective. The analogy between the successive lateral generations of the aphis, before males and females are formed to recom- mence the propagation by eggs, and the pro- gressive formation of bulbs before a flower-bulb is the result, is still more striking. The bulb, which is raised from the seed, produces one or two leaves only, and bulbs one degree more perfect than it- self; which in their turn yield stronger plants and more perfect bulbs; and in this manner a suc- cession of leaf-bulbs are annually evolved for four or five years, till at length a flower-bulb and a seminal progeny are produced. The vegetable, however, differs from the animal in producing, at the same time with the flower-bulb, other * Amcenit. Academ. vol. vii. Darwin's Phytologia, ix, 3, 1. LECT. IX.] HYBERNACULA CAULINAR BULBS. 459 leaf-bulbs also; so as still to secure its preserva- tion should its flowers be accidentally destroyed. Such are the separating Hybernacula. Al- though they resemble seeds in containing within them a perfect embryon, yet, like that contained in the attached buds, which we are now about to examine, it is an offset or continuation only of the parent, and not a renewal of the species. Many of these plants which have been improperly named imperfect, as for example the Confervas and Lichens, are supposed to be propagated by no other means; but as the plants thus produced dis- play not only the essential, but the accidental cha- racteristics of the parent, I am disposed to believe, as I have already stated, that the " propagation " by seed is, in their case," by no means, " out " of the question *." The attached Hybernaculum, or bud, or GEM -|~, as it is more generally termed, is a small oval or pyramidal body, enclosing the rudiments * Elements of the Philosophy of Plants, by A. P. Decan- dolle and K. Sprengel, 302, Eng. Trans. f The ancients used the terms Germen and Oculus to denote those buds which contain the rudiments of branches and leaves, and Gemma those in which flowers only are contained ; but by the moderns, Germen has been applied to denote the rudiment of the fruit ; thus, Linnaeus, Germen rudimentum fructus im- maturi in flore (Phil. Bot. $96), or as a generic term for all buds (see Gaertner de FructibusJ, while Gemma is employed exclusively to indicate caulinar buds. 460 CONSERVATIVE ORGANS. j^LECT. IX. of branches and leaves, and sometimes flowers, and never separating from the parent *. Gems are found on all trees and shrubs in temperate climates. In the majority of instances they are visible from the first, in which case they are axillary, that is, seated in the axillae of the leaves, or the angle which the upper part of the footstalk of the leaf makes with the surface of the stem ; and terminal, or at the extremities of the branches : but in some instances, for example, the Sumachs, Rhus, and the Planes, Platanus, they are latent ; being hid within the base of the footstalk, and never seen until the fall of the leaf. In the marginal cut, a. repre- sents the footstalk of a leaf of the Oriental Plane, Platanus orienfalis, split longitudinally to show the ca- yity b. in which is seated c., the gem. Gems are, however, some- times protruded from the trunk, long after it has ceased to produce * The above definition is a modification of that of Gaertner, who thus defines the gem, " a compound subulate or pyramidal " germ, with manifest herbaceous leaflets, containing the rudi- " ments of branches, and never separating from the mother." Linnaeus's definition is less correct, " Gemma est hybernaculum " caudici adscendenti insidens. Constat vel stipulis, vel " petiolis, vel foliorum rudimentis, vel squamis corticalibus/' Phil. Bot. 85, 2. LECT. IX.] HYBERNACULA GEMS. 46 1 leaves, as in the case of adventitious buds, already treated of; they are also situated on roots, and on tubers; but in these cases they are usually deno- minated eyes, oculi. Annual plants are supposed not to be furnished with gems; but although they are devoid of covered gems, yet their lateral shoots proceed from naked buds, which immediately spread into foliage. The relative position of axillary gfcms is ne- cessarily regulated by that of the leaf; and there- fore we find them, 1. opposite, or placed exactly on the same line on opposite sides of the stem or the branch; 2. alternate, or placed alternately, al- though on opposite sides; and 3. spiral, that is, placed round the stem or the branch in such a manner, that a cord wound in a spiral manner round it would touch each gem. They are said to be simple or solitary, when one gem only is seen in the axilla of each leaf, as in the greater number of in- stances ; and aggregate, when, as in some plants, two, three, and even more, are protruded at the same time: thus we find two on the common Elder, Sambucus nigra, three on broad-leaved Birth-wort, Aristolochia sipho, and on Blue-ber- ried Honeysuckle, Lonicera ccerulea, and many on common Toothach tree, Zanthoxylum fraxi- neum ; but as these are natives of cold countries, it is supposed that the intention of Nature in this double and triple supply is to secure the plants 462 CONSERVATIVE ORGANS. [LECT. ix. against the effects of frost, and other accidents of climate to which their situation exposes them. In this climate, however, one of the additional buds is always evolved the same season in which it is protruded. Du Hamel first noticed the fact, that stems and branches furnished with alternate axillary gems, have generally one terminal gem only, and those with opposite have generally three terminal gems. In the Horse Chesnut the middle terminal gem is the largest (see a. in the marginal cut), and bursts soonest into foliage; while those on its sides (b. b.) are much smaller, and sometimes never open, but decay and drop from the branch. In the Lilac, on the contrary, the middle terminal gem is always the smallest, and scarcely ever pullulates. In some trees, as for example the Pine tribe, all the gems are terminal. The gems on most trees and shrubs rise with a broad base from the surface where they are pro- truded, and, consequently, being in close contact with it, are said to be sessile (sessiles) ; but they are distant or stalked on some; as for example, the common Alder, Alnus glutwosa, on which LKCT. IX.] HYBRRNACULA GEMS. 463 they are supported on a short footstalk, and are then termed stalked (pedicillatce) . In employing these terms, however, the student must bear in mind, that they refer to the entire gem ; for a writer of great merit, in describing the gem, says, it " is connected with the stem or branch by means " of a short and fleshy pedicle, in which the " scales originate * ;" whereas this is merely the base of the young branch it encloses. The angle, also, which gems form with the stem or branch, varies considerably in different trees ; thus on the Willow they lie almost parallel to it, while on Apple and Pear trees they project so as to form nearly a right angle with it ; but by this the di- rection of the future branch is regulated. Let us detach a gem from any tree, for in- stance the Horse Chesnut, which affords the most magnificent specimen of a gem known in this country, and examine its structure. We find that it consists exteriorly of eight pairs of hollow scales, each pair consisting of scales of the same form and magnitude, placed directly opposite to each other, in such a manner as to cover the opposing edges of the pair within them. The inner scales are longer, and more tender and succulent than the outer, which are hard and covered with a viscid resinous exudation, that unites them together, and is found * Keith's Syst. of phys. Bot. vol. i. p. 65. 464 CONSERVATIVE ORGANS. [tECT. IX. on the inner scales also; but of a thinner and more transparent quality. On removing all the scales, the rudiments of the young branch and the leaves are discovered, embedded in a soft hairy or woody substance. If this examination be made, by dividing a terminal gem longitudinally, in the very early part of spring, before the buds begin to swell, we find (see Jc. fig. 15, Plate 5) the rudi- ment of the new branch apparently quite distinct from the old ; separated by a partition which, as the season advances and the scales begin to open, is gradually obliterated, while at the same time the quantity of woolly matter surrounding the leaves is greatly increased, and these acquire their determinate forms, folded up, however, so as to occupy the smallest possible space. But although the examination of any gem will afford a general idea of the structure of all, yet, gems differ very considerably in the number and characters of the enclosing scales, their contents, the folding up of the leaves within them, and the manner in which these are evolved in the spring. a. The scales which constitute, in fact, the hy- bernaculum, differ, as has been already stated, in size and texture even in the same gem ; in the gems of different plants they differ also in num- ber and in the nature of their coverings. Some gems, indeed, are entirely destitute of scales, for example, those of annual plants, and of many LKCT. IX.] HVBKKNACULA GEMS. 465 perennials of tropical climates, in which the in- terval between the formation and the evolution of the bud is so short as to require no protection for the young shoot. The scales in some instances are smeared with a resinous matter ; in others they are perfectly free from any moist exudation, but are smooth and polished, being covered with a dry gummy var- nish ; or they are externally hairy, or enveloped in a velvety down. In their organic structure, they closely resemble the scales of bulbs, being com- posed of a layer of cellular substance, enclosed in an epidermis, and containing fasciculi of ves- sels running in lines from the base to the apex. The vessels enter the scale in distinct fasciculi ; and to this arrangement is attributed the differ- ence in figure which always exists between the scale and the real leaf, into the latter of which the vessels enter in one fasciculus only, from which, as from a common centre, they are distributed through the leaf. The inner scales perform the functions of the leaves until several of these are expanded, and then they generally drop off; displaying in this respect a striking analogy to seed lobes, which, in many seeds, rise above the surface of the ground, and become green, executing all the functions of leaves until these attain a certain degree of ma- turity; after which they shrivel and fall. The VOL. I. H H 46(5 CONSERVATIVE ORGANS. [LKCT. IX. period at which the scales drop, differs in different plants; in some, as the common Lilac for instance, the more succulent inner scales remain attached until the shoot has attained to a considerable length ; whereas in the Lime tree, Tilia Europea, they drop before the leaf is fully expanded. Gems are scarcely ever formed in the axillae of the scales. Gems differ in their characters in the same family of plants * 3 and even when found on the same tree; and, as the external form of the gern indicates the nature of its contents., Botanists have arranged them into three species, leaf-gems, flower- gems, and mixed gems. 1. Leaf-gems, or buds ( Gemmce foliiferce), are long, slender, tapering, and acute, generally containing, besides leaves, the rudiments of a shoot, on which account they are also termed wood buds; and are, in truth, embryon branches^. * " Gemmae in eodem genere saepe diversissimae, uti constat " ex genere Rhamiii, ubi Cervispina, Alaternus^ Paliurus, " Frangila, gemmis diversae sunt.'' Phil. Bot. 278. f On this fact is founded the process of budding or inocula- tion, which is generally performed in July and August, and is preferred to grafting for such trees as are liable to exude much gum. To perform the operation, a transverse incision is made in the bark of the stock through to the wood ; then a longitudinal one downwards, so that the two incisions shall re- semble the letter T; and, lastly, the bark on each side of the longitudinal incision, is gently raised with a flat instrument, or the handle of the pruning-knife. The bud to be inserted should be selected from the middle of a shoot, and being separated with a slice of the bark about an inch above and below it, the LUCT. IX..] HYBERNACULA GEMS. 467 2. Flower-gems ( Gemmce ftoriferce vel fructi- feros) are short, thick, swelling and rounded at the apex. Whenever the fruit ripens, all the parts protruded from a flower-bud die, but those from a leaf-bud give a permanent addition to the tree. 3. Mixed gems (Gemmae mlxtcc vel foliif'ero- florlf'erce) are intermediate in respect to form ; but generally larger than either of the other kinds-^. The Peach tree, Arnygdalus Persica, the Mezereon, Daphne Mezere- um, and many other plants, afford examples of distinct leaf and flower gems ; the Lilac and the Horse Chesnnt of mixed gems; and Pear and Apple trees of both leaf and mixed gems. The mar- ginal cut represents the twig of a Pear tree, in which . is leaf is then to be cut off, leaving half an inch of the stalk ; and any wood that may remain attached to the bark, must be stripped off by pulling it downwards. The lower part of the bark, attached to the bud, is now to be introduced into the cross incision in the stock, and pushed downward*: and, the upper part being cut across, the bark of the bud and that of the stock are brought into close contact. A piece of bass tied round the stock over the incision is the best mode of securing the bud in its place; and on loosing it, about three weeks after the operation, if the bud appear swelled and the footstalk of the leaf drop off, the operation has succeeded. f Linnaeus enumerates seven species of gems: "Deciduae, in H H2 CONSERVATIVE: ORGANS. [LECT. ix. a leaf or branch gem ; and b. b. are gems which produce a small tuft of leaves terminated by a bunch of blossom. The information afforded by the external characters of gems is import- ant to the practical gardener, in pruning fruit- trees in winter and in early spring ; for, without it the whole of the floriferous gems might be de- stroyed, and the expectations of the cultivator al- together disappointed. But change of soil or of climate, the art of the horticulturist, and many accidents, may change one kind of gems into the other kinds. Thus a Solandra, Solandra grandi- flora, in the Kew garden, which had never flow- ered, being by an accidental neglect left without water, the too luxuriant growth of the plant was checked, and flower-buds were formed in the en- suing summer *. From the same cause, a tree newly transplanted, is often covered with blossoms, although it be nearly destitute of foliage. The leaves, as has already been mentioned, are variously folded up, so as to occupy the smallest possible space within the hybernaculum. This regulates the expansion of the leaves when / Deritraria, Ornithogalo, Lfiio, Saxifraga. Foliiferce, nonjlo- " rifercK : Alnus. Foliiferce, et Jloriferce distinct^ : Populus, " Salicis species, Fraxinus. Foliiferce et Jloriferce jeminece : " Corylus, Catfpinus. Foliiferce et jloriferce mascultB : Pinus, " Abies. Foliiferce etjlorifera hermaphrodite : Daphne, Ulmus, " Cornus, Amygdalus. Foliifero-Jlorifera:, ut pleraeque ar- 4< bores." Phil. Boi. 85. * Smith's Introduction^ p. 190. LECT. IX.] HYBKRNACULA GEMS. 469 the gem opens in spring; and it is invariably the same in individual plants of the same spe- cies. The process is termed FOLIATION *, and the figures which the leaves assume at the time, have received different appellations. In noticing these I shall arrange them under the three fol- lowing heads : Folded, jo Overlapping, and y Rolled; and mention the varieties of each kind. The best method of ascertaining the character of a gem, as respects its foliation, is to cut it across while it is opening, and to examine the sections of the leaves. oc. Folded. This kind of foliation displays the leaf- or its parts variously doubled together. There are two varieties of it : the doubled and the plaited. 1. In the doubled (condu- plicata), the two sides of the leaf lie parallel to each other, as exemplified in the Oak, the Walnut, the Cherry, the Beech, the Rose, &c. In the marginal cut, a. shows the section of a leaf, and b. the entire opening gem of the Lime tree, Tilia Europea, which is an excellent illustration of this variety. * FOLIATIO est complicatio ea, quam servant folia, dum intra Gemmam aut Asparagos plantarum latent. Phil. Bot. 163, iv. H H 3 470 CONSERVATIVE ORGANS. [LECT. IX. 2. In the plaited (Plicata), the leaf is folded up like a fan ; as ex- emplified in many of the Palm tribe; in the Birch, Betula alba, and La- dy's-Mantle, Alchemilla. In the mar- ginal cut, a. represents an unexpanded leaf of Alchemilla alpina ; b. its trans- verse section. Overlapping. Under this head are ar- ranged those gems in which the margins of the leaves overlap those within them, or opposite -to them, without being rolled. It comprehends the three following varieties. 1 . The Imbricate (Imbricata), in which the edges of two opposite leaves touch each other, embracing those within them, which they cover like tiles. In some instances the edges of the one leaf ex- tend a little over those of that to which it is opposed ; while in others the opposed edges scarcely touch. This variety is exemplified in Privet, Ligus- trum vulgar e, and Lilac, Syringa vul- garis, &c. In the marginal cut, a. represents the opening gem of Lilac, and b. its transverse section. 2. The Equitant (Equitantia), in which the leaf is so folded, that the two sides LECT. IX.] HYBERNACULA GEMS. 471 deeply embrace the opposite leaf, which in its turn encloses the one op- posed to it ; and so on to the centre of the bud. It is beautifully exemplified in Day Lily, Hemerocallis ; in the Iris family ; and in Solomon's Seal, as represented in the marginal cut, in which fig. 1. shows the character of the entire bud, with the leaf a. embraced by the opposite one b. ; and fig. 2. the transverse section in which a. b. and the other leaves are drawn slightly asunder, so as to show more distinctly their arrangement. 3. The Ob volute (Obvoluta), in which one leaf, doubled lengthways, embraces within its doubling one half of the opposite leaf, folded in the same manner ; as in the genus Valerian, Valeriana ; Scabious, Scabiosa; Teasel, Dipsacus ; and Sage, Salvia. In the marginal cut, a. represents the opening bud of Common Sage, b. its transverse sec- 'tion. Rolled. This division contains all those gems in which the leaves are rolled, either on their lateral margins, or from the apex to the, base. There are five varieties of this form of foliation. l.The convolute (Oonvoluta}, in which the leaf H H 4 7- 472 CONSERVATIVE ORGANS. [LECT. IX, is rolled lengthways in a spiral manner, one mar- gin forming the axis round which the other turns, as in the Plum genus, Prunus ; the Lettuce tribe, Lactuca ; the Cabbage, Brassica, and many Grasses, &c. In the marginal cut, g. displays this form of foliation in a section of the unrolled leaf of Indian Corn, Zea Mays. 2. The Involute (Involuta), in which each lateral margin of the leaf is rolled in- wards ; as in the gems of the Honey- suckles, Lonicerae ; and the Violets, Violae, &c. In the marginal cut, e. is a section of the unexpanded leaf of the Yellow Water Lily, Nuphar lutea. 3. The Re volute (Revoluta), in which the lateral margins are rolled as in the gems of Rosmary, Ros- marinus officinalis; and of the Primrose genus, Primula, &c. In the marginal cut, f. is a sec- tion of the unexpanded leaf of Patience Dock, Rumex patientia. 4. The Circinal (Circinata), in which the leaf ^ is rolled from the apex to the base, as in the Ferns, Filices; the divisions of the leaf, as represented at a. a., in the marginal cut, being rolled upon the mid-rib, which is also rolled from to c. carrying the divisions in its turns. 2 outwards LECT. IX.] HYBERNACULA GEMS. 473 5. The Turned down (Redinata), in which the leaf hangs down and is wrapt round the footstalk, as in the buds of officinal Wolfsbane, Aconitum neo- montanum ; the genus Anemone, &c. In the marginal cut representing an unexpanded leaf of Duck's-foot, Po- dophyllum peltatum, a. a. shows the leaf wrapped round the footstalk b. c. As the gems open, the leaves gradually unfold themselves, and assume their natural forms ; but the opening of the bud does not, in every in- stance, immediately set free the leaves; for, in some gems, each leaf is separately enclosed in a membranous cover, which opens either laterally or at the apex, and permits the leaf to expand. This covering is generally regarded as a stipule (stipula); but it scarcely accords with Linnseus's definition of that appendage* ; and may rather be considered as a protection to the embryon leaf, until it has attained sufficient vigour to bear the stimulus of light, and to admit of that degree of perspiration, which its exposure to the atmo- sphere occasions. The gem of the Tulip tree, Li- riodendron Tulipifera, affords a very beautiful ex- ample of this form of foliation. The leaf before expanding is conduplicate and arched, or bent * " Stipula est squama, quae basi petiolorum aut pedimculo- " rum enaseentium utrinque adstat." Phil. Bot. 84. 474 CONSERVATIVE ORGANS. [LECT. ix. down as represented at a. in the marginal cut ; and the sti- pules, as they are termed, which are flat foliaceous plates (see b. which is one of them separated to show its form and vessels), form a bivalve case for it, con- taining at the same time all the younger leaves, each arched and enclosed in a similar man- ner. As soon as the leaf is capable of bearing the exposure, the two plates of the case separate, bending down as at c. c. and in a short time drop off; meanwhile the footstalk of the leaf becomes straight and the disk is spread out to the light and air. In some in- stances this covering is univalvular and sepa- rates at the base, at the same time that it opens laterally, as exemplified in the Magnolias; and I very elegantly in the Elastic-gum tree, Fie us elastica; the sheath of which (see fig. 1. a. in the marginal cut) is of a deep red colour, LECT. IX.] HYBERNACULA GEMS. 475 thick, opaque, and covering the whole of the terminal shoot above the insertion d. of the last evolved leaf, the footstalk of which b. is pur- posely left in the figure : in fig. 2, the sheath is re- moved to display the leaf a. seated on its footstalk e. and wrapped round the sheath of the next ex- pected leaf. In other instances this sheath is thin, semi-transparent, and filled with a gelatinous matter, which involves the young leaf; as exem- plified in the Dock tribe, Riimex. These sheaths or utricular coverings cannot be regarded as hyber- nacula, as they are present in every season of the year; but, inasmuch as they preserve the young leaf from the stimulus of light and the effects of mois- ture, they bear a close analogy to the hybernacu- lar scales. Their chief use, however, is to re- strain the perspiration of the young leaf, till such time as its vessels are sufficiently perfect to supply by absorption the exhaustion of moisture which that function necessarily occasions. The origin of the gem has been already suffi- ciently investigated (p. 383-406). It is evidently nourished during the summer by the leaf, which is, perhaps, to the embryo in the gem, what the flower is to the fruit * ; but when the leaf falls, the gem is left to its own resources; and scarcely any visible change occurs in its aspect or its magnitude * " La feuille est au bourgeon ce que la fleur est au fruit " et a la graine. " Essais sur la Veg. par A. Aubert du Petit- Thouars, p. 14-5. 476 CONSERVATIVE ORGANS. [LECT. IX. until the following spring. During this torpidity, which is maintained throughout the winter, the scales are supposed to preserve the enclosed em- bryo from the effect of cold ; but if we reflect on the insufficiency of so feeble a guard, and further consider the great degree of cold which a seed can withstand without losing its vitality, we shall be able to appreciate justly the value of this opinion. The fact is, the vitality possessed by the embryo in the gem, like that it enjoys in the seed, is not susceptible of the stimulus of heat under a certain degree of temperature; and until it re- ceives this, in combination with circumstances otherwise favourable for vegetation, no change of organization is produced in it, and the vital prin- ciple remains unaffected, even in very low tem- peratures. Nor is this wonderful when we con- sider that a caterpillar may be frozen, and yet live after it has been thawed. But if the gems remained uncovered duiing the long period which intervenes between their formation and evolution, they would run great hazards from the effect of moisture, and from the depredations of insects, against which the scales and the varnishes which cover them are excellent safeguards. We, besides, know that light is unfavourable to the evolution of the embryo in the seed, and may we not conclude from analogy, that this is the case also to the young branch in the gem? When the spring returns, and the temperature LECT. IX.] HYBEKNACULA GEMS. 4.77 of the atmosphere has arrived at that point which the excitability of the gem demands for arousing its vital energy into activity, the outer scales being no longer useful, drop off; but the inner ones remain and assume the functions of leaves, until the real leaves are fully expanded. All the gems on a tree, however, do not open at the same time, for the current of fluid in the sap vessels communicating with the terminal gems being more direct than that which supplies the lateral gems, the former always open sooner than the latter. Flower gems almost always open before leaf gems on the same tree; but flower gems which are at a distance from leaf gems, generally fall with- out producing fruit ; and perhaps they possess al- together less individuality than leaf gems, yet flower gems live and flourish when they are de- tached and then budded near a leaf gem on the same or on another stock. We are now treating, however, of the conservative organs only of plants ; I shall, therefore, at present, not enter upon the examination of the flower gem; but pass on to treat the contents of the leaf gem ; or leaves and their appendages. LEAVES are organs of essential importance to the vegetable. They are, also, objects of great de- light and interest, whether we examine them indi- vidually as the clothing of a single plant, or col- lectively as producing the lively freshness of the 478 CONSERVATIVE ORGANS. [t,ECT. IX. verdant vale, and the massive luxuriance of the darkened forest. The most beautiful flower loses half its charms when it is displayed on a naked stem; the miserable hovel becomes picturesque when spread over with the foliage of the Vine ; the ruins of former magnificence acquire more reverence, and command a double share of our respect, when seen through the tracery of the Ivy; and the horrors of the frowning rock are softened into beauty when mantled with pendent creepers or with Alpine shrubs. Leaves are still more important when we regard them as af- fording food to man and the rest of the animal creation; and supplying medicinal agents to re- lieve their sufferings in disease. Notwithstanding, however, the interest which they thus excite ; and our familiarity with leaves, as objects of sight, from our earliest years, it is impossible to form an un- exceptionable definition of the leaf. This difficulty arises from the great diversity of figure, substance, surface, and colour which it assumes in different plants. If we cannot, therefore, define it accu- rately from its external characters, we must have recourse to its functions; and perhaps the following is the least exceptionable definition we can offer : The leaf is, a temporary organ of plants., which performs nearly the same function in the economy of vegetable life as the lungs perform in that of animal life : or, in fewer words, leaves are the LECT. IX.] LEAVES. 479 respiratory organs of plants*. It may be ob- jected to this definition, that some plants, as for example the Dodder, Cuscuta Europea, 'the Sta- pelias, and many of the Cactus tribe, are devoid of leaves ; but in these instances, and in all aphyl- lous plants, the surface of the stem performs the function of the leaves. The diversity of character which leaves dis- play is taken advantage of by systematic Botanists for determining species, and consequently every circumstance connected with that diversity, as form, substance, position, attachment., and direction should be made familiar to the student ; as well as the more intimate or internal structure of the leaf itself. In our examination, therefore, as in the case of stems, I shall first demonstrate the external characters of leaves, and then investigate their anatomy or internal structure. Let us take any leaf from among those now scattered before us ; this for instance of the Lilac. * " Folia transplant et adtrahunt (uti pulmones in anima- " libus) umbramque praebent." Phil. Bot. 81. 480 CONSERVATIVE ORGANS. [liECT. IX. We find that it consists of two parts ; the one, a. (see the figure) thin and expanded, and which in common language is named the leaf (folium); the' other b. long, equally thick as broad, and stalk- . like, which is denominated the footstalk or petiole (petiolus). The footstalk and the expansion, however, constitute but one organ or proper leaf, the footstalk being merely a prolongation of the mid-rib c. c. which in this leaf divides the expansion into two equal portions. This is further proved by the fact, that the expansion cannot i>e separated from the footstalk without tearing or cutting ; and that in autumn, when a leaf withers, both parts fall together, the whole leaf separating, on the slightest touch, at the point where the foot- stalk is attached to the branch*. But many of the leaves before us have no prolongation of the mid-rib, thence we conclude that the petiole is not universal. Continuing our examination, we observe that the two surfaces of this leaf are not alike; that one is of a deep green colour and smooth ; the other is a pale green and marked by a num- ber of elevated ridges (costulce), branching off from the mid-rib : the deeper green and smoother surface is always turned upwards or towards the light, and is named the upper disk (pa- * The error of Linnaeus's definition of the footstalk is very apparent: " Petiolus, trunci species, adnectens folium, nee " fructificationem.'' PhlL Bol. SI. F. LECT. IX.] LEAVES. 481 gina superior), or face of the leaf; the latter, which has of course the opposite direction, is termed the under disk (pagina inferior)) or back of the leaf: but, as I remarked with respect to the footstalk, these distinctions of surface are not universal, for we meet with some leaves which stand vertically on the branches and have both surfaces alike*. That part of the leaf d. d., which is next to the footstalk or to the point of attachment, is always considered as the base ; and the part e., which is directly opposite, the apex ; whatever may be the shape of the leaf. The line e.f. d. d.f. e. forming the contour of the leaf, is named the margin. The angle which the leaf or its footstalk forms at its point of attachment with the stem, or the branch, is termed its axilla. Leaves are either Simple, consisting of one expansion only, with or without a footstalk, as those of the Lilac, the Apple tree, the Nettle, Urtica dioica, and many other plants; or Com- pound, consisting of several distinct expansions, with or without distinct footstalks united to- gether on one common footstalk, as those of * Mirbel considers these as transformed footstalks. He re- marks, speaking of the Acacia of New Holland, " A mesure " que les folioles disparaisserit, les petioles changent visible- " ment de forme et de structure. La plupart s'elargissent vers " les deux bouts, a la maniere d'un fer de lance." Element de Phys.veg. 1. 149. VOL. I. I I 482 CONSERVATIVE ORGANS. [LECT. ix. Buck-bean, Menyanthes trifoliata, of Horse Ches- nut, <#sculus, the Vetch tribe, Vicia, &c. A. SIMPLE LEAVES (Folia simplicia) differ in respect of general Jigure ; form or solid configu- ration ; apex; base; margin; surf ace ; and sub- stance. a. The general Jigure or superficial aspect of a leaf is derived from the line which circumscribes its flat surface, or which is described by its mar- gin. In demonstrating the diversities which it displays, I shall begin with the simplest and pass progressively to the more complex. A leaf is termed Capillary (folium capillare), a. } when it is long, fine, and flexible, resembling a hair. Linear ( linear e), b., when it is long, about a geome- trical line in breadth, and the sides parallel, or is the same breadth near the apex as at the base. Gramineous or riband-like (fasciarium), c. y when it resembles the linear, with which it is sometimes confounded, but from which it differs in being broader and not parallel towards the apex. Needle-shaped (acerosum *), d., when, resembling * " Acerosum est lineare persistens : ut in Pino, Abiete, " Junipero, Taxo.'* Phil. Bot. Notwithstanding this high au- LECT. IX.] GENERAL FIGURE OP LEAVES. 483 the linear, it is rigid and acute. Awl-shaped (subulatum), e. } when it is thick at the base and gradually attenuated to a sharp point. Lanceolate (lanceolaturn) , f., when it is four or five times longer than it is broad, and tapers towards both the base and apex. Sword-shaped (ensi- forme), *., when it is long, tapering to a point; very thin on both edges, and slightly curved. Spatu- late (spathulatum), h., when round at the apex it gradually tapers towards the base. Wedge- shaped (cuneiforme vel cuneatum), i., when broad and abrupt at the apex it tapers towards the base. Fan-shaped (flabelliforme) , A:., when it re- sembles the wedge-shaped leaf in the base, but is more dilated and rounded at the apex. Ob- thority, neither the leaves of the Yew nor of Juniper can be re- garded as needle-shaped. Linnaeus adds another circumstance connected with the needle-shaped leaf, more correct, " ple- " rumque basi articulatione ramo inserta." Phil. Bot. 277. 484 CONSERVATIVE ORGANS. [LECT. IX' long (oblongum), when the length exceeds the breadth at least more than three times, with the ends generally rounded: but the base and apex may be variously defined ; " and this term," as Sir J. E. Smith justly remarks, " is used with great " latitude." Oval or elliptical (ovale vel ellip- ticum), /., when it is twice as long as it is broad, and is nearly equally rounded at both extremities. Ovate (ovatum), m., when the length is greater than the breadth, with both extremities rounded, but the base much broader than the apex. Obovate (obovatum),n. } when it has the ovate shape reversed ; and is consequently attached by the narrower ex- tremity. Roundish (subrotundum) , o., when it ap- proaches to the circular figure. Circular (orbicu- lare),p., when its length and breadth are equal and the circumference is a circular line. Crescent- shaped (lunulatum s. semilunatum), q., when it is curved, as the name implies, like a crescent: whe- ther the footstalk be inserted into the concave or the convex edge of the crescent. Angled (angu- LECT. IX.] GENERAL FIGURE OF LEAVES. 485 mmm fi latum), when the circumference has considerable projections, which are not lobular: and the leaf is termed three-angled (triangulatum), r., four- angled (quadrangulatum) , and five-angled (quin- quangulatum), s., as the angles are either three, or four, or five. If the angles be obscure, the leaf is said to be repand (repandum). It is trowel-shaped (deltoides*), t., when it has three angles, or * " Deltoides, rhombeum est ex quatuor angulis, e quibus " laterales minus a basi distant quam reliqui." Phil. Bot. Sir J. E. Smith states that " a wrong figure is quoted for this in " Philosophia Botanica, which has caused much confusion." Introduc. p. 155. He might have added, that the whole de- scription is erroneous. n3 486 CONSERVATIVE ORGANS. [LECT. IX. resembles the Greek A, one of the angles forming the apex of the leaf: and diamond-shaped (rhom- boideum), u., when the lines describing the edges of the leaf, instead of being curved, form obtuse angles pointing outwards on each side. A leaf is fiddle-shaped (pandurceforme) , v., when it is oblong and has a curvilinear indentation in both its sides : and lyre-shaped (lyratum), *, when there is one large circular or elliptical lobe towards the apex, and several small lateral lobes towards the base. It is termed lobed (lobatum), when it is deeply divided into rounded segments ; and is, therefore, said to be two-lobed (bilobum), w. ; three-lobed (trilobum), x. ; four-lobed (quadrilo- bum), &e. according to the number of the lobes, LECT. IX.] GENERAL FIGURE OF LEAVES. 487 It is arrow-shaped (sagittatum), y., when the disk is triangular, and the sides are produced down- wards into two pointed lobes, like a barbed ar- row: and halberd-shaped (hastatum), z. 1., when the sides are produced into two lateral spread- ing points or lobes near the base. Sometimes the lateral lobes are distinct, as represented at z. 2, A leaf is heart-shaped (cordatum), a., when it is hollowed at the base into two lobes and pointed at the apex, so that the leaf has some- what the appearance of the heart on a card. When the apex 5 instead of being directly oppo- site to the base, is thrown off at one side, the leaf is said to be oblique cordatum, , as beautifully illustrated in Begonia. Kidney-shaped (reni- n4 488 CONSERVATIVE ORGANS. [LECT. IX. forme), i/., when the apex is broad and rounded, and the base deeply hollowed out. A leaf is termed palmated (palmatum), , when it is cleft into oblong or finger-like lobes, not, however, ex- tending to the base; but leaving an entire flat space, which has been likened to the palm of the hand. Laciniated or incised (laciniatum sen sec- turn) , s, when it is cut into numerous irregular divisions, which are termed segments. Parted (partitum), i., when the clefts reach nearly to the base : and according to the number of these, the leaf is said to be bipartitum, tripartitwn, qua- dripartitum, qmnquepartitum, multipartitum. It is said to be cloven (fissum), when the margins of the segments are nearly straight lines : and accord- ing to the number of the clefts the leaf is termed bifidum, trifidum, multifidam^ x. (page 491), &c. Runcinate (runcinatum) , A, signifies that the ex- pansion is deeply cut into many transverse acute- angled segments, the points of which tend to- wards the base of the leaf. When the segments are deeper, and more regular and distant from LECT. IX.] GENERAL FIGURE OF LEAVES. 489 each other, the figure of the leaf is termed pin- natifid (pinnatifidum) , ^; and pectinate (pectina- tum), v, when the segments are very narrow, linear, and parallel like the teeth of a comb. These terms are frequently combined to ex- press modifications of two or more of the forms they imply conjoined in one leaf. Thus you will find in the descriptions of leaves, in systematic works, such terms as ovato-lanceolatum, cordato- lanceolatum, kastato-lanceolatum, cordato-ovatum, lineari-lanceolatum, cordato-sagittatum, sagittate - ovatum, lanceolato-ellipticum, mbrotundo-corda- tum, &c. the meaning of which can be accurately acquired by practice only in the examination of plants. The three last-mentioned terms, I ought to notice, involve a contradiction; or, at least, are too nice for practical purposes ; and con- sequently we find, that several of the best sys- tematic Botanists have confounded and misap- plied them. The terms incisum and dissectum are merely modifications of lacmiatum. I may take this opportunity of remarking that the 490 CONSERVATIVE ORGANS. JJLECT. IX. terms angled and lobed are, also, not unfre- quently misapplied, although the distinction is perfectly obvious: thus, each segment of an angled leaf has the margins, which meet to form the angle, straight or nearly so; whereas in a lobed leaf the margins are always curved, so as to give the lobes a rounded appearance, whether their apexes be obtuse or pointed. The import- ance of accuracy in the application of terms sig- nificant of the forms of leaves, will be fully seen when I come to treat of the classification of plants. b. The solid configuration of a leaf is taken from its real form, including length, breadth, and thickness; determined by transverse and longi- tudinal sections. It is termed Cylindrical (teres, cylindrica), 1, when a transverse section, made any where throughout the greater part of the length of the leaf, is circular. If the diameter be very small, so that the leaf is as fine as a hair, the configuration is termed capillacea, the distinction of which from capillary, consists in the form of the capillaceous leaf being exactly that of a hair, whereas the capillary is only as small as a hair : Semicylindrical (semicy- lindracea), 2, when one side of a leaf is flat and the other convex: Tubular (tubulosa), 3, when the greater portion of the leaf is cylindri- cal, or nearly so, tapering to a point, and hollow within. Sometimes the hollow appears as if it LECT. IX. j SOLID CONFIGURATION OF LEAVES. 491 were formed by the two sides of the leaf being compressed together, but separated near the midrib, so that one part of the leaf is flat and another tubular, as beautifully exemplified in the genus Sarracenea, 4. The configuration is four-edged (tetragona}, when there are four longitudinal sides, and consequently four corners: but if there be three sides only, as in Mesem- bryanthemum aureum, the configuration is termed trigona, 4 (see page 492) ; Linnaeus uses trique- trum to express an awl-shaped leaf, which has three flat sides; but the term is superfluous. Tongue-shaped (UngulataJ, 5, implies that the leaf is thick, oblong, and blunt. This form of leaf is often cartilaginous at the edges, as in some of the Aloe tribe. The configuration is gibbous (gibbaj, 6, when it is thick and swells out, or is humped on one or both sides. Symitar- shaped (acinaciformis) , 7) when one edge is thick, flat, and nearly straight, and the other thin, sharp, and curved like a symitar. Hatchet- shaped (dpfabriformis), 8, when it resembles the 492 CONSERVATIVE ORGANS. [LECT. IX. former, but has the keel or compressed part more abruptly prominent, and the base nearly cylin- drical. Sir J. E. Smith remarks, that " these two " last terms might well be spared, as they seem con- " trived only for two plants," but this is supposing that the whole vegetable world is already known, which is very far from being the case. The same leaves are examples, also, of the two next terms of configuration: the Compressed (compressa), which is used when a thick leaf is flattened laterally, so as to make it thicker than it is broad ; and the flat (plana), when both surfaces of a thick leaf are flat and parallel to each other. A two- edged leaf (folium anceps), JO, displays both the edges, in a transverse section, produced to a very acute angle. The configuration is spherical (sphceroidea) , when it approaches to the globular form : Ovoide (ovoideaj, 1 1 , when it some- what resembles that of an egg: Coccoon-shaped (fusina), 12, when it is cylindrical in the middle and tapers to a point at each end : Club-shaped (clavata), when it is round and stem-like, with a LECT. ix.] APEXES OF LEAVES. 493 thick, blunt apex : Hooked (uncinata), 1 3, when it is curved so as to resemble, in some degree, a hook : Lenticular (lenticularis) , when it is flat, roundish, and convex on both surfaces, and a transverse section of the leaf has the appearance represented at 14, in the cut. c. The apex of a leaf, as has already been de- scribed, is that part which is opposed to the base or the footstalk ; or to the point of attach- ment when it has no footstalk. It differs very considerably in shape in different leaves. The apex is termed acute (acutus), 15, when the conjunction of the two lines of the edges forms an acute angle : acutiusculus, 1 6, when there is a slighter degree of this kind of termination : acu- minate ( acuminatus) , 17, when it is long and very tapering : spine-pointed (cuspidatus), 1 8, when it runs out gradually into a small, awl-shaped, rigid spine; and mucronate (mucronatus) , ]Q, when it is rounded with an herbaceous spine standing on it. The apex is awned (aristatus) ,* , when it is 494 CONSERVATIVE ORGANS. [LECT. IX. terminated by a long rigid spine, which does not appear as a continuation of the leaf: cirrose (cir- rosw, circinatus), 20, when it is produced into a kind of tendril ; and this is in a few instances " fur- " nished with an additional organ for some par- " ticular purpose not essential to a leaf J." Thus, in one species of the genus Nepenthes-}-, 21, the J Smith's Introduction, p. 173. f This singular family of plants was first noticed by Hiero- nymus Benzoni, an Italian, who visited India about the middle of the sixteenth century (1542 1556); and was described by him in a work entitled, " Nova novi Orbis Historia," Genevae, 1578. One species of it, the Phyllamphora (Lin.) 9 was af- terwards fully described and figured in the Herbarium Amboi- nense of Rumphius, who was appointed Governor of Amboyna in 1706. Rumphius regards the fluid found in the pitcher as a secretion of the plant itself, and says it increases during the night ; that it has a sweetish taste and attracts worms and other insects into the pitcher; who, however, all die, except a species of squilla, " squilla gibba/' that seems to prey upon the carcasses of the others. He describes the pitcher itself as being beautifully coloured in the inside with purple streaks and spots ; and the lid opening and shutting. This species is found in LECT. IX.] APEXES OF LEAVES. 495 apex of each leaf terminates in a long rigid thread, a continuation of the midrib, bearing a small covered pitcher, which is generally found nearly full of water : in another species this pitcher is sessile/* 1 ; and, in Dionsea muscipula, the ap- pendage is composed of a pair of toothed lobes, 22, which are irritable; and which close together and imprison insects that alight upon them-f~. The apex is obtuse (obtmus), 23, (p. 493), when it forms the segment of a circle or is rounded. The rounded apex of a solid leaf, when a little thick- Amboyna growing in dry waste places; and also in Ceylon; but I suspect that the Ceylon plant is a variety, if not a distinct species, for the pitcher is not contracted at the neck, as in Rum- phius's figure, and it is found growing only in moist valleys and on the banks of rivers. At this time (August 1821), there are several plants of Nepenthes distttlatoria vegetating in pots, in the magnificent hothouse of Messrs. Loddiges, at Hackney. The pitcher in this species is attached to the apex of the leaf, without the medium of the twisted wire, which is found in Phyllam- phora ; and there are, also, two leafy appendages running the whole length of the pitcher, on that side of it which is next to the plant. The lid exactly resembles that of Phyllamphora. A more beautiful vegetable pitcher is found in the Cephalotus^/^ttfo- ris, a New Holland marsh plant, which was discovered and de- scribed by Mr. Robert Brown, to whose exertions and talents Botanical science is most extensively indebted ; and is figured in the atlas of Capt. Flinders's voyage ; but, as it is not ap- pended to the leaf, I shall describe it among the general ap- f The cause of this vegetable phenomenon shall be after- wards investigated. 496 CONSERVATIVE ORGANS. [LECT. IX. ened, but not sufficiently so as to render it clubbed, is termed incrassatus. If a small point project from the middle of the obtuse apex, the leaf is said to be obtusum cum acumine, 24 (p. 493). The apex is termed retuse (retusus), 25, when it is obtuse, with a broad shallow notch in the middle: emarginate (emarginatus), 26, when the notch is sharper, or nearly triangular : truncated (truncatus), 27, when it appears as if cut across in a straight line; as beautifully exem- plified in the leaf of Liriodendrum tulipifera. It is jagged (prcemorsus) , 28, when it appears as if gnawed off, and the cross lines describe several ir- regular points. Ex. Caryota urens (28) ; and tri- dentate (trident atus) , 29, when it forms three teeth. 25" d. The differences in the bases of leaves depend on the general superficial configuration ; I must, therefore, refer you to what has been said on that subject, and notice here one circumstance only, LECT. IX.] LEAVES. 497 connected with the base, which has not already been described. When the two halves of the ex- pansion are of different lengths, this is observable chiefly at the base, and the leaf is said to be un- equal or oblique (basidncequale), 3O, (page 496). e. The Margin of a leaf may be either entire, indented, bordered, or rolled. * Entire. An entire leaf (folium integerrimum) has the line of the margin uninterrupted, or free from every kind of incision or indentation. Sir J. E. Smith properly remarks that this term refers solely to "the margin of a leaf; whereas in- " tegrum respects its whole shape, and has no- " thing to do with the margin-^-." I may add, it is used in contradistinction to compositum. ** Indented. A leaf is termed sinuated (smuatum), 31, (page 498), when the margin is cut into roundish scollops, as in the Oak, Quercus robur; but when the notches or scollops are very irregular, as if formed by the gnawing of some insects, the margin is then said to be gnawed (erosus), 32. It is termed toothed (den- tatus), 33, when it displays pointed marginal pro- jections of the expansion, with interstices between them ; and the following terms are employed to express the character of the margin as far as re- f Introd. to Phys. and Syst. Bot. p. 161. VOL. I. K K 498 CONSERVATIVE ORGANS. [LECT. IX. gards its denticulations: equally toothed (cequaliter dentatus), 34 ; unequally toothed (incequaliter den- tatus), 35 ; deeply toothed (profunde dentatus) ; obscurely toothed (obsolete dentatus), 36. If the denticulations themselves be again dentated, then the margin is termed doubly dentate (duplicato- dentatusj, 37. The margin is denominated serrated (serratus), 38, when the teeth are sharp, and lie as it were upon each other, as in a saw, all pointing towards the apex of the leaf. The nature of the serratures is distinguished by the following terms applied to the margin : equally serrated (cequaliter serratus); unequally serrated (incequaliter serratus); sharply serrated (argute serratus) ; deeply serrated (pro- funde serratus), 3Q; and doubly serrated (du- pUcato-serratus), 40. When the serratures are minute, or not distinct, the margin is termed serrulatus, 41. LECT. IX.] LEAVES. 499 Crenated (crenatus), 42, implies that the in- dentations of the margin are blunt and rounded, and do not incline to either extremity of the leaf. The crenatures themselves may be crenated, in which case the margin is termed doubly crenated (duplicato-crenatus); or they may be of a doubt- ful form, being neither completely rounded nor yet pointed, in which case it is termed dent at o- crenatus, 43 ; and crenulated (crenulatus), 44, if they are very shallow and at the same time perfect. When the marginal denticulations, whatever form they assume, are terminated with sharp, rigid spines, the margin is termed spinous (spinosus), 45; and as a comparative term, a leaf is said to be unarmed (inerme), when it occurs in a species or a variety belonging to a tribe of plants, which has for the most part spinous-leaved species. Bordered. When the substance of the margin of a leaf differs from that of the expansion, the leaf is con- sidered as bordered ; and according to the charac- KK2 500 CONSERVATIVE ORGANS. [LECT. IX. ter of the border, the margin receives different ap- pellations. Thus it is termed cartilaginous (carti- lagineus), when it is firmer than the disk and somewhat elastic. This cartilage is generally whitish, yellowish, pinkish, or some other colour, but is seldom green. It is termed horny (cor- neusj, when it resembles the cartilaginous, but is harder and less elastic: ciliated (ciliatm), 46, when beset with soft parallel hairs, not closely set together ; but if the hairs be stiff and like bristles, it is then said to be aculeato-ciliatus. (See fig. 1 . Plate 4). If the margin be studded with small granular, either opaque or semitransparent bodies, exuding some kind of fluid, it is termed glandular (glan- dulosus) ; but, if these glands be supported on hairs, glanduloso-ciliatus, 48. **** Rolled. When the margin is rolled backwards, or upon the under surface of the leaf, it is said to be re- volute (revolutus), 49; when forwards, involute (in- volutus). In some instances the margin, compre- hending a portion of the disk of the leaf, is so much more expanded than the rest of the disk, that it assumes a waved character, or is undulated (un- dulatus), 5O ; and when it is still more expanded, so that the margin is variously curled and twisted, it is termed curled (crispus). These appearances occur in leaves, which have very different kinds of margins in other respects. LECT. IX.] LEAVES. 501 f. The surface of a leaf comprehends both the upper and the under disks. In general, the upper disk is smoother than the under disk ; for, although the vascular fasciculi can be traced in the form of white or coloured lines on the upper, yet, they very rarely produce those elevated ridges which mark their course on the under disk. In treating of the different characters that distinguish the surfaces of leaves, the upper disk only is alluded to. It is necessary to state that the greater number of the following terms are equally applicable to the surface of the stem as to that of the leaf, when the contrary is not expressed. The surface of a leaf which exhibits no in- equalities is said to be flat (plana vel lasvis) ; if there be no hairs nor spines, it is smooth (glabra, nuda); and if no fasciculi of vessels be apparent on it, veinless (aveniaj. If it be smooth and shined, it is termed polished (nitida); and lucid (lucidaj, if it be so considerable as to present the appearance of being varnished. If the upper surface be con- vex and the under concave, the leaf is termed convex (convexum); and the opposite of this state, concave (concavum) . Both these states are the consequence of a tightness of the margin. When an oblong or a linear leaf is longitudinally hollowed, and a transverse section of it is a se- micircle, the surface is said to be channelled (ca- naliculata) , 52, (p. 503); but when the transverse KK3 502 CONSERVATIVE ORGANS. [LECT. IX. section is angular, and the midrib on the under surface resembles the keel of a boat, it is keeled (carinata), 53. When, instead of one longitudinal hollow, there are several linear depressions, the surface is said to be furrowed (sulcata), 54 ; and streaked (striata), if the depressions be super- ficial, very narrow, and in parallel lines. If the surface be depressed in the centre, and the leaf is peltate, it is said to be navel-like (umbillcata) , 55: but if it rises and sinks alternately, in straight angular furrows, like the folds of a fan, it is termed folded (plicata) ; and waved (undulata) if in un- dulations commencing from the midrib. When the inequalities proceed from the portions of the expansion between the network of the vascular fasciculi being fuller than is requisite to fill the vacant spaces of the meshes, and rising upwards, the surface is wrinkled (rugosa), 56; and blis- tered (bullata), when these elevations are still more considerable; but when, on the contrary, the fulness between the vascular fasciculi produces der pressions, the surface is said to be pitted (lacu- nosa velfavosa). When a leaf is covered with small hard tuber- cles, which are more easily distinguished by the finger than the eye, the surface is said to be sca- brous (s cobra) : it is rough (asp era) when these are more visible ; warty (verrucosa), 57, when they are still larger and more solid ; and pustular, LECT. IX.] LEAVES. 503 or vesicular (papillosa), when they are evidently elevations of the cuticle filled with aqueous fluid, as in the Ice plant, Mesembryanthemumtr^frz/Kmtm. If the surface of a leaf be studded with short her- baceous spines, it is termed muricated (muri- cata); when these have stiff points, it is prickly (echinata) ; and aculeated (aculeate), when., in- stead of being herbaceous, the spines are hard and pungent. The surface is termed hispid (his- pidajy when it is beset with short stiff hairs ; when these are longer, and consequently less rigid, it is hirsute (hlrsuta) ; bristly (setosa), 59, if they stand 2. singly and resemble bristles ; strigose (strigosa), if they are firm, and stand upon small prominences, or papillae* : and bearded (barbata), when they are rather long and crowded together. Soft hairs are generally termed pubescence, and the surface of the leaf receives various appellations from the * " Strigae arcent setis rigidis animalcula et linguas. Cae- " tus, Malpighia, Hibiscus, Rubus." Phil. Bot. 163. K K 4 504 CONSERVATIVE ORGANS. JJLECT. IX. character of this description of covering. Thus it is hairy (pilosa), when the hairs are soft, distinct, somewhat long, and bent ; shaggy or velvety (vil- losa), when they are soft, nearly erect, and paral- lel; silky (sericea), when they are soft, and 'lie thick and flat on the surface, giving it a satin-like lustre; downy (tomentosa*), when they are very soft and matted together, so that the individual hairs are not distinguishable; woolly (lanata^-), 60, (page 503), when they are also matted together, but yet individually distinguishable ; tufted (floccosa), when they are soft and matted, and can be easily detached in small tufts; and starred (stellata), when the hairs or the spines are radiated like stars. In some instances the stellated appearance proceeds from pedicillated tufts ; which under the microscope appear like branched shrubs de- void of foliage. It is necessary to remark that differences of age, climate, soil, dryness and hu- midity, alter very considerably the pubescence of plants ; some which are tomentose when young, become hispid, or smooth, when old; others which are naturally hairy, become quite smooth when cul- tivated ; and others, which in their native climate are smooth, become hairy when removed. Sir J. E. Smith, nevertheless, in opposition to the * " Tomentum servat plantas aventis; gaudet saepius co- " lore incano." Phil. Bot. 163. f " Lana servat plantas ab sestu nimio." PhiL Bot. 163. LECT. IX.] LEAVES. 505 maxim of Linnaeus, " pubescentia ludicra est dif- " ferentia, cum cultura ssepius deponantur *," remarks that " the direction of the hairs or " bristles proves a very sure means of distinguish- " ing species, especially in the genus Mentha, the " hairs about whose calyx and flower-stalk point " differently in different species; and/' he adds, " I have found it the only infallible distinction " between one Mint and another ^." The surface of a leaf may be furnished with visible glands. When these are elevated, or on pedicils, the surface is said to be glandular (glan- dulosa) ; but, when they are not raised, they ap- pear like punctures, which either penetrate the substance of the leaf, or are merely superficial and visible on one disk only. In either case the surface is said to be dotted (punctata), 58, (page 503). Any glandular exudation, if considerable on a surface, gives a character and denomination to it: thus, if it be moist and tenacious, the surface is termed viscid (vlscida vel glutinosa) ; it is hoary (pruinata vel incana), if the secretion be a dry, very fine, waxy powder of a bluish colour, devoid of gloss, and easily wiped off; and mealy (farinosa), i it resemble a mealy powder. The majority of leaves is green ; and, as has bqen already mentioned, this colour is of a deeper * Phil. Bot. 272. f Introd. p. 228. 506 CONSERVATIVE ORGANS. [LECT. IX. tone on the upper than on the under disk: but the green colour, from its being nearly uni- versal, is not noticed in the description of a leaf, except as regards its depth or tone ; thence, the comparative terms dirty (sordidus) ; intense ( tn- tensus) ; full (saturatus) ; pale (pallldus) ; and diluted (dilutus). The colour is denominated olive (olivaceus) when it is green with a shade of brown ; and sea-green (glaucus), when it appears bluish, or as if formed from a mixture of blue and green. When other colours mingle with the green, the leaf is termed coloured (coloratum) ; and the differences of the colouring receive a determinate designation : thus, when the coloured portions afford the semblance of decay or disease, the leaf is termed variegated (variegatum) ; spotted or maculated (maculatum), when the colour is dark and in spots or blotches; and zoned (zonatum), when the colours are displayed in one or more curved lines. The term discolor is employed to denote that a leaf displays one colour on one sur- face, and another on the opposite surface, as in purple-leaved Spiderwort, Tradescantia discolor,. two-coloured Begonia, Begonia Evansiana, &c. The vessels of every leaf enter it by the petiole, or by the point of attachment, in fasciculi ; which afterwards separate, and are spread in various ways through the substance of the leaf. These, when visible, form lines, which are very LECT. IX.] LEAVES. 507 prominent on the back or under surface of the leaf, and have been very improperly denominated nerves and veins-, for nothing has yet been dis- covered in vegetables bearing any analogy to nerves ; and as the fasciculi ' contain both con- ducting and returning vessels, they may be termed arteries with as much propriety as veins. But the terms, notwithstanding their absurdity, being in general use, require to be explained. The larger fasciculi are denominated nervi and costas, and the smaller vence, in whatever man- ner they are disposed throughout the leaf; but the terms nervosum, costatum, and venomm, imply distinct dispositions of the fasciculi. A leaf is said to be nerved (nervosum) when the larger fasciculi run, in simple, slightly curved lines, from the base to the apex ; and the leaf is named according to their number, among which the midrib is reckoned as one. Thus three-nerved (trinervis), 6i, (page 509), means that the leaf has one longitudinal fasciculus of vessels on each side of the midrib, taking its origin from the base ; and five-nerved (quinquenervu), that it has two lon- gitudinal fasciculi on each side of the midrib, un- der the same circumstances. It is seven-nerved (septemnervis), 62, when there are three fasciculi on each side; nine-nerved (novemnervis), when there are four ; and many-nerved (multinervis}, when the number on each side exceeds four. The 508 CONSERVATIVE ORGANS. [JLECT. IX. nervation is in some instances not regular ; but one large fasciculus is given cff on each side of the midrib, at the very base of the expansion, dividing and subdividing as it proceeds, as in the annual Sunflower, Helianthus annuus; in which case the leaf is said to be basi-trinerve, 63. The same leaf is also an example of the term venoso-nervosum, which is used by some authors to imply that the nerves pass into veins. If the lateral longitudinal fasciculi do not spring directly from the base, but so far above it, that part of the expansion of the leaf is below them, as in the leaf of the Cinnamon tree, Laurus Cinnamomum, 64, the terms em- ployed are triplinerve, quintuplinerve, multipli- nerve. In one example of the multiplinerved leaf, Hydrogetoufenestralis*, 65, the longitudinal fas- ciculi are united by transverse bands, forming square meshes, which are perfectly void, like wire-work, giving that character which Botanists have termed fenestrated, or open ( fenestratum) . But when the longitudinal fasciculi, forming ex- current nerves, are united by transverse fasciculi, however elevated or strongly marked these may be, if the intermediate spaces be filled with the expansion, as in 66, the leaf receives no particular appellation. Ribbed (costatum), is sometimes employed as * This singular plant is an aquatic, a native of Madagascar. LECT. IX.] LEAVES. 509 the synonime of nerved, and is certainly the more proper term of the two ; but it is, also, specifically used to imply that the veins proceed from the mid- rib in lines nearly straight towards the margin, and parallel t6 each other. In some instances the costae are close together and scarcely elevated, as in the Banana, Musa sapientium; but in others they approach to the character of nerves, except that the angle they form with the midrib is less acute, 67, and they do not appear to be split off from the midrib as in the multiplinerved leaf. 61 510 CONSERVATIVE ORGANS. [LECT. IX. Veined (venosum), expresses merely that 'the fasciculi of vessels are so elevated or marked in their branching and sub-branching, as to form a kind of network over one or both surfaces of the leaf. When the 'surface is altogether free from any appearances of the vascular fasciculi, the leaves are said to be nerveless, or veinless (enervia, aveniaj, 70. In speaking of the substance of a leaf, as far as that can be determined from its external cha- racters, we refer to the expansion only ; for, al- though the substance of the petiole is often the same as the expansion, yet this state is neither con- stant nor essential. All leaves may be regarded as herbaceous; but the term is occasionally employed in a determinate sense to denote a soft, green, pliable leaf, the vascular fasciculi of which are as succulent, and scarcely firmer than the rest of the leaf; as exemplified in Spinage, Spi- nacia dleracea, the different species of Goose-foot, Chenopodeum, &c. A leaf is termed membra- nous (membranaceum) , when it is thin and pliable, the quantity of parenchymatous matter being so small that the cuticle of the one surface appears almost closely applied to the other; as in Broad- leaved Birthwort, Aristolochia sipho; Scented Bramble, Rubus odoratus ; and most trees and shrubs : but this term does not imply that uni- formity of substance which the ^ appellation herba- LECT. IX.] LEAVES. 511 ceous denotes. The terms scariose (scariosum), and chartaceous (chartaceum, papyraceum) r are given to varieties of the membranous leaf; the first im- plying that it is dry, or apparently sapless, and somewhat translucent ; the second, that it re- sembles paper, as in Draco terminalu. Leathery (coriaceum) implies that the leaf is thick, tough, and elastic; as in the Mistletoe, Viscum album, Changeable Hydrangea, Hydrangea hortensis, &c. : and rigid (rigidum), that it is hard, with little elas- ticity, as in Butcher's broom, Ruscus aculeatus, Scotch fir, Pinus sylvestris, &c. Fleshy (carnosum) denotes that the leaf is thick, and consists chiefly of a juicy but firm cellular parenchyma, as in Houseleek, Sempervivum tectorum ; and succulent (succulentum), that the pulp is laxer and more juicy, as in Mesembryanthemum echinatum, &c. Mirbel says the consistence of the fleshy leaf is that of the Apple ; of the succulent that of the Plum *. As all the characteristics of leaves, which I have yet described, relate to the expansion, they are consequently peculiar to simple leaves only, or to the individual leaflets of compound leaves. Be- fore detailing those common to both descriptions of leaves, it is requisite to understand the pecu- * Elemens de Phys. veg. iii. p. 642. 512 CONSERVATIVE ORGANS. [LECT. IX. liarities connected with the petiole or footstalk ; and, as naturally arising out of these, the circum- stances which constitute the compound leaf, with its species and varieties. The description of these, therefore, shall form part of the subject of our next Lecture. Before closing this Lecture, however, I may remark that, although the study of the terms which denote the peculiarities of formation in the vegetable body, be apparently the least amus- ing part of our subject, yet, it is not altogether devoid of interest, from the great variety which Nature displays in this part of the creation ; and I can assure those who may think it tedi- ous, tKat the readiness with which it enables us to describe plants in proper Botanical language, and to understand the descriptions of others, fully repays the labour of the acquirement. Like learn- ing a language of any kind, its utility becomes evident only after its acquisition; but then the stores of knowledge to which it serves as the key, are opened with a facility which is not less gratify- ing, than their magnitude and richness are asto- nishing and their possession delightful. 3 513 LECTURE x. i OP THE PETIOLE * COMPOUND LEAVES. THE SITUA- TION, POSITION, INSERTION, DIRECTION, MAGNI- TUDE, AND EXTENSION OF LEAVES. ANATOMY OP THE LEAP. THE FOOTSTALK (petiolus) in every instance in which it is present, constitutes a part of the leaf. It is simple, in some instances, consisting of one piece only, as in all simple leaves; or it is com- pound (composites), consisting of one common stalk divided into several distinct parts. The com- mon petiole is generally termed primary (pri- marius) ; the immediate divisions of this se- condary ( secundarii) ; the divisions of these ter- nary (ternarii), and those which support the leaf- lets partial (partiales seu proprii). The petiole further differs in form, in the nature of its ap- pendages, its mode of insertion into the leaf, and its articulations. In respect of form, some of the terms em- ployed in the description of the stem are applicable to the petiole. Thus it is said to be round (teres) ; half-round (semiteres); compressed (compressus), &c. according to the figure of its trans verse section. The compressed petiole occasionally assumes the VOL. I. L L 514 CONSERVATIVE ORGANS. [LECT. X. aspect and functions of a leaf, in leaves which suffer that change which is termed alienated (alienatum) ; that is, when the compound leaves which are the first leaves of the plant, and na- tural to the tribe, afterwards give place to simple leaves, which are merely dilated petioles ; as exemplified in several of the Mimosas from Botany Bay. De Candolle has named these in- termediate forms phyllodia. The transition might be supposed to depend on the energy of the plant diminishing with its growth; for, when a plant of this kind is topped, the new leaves which are put forth display the same characters as the original foliage * ; but, that this is not the case is evident, from the transition occurring constantly in New Holland, where the genus has every op- portunity of following its natural habits. The ma- jority of petioles are slightly furrowed on the up- per side ; but, when this is deep, it is particularly denoted by the term channelled (canaliculatus), and when much dilated the petiole is said to be concave (concavus). In the last-mentioned in- stance it sometimes partially embraces the stem * I saw an illustration of this fact yesterday (25th September 1821), in the garden of Comtesse de Vandes, near Bayswater. An old plant of Acacia falcata, on which no pinnated leaves had been seen for several years, had been cut down and had put forth pinnated leaves, resembling in every respect, except in size, the first leaves of the plant. The second series of shoots, however, displayed falcated leaves only without any pinnae. LECT. X.] LEAVES THE PETIOLE. 515 (amplexans), 7^; or wraps completely around it, and then is termed sheathing (vaginans), as in the Grasses. In some cases the petiole is inflated (in- flatus), and the plants in which this occurs being aquatic, it serves to float the leaf, as in Trapa natans, 73, &c. The appendages of the petiole are not nume- rous. A footstalk is said to be winged (alatus), when it has on each side of it a portion of the expansion, isolated from the rest, as in the Orange tribe, Citrus, 74. But there are leafy appen- dages of a distinct character, denominated sti- pules, which are yet to be described, in many in- stances attached to the petiole; as in the Rose tribe, Rosa, 75, in which case the petiole is termed stipuliferous (stipuliferus). It is termed glanduliferus, 76, when one or more glands are seated on it, as in the genus Passiflora ; in the Castor-oil plant, Ricinus communis, &c.: and floriferus when it bears the flower, as in Elm- leaved Turnera, Turnera ulmifolia. In respect to insertion, in the majority of in- LL*2 516 CONSERVATIVE ORGANS. [LECT. X. stances the petiole and the expansion are in the same plane; the insertion of the footstalk being at that portion of the margin of the expansion, which is regarded as the base of the leaf; but in some instances it is in the centre of the disk, and then the leaf is termed shieldlike (peltatum), 77. The consideration of terms denoting the other inser- tions of the petiole is deferred until we examine the various insertions of the leaf, of which the footstalk is merely a part. The term articulation, as applied to the petiole, is intended to denote that it consists of more than one piece, the pieces being generally united by a small intermediate portion, thicker and more spongy than the parts it unites, commonly of a different colour and capable of motion. The pe- tiole of the simple leaf is rarely, although some- times, articulated, as in Citrus aurantium, 74, a. ; and, also, in the majority of Grasses, at that point which separates the real leaf from its sheathing petiole, and which is generally marked on the inner side by a small membranous appendage, which the elder Botanists termed ligula and mem- brana foliorum, but which is now properly con- sidered a stipule. Articulation is very common in compound leaves. It occurs generally at the attachment of the partial to the common pe- tiole, in such leaves as fold together their leaflets during the night; but, in some instances, there is an articulation, also, near the point where the LECT. X.] LEAVES THE PETIOLE. 517 primary petiole springs from the branch, particu- larly in those plants which have irritable leaves, as Mimosa sensitiva, &c. At these secondary arti- culations the leaflets separate, and fall in autumn; sometimes before the common petiole falls, par- ticularly when it is not articulated near its in- sertion. The term inarticulatus is employed com- paratively, when leaves belonging to a tribe of plants, in which articulation is common, are de- ficient in this peculiarity. The composition of the petiole necessarily im- plies that of the leaf; but a compound leaf does not necessarily imply the existence of either se- condary, ternary, or partial petioles. The COM- POUND leaf may be thus defined: a leaf which consists of seveal distinct expansions or leaflets, connected by articulations either directly or in- directly, and a common footstalk. This defini- tion differs from that of the majority of Botanical writers, in regarding the connexion of the leaflets by articulations as the chief charasteristic of the compound leaf * ; but I cannot consider any leaf * Mr. Keith seems to be fully aware of this characteristic of compound leaves (see Syst. of Phys. Bot. vol. i. p. 54) ; and Bulliard thus defines the compound leaf: " La feuille composee " a une ou plusieurs folioles attachees au petiole commun par " une articulation, au moyen delaquelle chacune d'elles peut se " mouvoir dans certaines circonstances, et 6tre detachee sans " lesion, soit spontanement, soit artificiellement." Diet, ele- ment, de Bolanique, p. 59. L L 3 518 CONSERVATIVE ORGANS. [LECT. X. as compounded, or consisting of distinct parts, in which the divisions of the expansion, however multitudinous they may be, are still parts of the same expansion, inasmuch as they are con- nected by the direct and immediate prolongations of the same petiolei That the leaflets in a true compound leaf are distinct organs, is confirmed by the articulations being conspicuous by anato- my, and by the natural habits of the leaf. The articulations in many leaves are rendered conspi- cuous by a cartilaginous structure ; and in others by a swelling of the partial petiole, at its point of union with the common footstalk, which almost impresses the idea of an artificial attachment: in some instances, however, this is so obscure, that it is by dissection only that the fact can be deter- mined. In a longitudinal section of the petiole of a simple leaf and its ramifications, the vascular bundles distributed to the latter appear merely slips from those of the common petiole; but in a compound leaf we find a peculiar vascular arrange- ment, closely resembling that which occurs in the knotted culm and other articulated stems (see p. 312), marking the place where the vessels of the common petiole apparently terminate, and those of the leaflets commence ; thus demonstrat- ing the individuality of the leaflet. At this point, also, as has been already stated, the leaflets spon- taneously detach themselves from the common LECT. X.] COMPOUND LEAVES. 519 footstalk in autumn, in the same manner as the leaf detaches itself from the tree. Every ex- pansion, therefore, in a compound leaf, is an in- dividual leaf; and it assumes different forms on different plants, in the same manner as the simple leaf. All compound leaves may be arranged under the three following divisions : simply compound (composita) ; doubly compound (decomposita) ; more than doubly compound (supradecomposita). 1. The simply compound leaf consists of a common petiole only, supporting two or more leaflets. When the leaflets are sessile, attached to the apex of the petiole, and three in number, the leaf is termed trifoliate or ternate (trifoliolatum, ternatum), 82 ; as in the genus Clover, Trifolium, 520 CONSERVATIVE ORGANS. [LECT. X &c. It is quadrinate (quadrinatum), when there are four leaflets, as in Marsilea quadrifolia, 83, and some of the Hedysarums: quinate (quinatum), when there are five, as in ^Esculus Pavia, 84; digitate or fingered (digitatum), 85, when there are seven, as in several of the Potentillas ; and multifoliolatum when the number exceeds seven; or umbellatum, 86, when being numerous they are so arranged as to form the figure of a parasol, as exemplified in many of the Lupine tribe. When the leaflets, instead of being supported on the very apex of the petiole, are attached on its sides, the leaf is termed yoked and pinnate. It is termed simply yoked (conjugatum}, when one pair only of opposite leaflets is supported on a common footstalk, as in the genus Zygophyllum, 87, (p. 521), which is named from this character of the leaves : and bijugum, when there are two pairs. But the terms bijugum, trijugum, quadrijugum, quinquejugum, and multijugum, are not employed, except to denote the particular number of opposite leaflets on a common footstalk, when such an enumeration is requisite for establishing specific characters; otherwise all compound leaves having more than one pair of opposite leaflets attached along the side of a common petiole, are, usually, regarded as pinnate (plnnata). The individual leaflets are termed pinncc; and according to the COMPOUND LEAVES. 521 LECT. X.] arrangement of these on the petiole, pinnate leaves receive different appellations. When all the leaflets are in pairs, and the com- mon petiole, or rachis, is not terminated either by a leaflet or a tendril, the leaf is termed abruptly pinnate (pari-pinnatum, sen abrupti-pinnatum) , 88 ; but it is pinnate with a terminal or solitary leaflet (impari-pinnatum, pmnatum cum impari), when, there is a single leaflet at the apex of the petiole, 8p, .; and the expression, with a tendril (cirroswn), is added, if, instead of a leaflet, the termination be a tendril, as in the Vetch tribe, Vida *, 90, a. If the terminal leaflet be much * There is no specific term to designate a leaf, which has one pair of leaflets only, on a common petiole which, extending beyond them, is terminated by a solitary leaflet, as in the Kidney Bean, Phaseolus vulgaris; for, although the term ternate is usually applied to it, yet it cannot be regarded as ternate on ac- 522 CONSERVATIVE ORGANS. [LECT. X. larger than the other leaflets, the leaf is said to be lyrato-pinnatum ; but this is a term of very rare application in compound leaves ; those so deno- minated being, for the most part., truly simple lyrate leaves, altogether devoid of articulations, or that modification of vessels at the union of the lateral divisions, which is the requisite charac- teristic of the leaflet, and consequently of the compound leaf. If the leaflets be in opposite pairs, and it be not essential to enumerate them, the leaf is said to be oppositely pinnate (opposite pinnatum) y 88 ; b.b. b. b. (page 521 ), and alternately pinnate (alternatim pinnatum), when they are al- ternate, QO, b. c. It is interruptedly pinnate (inter- rupte pinnatum), when the leaflets are alternately large and small on both sides of the petiole, 89, b. c.b.c. ; jointedly pinnate (articulate pinnatum) , when the common petiole is jointed betwixt each pair of leaflets, .91 ; and decreasingly pinnate (pinnatum foliolis decrescentibus), when the leaflets gradually diminish in size from the base to the apex of the leaf, as in Vicia sepium. In al- most every writer on elementary Botany, the term decurrently pinnate (decursive pinnatum) is de- scribed as denoting that the leaflets are decurrent, or united by a foliaceous membrane, extending count of the lateral leaflets, this term implying that all the leaflets are on the apex of the petiole; I would, therefore, propose the term impart '-conjugatum, for the above-mentioned form of leaf. See cut, fig. 97, page 523, which is the leaf of Hedysarum gyrans. LECT. X.] COMPOUND LEAVES. 523 from one leaflet to another, down the common petiole; but this is the description of the pin- natifid leaf; for the real decurrently pinnate is a modification of the jointedly pinnate, each joint consisting of one pair of pinnae supported on a winged petiole (see fig. 91, page 521, a. a. a.). In some instances the leaflets, instead of being ar- ranged in the same plane on each side of the com- mon petiole, are placed around it; and the leaf is consequently termed verticillato-pinnatum. 2. The second division of compounded leaves, the doubly compound, comprehends those leaves in which the common petiole is divided into, or sup- ports secondary petioles. When near the apex of the common petiole, there is a single pair of secondary petioles, each of which supports a pair of opposite leaflets, as in Mimosa unguis cati, the leaf is termed twice paired (bigeminatum, seu biconjugatumj , Q2: it is 92 524 CONSERVATIVE ORGANS. [LECT. X. thrice paired (tergeminatum), when the leaf re- sembles the twice paired in its petiolar divisions, and has besides a third pair of leaflets at the point where the secondary petioles originate, as in Mi- mosa tergemina, Q3; but if the common foot- stalk supports three secondary petioles on its apex, and each of these support three leaflets, then the leaf is termed doubly tern ate (biternatum, dupli- cato-ternatum) , Q4. A leaf is termed doubly pinnate (bipinnatum), when the secondary petioles are arranged in pairs on the common petiole, and each secondary petiole is pinnate, or displays the characters of the simply pinnate leaf, Q5 ; it is conjugated and pinnate (conjugatum-pinnatum), when a common petiole supports a single pair of secondary petioles, each of which is pinnate, as in Mimosa purpurea, 96; ternated and pinnate (ternatum-pinnatum) , when the common petiole supports on its apex three pin- nate leaflets, as in HofFmannsegia: and digitated and pinnate (digitatum-pinnatum) , when the num- ber of these exceed three, as in Mimosa pudica. The pedate leaf (folium pedatum) is general- ly described as a decompound leaf, composed of a common petiole divided at its summit into two diverging branches, with an intermediate leaflet, and each branch supporting two or more lateral leaflets on their anterior edge; as LECT. X.] COMPOUND LEAVES. 525 in Helleborus niger, Q8; but according to the definition of the compound leaf which we have adopted, the pedate is merely a deeply divided simple leaf, having a close affinity to the palmate, or rather, as Sir J. E. Smith observes, who never- theless regards it as a compound leaf, to " those " simple leaves which are three-ribbed at the base." 3. The third division of compounded leaves, the more than doubly compound, comprehends those in which the common petiole supports se- condary petioles, which in their turn support ter- nary petioles. When the common petiole supports, on its apex, three secondary petioles, which each sup- ports three ternary petioles, and every one of these three leaflets, the leaf is termed thrice ternate (tritematum s. triplicato-ternatum), 99, (p. 526) ; and when along the sides of the common petiole, 526 CONSERVATIVE ORGANS. [LECT. X. there are secondary petioles, supporting ternary, which are pinnate, it is triply pinnate (tripinna- tum s. triplicato-pinnatum) , as in the Carrot genus, Daucus, &c. 100, (p. 525). These are the only species of the decompound leaf; but the term is occasionally applied to leaves which are very much and irregularly divided. There is, however, one species of the compound leaf, which we have not yet examined, because it cannot properly be classed in any of the foregoing divisions. It is sometimes termed vertebrated (ver- tebratum), and sometimes jointed ( articulatum) . It consists of several leaflets, which appear to grow out of each other, or are attached one upon the summit of another, with an evident LECT. X.] COMPOUND LEAVES. 527 joint at the point of attachment, as in prickly- leaved Fagara, Fagara tragodes *, 1 00. f02 In the majority of plants the form of the leaf, whatever that may be, is the same over the whole of the plant; but there are exceptions to this rule ; and when the diversity in the form of the leaves is constant, the term heterophyl- lus is occasionally adopted as the specific naine of the plant. Thus various-leaved Spurge, Eu- phorbia heterophylla, is named from having the lower leaves short, wedge-shaped, emarginate and mucronate, while the upper are long linear- lanceolate, acute, and entire, 101. Heterophyllous plants, however, must not be confounded with * Mirbel (Elem. de Phys. veg. Zde Partie, p. 655) quotes the leaf of spike-flowered Cussonia, a curious Cape plant, as an example of the vertebrated leaf; but this Cussonia presents an instance only of a digitated leaf, with articulated, winged, se- condary petioles. 528 CONSERVATIVE ORGANS. [LECT. X. those which have alienated leaves, as exempli- fied in Mimosa verticillata (see 102, p. 527 ; a. the forn^of the earlier leaves, b. those which succeed them) ; for, in these, the leaves, which are alien- ated or different from those which first appeared on the plant, and from those peculiar to the genus, are ultimately the only leaves of the plant. Besides those characteristics of leaves which have been described, there are others common both to simple and to compound leaves, whatever may be their form and structure. These refer to the situation of the leaves ; to their disposition on the stem and branches ; to the mode of their attach- ment or insertion ; and to the direction of their surfaces with respect to the stem and the branches as well as to the plane of the horizon. In point of SITUATION the majority of leaves may be regarded as aerial; being suspended on the stem or the branches, in such a manner that the air is applied to both surfaces of every leaf. The position of such leaves in relation to the hori- zon, is determinate, or always the same in the same description of plants, but varies in different kinds of plants ; and this constitutes, in Botanical language, their direction: in ascertaining which, the plant must be in a healthy state, for an un- healthy state of the plant and the natural decay of the leaf in autumn, alter the ordinary direction of this organ. The following are the determinate LECT. X.) COMPOUND LEAVES. 529 directions of leaves which are employed in forming the spe- cific characters of plants. When the direction of a leaf is nearly perpendicular, it is said to be erect (erecta)*; in which case it forms an acute angle with the stem (see the diagram, a. a.) : but if the angle be so acute that the upper disk of the leaf is closely pressed to the surface of the stem or the branch, it is then termed adpressa. When the angle is moderately acute only, or about forty-five degrees (.), and the surface of , the leaf, consequently, approaches the line of the horizon, the direction is termed spread- ing (patens sen patula); and horizontal (hori- zontalis sen palentissima), when it spreads still more, approximating to a right angle with the stem or the branch when these are erect (c.), or to ninety degrees in relation to the horizon. When the feaf is spreading with a drooping apex (d.) 9 the direction is termed nodding or inclined (cernua seu inclinata): it is reflex (reflexa seu recurva), when * The term vertical is also employed by some writers, but it is superfluous, the position of the most vertical leaf being sufficiently described by the word erect. VOL. I. MM 530 CONSERVATIVE ORGANS. [LECT. X. the leaf forms a curve, the convexity of which (e.) is upwards; inflex (incurva seu inflexa), when the convexity is downwards (/!); and pendulous (dependens, pendula, seu demissa), when the whole leaf droops (g. g.). When the leaf is so twisted that one part of it is vertical and' another horizontal, it is termed twisted (oUiquum, deviatum), as exemplified in oblique- leaved Garlic, Allium obliquum; and reversed (resupinatum) , when the surface which is com- monly undermost *is found uppermost, as in spotted-flowered Alstraemeria, Alstrseineria pele- grina. If $1 the leaves lean or point to the same side, as in Polygonatum mitltiftora (see fig. c. p. 246), the direction is termed unilateral (secunda seu unilateralis) . Some leaves, instead of being 1 suspended in the air, lie on the surface of the ground, as for instance in several species of the genus Plantago, and the common Daisy, Bellis perennis ; in which case the direction is termed procumbent (procumbens seu humifusaj. In aquatic plants, the direction of the leaves is determined by their relation to the surface of the water. When the leaf is raised upon its pe- tiole, above that surface, and is, therefore, an aerial leaf although growing in water, it is said to be emerging (emersa), as exemplified in the greater Water Plantain, Alisma Plantago: if it lie on the surface of the water, and the up- LECT. X.] DIRECTION OF THE LEAVES. 531 per disk of it only be exposed to the light and air, it is termed floating- (natans), as in the White Water Lily, Nymphaea alba, floating Wa- ter Plantain, Alisma natans, &c. : and when it is surrounded on every side by the water, as the majority of leaves are by the air, its direction is denoted by the term sunk or immersed (submersa, immersa, demersa), as in perfoliate Pond-weed, Potarnogeton perfoliatum. Some aquatics have leaves which are immersed and also leaves which float, as, for instance, Various-leaved Crow-foot, Ranunculus aquatilu, the beautiful white flow- ers of which decorate our ponds in the spring and summer months: others, as verticillate Water-milfoil, have both immersed and emerging leaves. In these and similar plants, it is curious to remark the manner in which the form of the leaf is modified by the medium in which it is naturally placed: the leaves, for instance, of Ranunculus aguatUis, which grow under the water, are divided into capillary segments (a. a. fig. 1, page 532), while those which float on the surface are merely lobed and notched (b.J. In plants even which are not aquatics, but which may happen to be planted in water, we perceive the me- tamorphose from the flat to the capillary leaves taking place in the fresh shoots before they gain the surface of the water, after which they assume M M 2 532 CONSERVATIVE ORGANS. [LECT. X. the form consonant to the natural habitat of the plant: a, fig. 2, is the natural form of the leaf of Horehound, Ballota nigra ; when it grows on dry land; b. h. fig. 3, display the change which takes place when it grows under water. Regarding the situation of leaves in relation to the part of the plant on which they are seated, they may be referred to the root, the stem, and the branches, or they are radical, caulinar, and rameal. A radical leaf (folium radicals) is seated upon or springs directly from the root, as exem- plified in the Primrose, Primula vulgaris, the Dandelion, Leontodon taraxacum, &c. ; but ra- dical leaves must not be confounded with seminal leaves, which are the first leaves of the majority LECT. X.] SITUATION OF LEAVES. 533 of plants proceeding from seeds that have more than one seed-lobe. The seminal leaf (/'. semi- note) is, in fact, a transformation of a seed-lobe to a leaf of a very temporary duration, and which performs the functions of a leaf until the real leaves appear, after which it drops off. Although it may be anticipating the remarks we have afterwards to make on the nature and functions of the cotyledon, yet it is proper to state here, that the cotyledon is a nutritive organ, con- taining in its cells a large portion of farinaceous matter, which becomes saccharine during the germination of the seed, and is admirably adapted for the nourishment of the embryon plant. In those seeds, the cotyledons of which rise above the ground when they vegetate, as, for instance, the Lupine, the cotyledons acquire a green colour as soon as they are exposed to the light; and then, besides continuing to supply nutriment to the young plant, the root of which is still incapable of taking up any thing from the soil, perform all the functions of the real leaf. If the seminal leaves be destroyed before the other leaves appear, the plant dies ; and, therefore, as the saccharine qua- lities of the seminal or cotyledon leaves in the Turnip attract a species of small beetle, which does not attack the proper leaves of the plant, whole crops of this useful vegetable are often destroyed. Farmers, indeed, do not consider the crop of tur- MM3 534 CONSERVATIVE ORGANS. [LECT. X. nips safe until the second leaf appears, or, in the language of agriculture, until the plants come into rough leaf'. The seminal leaves, in almost every instance, are readily distinguished by their form, which al- ways varies more or less from that of the proper leaves; thus, in fig. 4 (page 532), a. shows the co- tyledons changed into seminal leaves, b. b. are the first proper leaves of the plant. A stem leaf (f. caulinare), as the term implies, grows upon the stem, and is attached to it either mediately, or immediately by means of a petiole. A branch leaf (f. rameum) is described as such only when it differs from the leaves on the main stem of the same plant, in which case it is requisite to distinguish it from them ; as exemplified in purple Cow-wheat, Melampyrum arvense. When stem or branch leaves are seated on joints, they are designated by the term articular (articulares) , Fig. 5. Fig. a ; and when situated close to or between the LECT. X.] SITUATION OF LEAVES. 535 flowers, they are termed floral (folia floralia), Fig. 6. Elementary writers usually distinguish these floral leaves, which are green and resemble the other leaves of the plant, from those which are of a different shape and colour, giving to the latter the name of bractes (bractece), and place them among those organs which I have yet to describe under the term appendages. Both varieties, how- ever, are real leaves, having the same anatomical structure, and differing in shape not more from one another, and from the other leaves of the plant, than these last differ among themselves. That colour is a bad cause of distinction is evident, for many plants have all their leaves coloured; and in purple-topped Clary, Salvia Horminum, and .many other plants, the transition from common leaves to green floral leaves (a. a. a. a. fig. 6), and from these to coloured (b. b. b. fig. 6), is so gra- dual as clearly to display their close affinity; and to prove that all the three kinds are merely modifi- cations of the same organ. The early Honeysuckle, Lonicera Caprifolium; Green Hellebore, Helle- borus viridis; and several of the Orchis tribe, af- ford examples of green floral leaves: the Laven- ders, Lavandulce ; crested Cow-wheat, Melampy- rum cristatum ; Purple bracted Monarda, Monar- da media, &c. display specimens of those that are coloured ; and several of the Sages, Salvise, M M 4 536 CONSERVATIVE ORGANS. X. exhibit both kinds. The floral leaves form a tuft above the flowers in the Crown Imperial, Fritillaria Imperialism above those of the Pine Apple, Bromelia Ananas, and of several other plants; and under the flowers of Anemone, they re- semble and have received the name of an involucre (involucrum) . All these, and such-like, are there- fore true leaves; but as some are more inti- mately connected with the flower, and in con- formity with the custom of other elementary wri- ters, I shall revert to the consideration of these as bractes, when we examine the floral ap- pendages *. Leaves, besides differing in situation, are also variously distributed on the stem and branches. * Linnaeus remarks, that no bractes have been observed in *ny plant belonging to the class Tetradynamia. LECT. X.] POSITION OF LEAVES. 537 The position is termed opposite (opposita),(6.) when they appear directly on opposite sides of the stem, in pairs; and when these alternately cross each other they are said to be decussated (decussa- ta), (7.)- The position is ternate (terna), when the leaves stand by threes around the stem, on the same plane of the horizon ; and quaternate (quaterna) when in fours ; which, however, is not applicable when the four leaves, lying in the plane of the horizon, point towards four opposite di- rections, the term cruciate (cruciata seu crwa*- formis), 8, being employed to denote this posi- tion. When the number of leaves grounding the stem or branch exceed four, and point to different directions, forming a starlike figure, the position is termed whorled (verticillata seu stellata), (.). In the majority of instances the term whorled is used without any reference to the number of rays; but in large natural genera, as, for instance, Galium (Bed-straw), it is necessary to designate these, and for this purpose the terms quina, sena, octona, &c. are employed. When, in- stead of being in pairs, leaves stand solitarily on the stem or branches, spreading in various directions, the position is termed alternate (alterna), (10, page 538); and in this state they form either a spiral line around the common axis (spiralia), as in the Norway spruce fir, Pin us Abies , or are irregularly scattered (sparsa); or they are two- 538 CONSERVATIVE ORGANS. [LECT. X. ranked (disticha), (1 l), which implies that they spread " in two directions, and yet are not re- gularly opposite at their insertion," as in the Yew, Taxus baccata, &c. When several leaves spring from the same point they are said to be tufted (fasciculata) ; and the numerical terms bina (a. b.) applied when there are two leaves, terna when there are three, and quiriu when there are five, are used to denote the number of leaves in each fascicle. With respect to proximity, in the distribution of leaves, the term Crowded (conferta) implies that their points of attachment are comparatively very LECT. X.] POSITION OF LEAVES. 539 close; and imbricated (imbricata), (12, and also fig. l . Plate 4), that they partly cover each other, like the tiles upon a house-top : but if they do not overlap very closely, and the leaves regularly di- verge, producing a figure somewhat resembling that of a rose, the position is termed roselata (13), and crowning (coronantia) , when they terminate the stem or its divisions like a plume of feathers, as in the Palm tribe. Remote (remota) is employed solely as a term of comparison, and implies that the leaves are at greater distances from one an- other than is usual in the majority of plants. In whatever manner leaves are disposed on the stem or branches, the mode in which they are connected with these parts is termed their in- sertion; and the diversities of this are taken ad- vantage of in forming specific characters. The majority of leaves are supported on footstalks ; and those thus furnished, whether the foot- stalk be long or short, siinple or compound, are said to be petiolate (folia petiolata) ; but if there be no footstalk, they are termed sessile (sessilia). There are several varieties of sessile leaves : thus, if the leaves clasp the stem with their bases, they are termed embracing, or clasp- ing (amplexicauliaj, (14, page 540); if the em- bracing leaves be opposite and united at their bases, they are said to be connate (connata), (15, page 540) ; and connato-perfoliata, ( \ 6, and fig. 6. 540 CONSERVATIVE ORGANS. [LECT. X. Plate 4), if the union be in the whole, or nearly the whole breadth of the leaves, so as to give the two leaves the appearance of being united into but one leaf. Connate leaves are .in some instances united by a membrane, which stretching from the margins of the opposed leaves, near the base, forms a kind of pitcher around the stem ( 1 6 *) in which the rain-water is retained ; as exempli- fied in Fuller's Teasel, Dipsacusfullonum, which has received its generic name SAJ/axof, or thirsty, from this circumstance. A perfoliate leaf (fo- lium perfoliatum) , (17), is itself perforated by the stem. When leaves embrace the stem with their bases, so as to enclose it as with a sheath, they are termed sheathing (vaginantia), ( 1 8, a LECT. X.] POSITION OF LEAVES. 541 the stem, b. b. the sheathing leaves) ; they are equitant (equitantia), when being opposite they clasp each other, as in the Iris tribe ; and de- current (decurrentia), when the lamellar part of the leaf runs down on each side of the stem (19. in ^ which a. a. mark the leaves, and b. b. their de- current portions). Some succulent leaves ap- pear as if they were unconnected with the stem, and merely resting upon it ; on which account they are termed loose (soluta). Leaves which produce spines in the same man- ner as the stem, are termed spiniferous (spini- fera), (21, page 542): when they give birth to other leaves, they are said to be foliferous (folii- fera), as exemplified in Duckweed, Lemna tri- sulca; and floriferous (florifera), if they bear flowers, as, for instance, on the petiole in Tur- nera cuneiformis, (22, page 542), on the upper disk of the leaf in Butcher-broom, Ruscus acule- atus, (23, page 542): and from the serratures of its margin, in Xylophylla (24, page 542). When they 542 CONSERVATIVE ORGANS. [LECT. X. throw out roots, and produce plants in every respect resembling the plant to which they belong, they are termed proliferous (prolifera). The plant which displays the most beautiful example of the proliferous leaf is the Cotyledon calydnum; see fig. 1. Plate 10, in which the letters a. a. a. a. a. a. mark the young plants springing from the serratures of the leaf, some of them sufficiently advanced to admit of their being separated from the leaf, and sustained by their own vegetative powers. The attachment of the young plants is extremely slight., and evidently effected by a cord of umbilical vessels, originating in the proper vessels of the leaf. Leaves of this description have the closest affinity with those bulb scales which, as I demonstrated to you in treating of bulbs, produce new bulbs when they are separated LECT. X.] LEAVES. 543 from the stem which bears them, and are planted. The young plants produced on such leaves are the lateral progeny of the adult plant, by which its existence is renewed, without the aid of the sexual stimulus; in the same manner as buds are formed on stems and branches, offsets on bulbs, and gems on tubers, to the last of which, indeed, such leaves are closely allied. They possess the same distinct vitality as tubers; for the leaves of Co- tyledon calydnum will produce perfect plants from their serratures, if even worn in the pocket, wrapped in a piece of paper; and, holding, as they do, an intermediate place between tubers and leaves, they might be almost regarded as foliaceous tubers. It may be asked, are the germs of the plants, which rise on the margins of such leaves, coexistent with the origin of the leaves, as is the case with the germs in the eye or gem of the tuber? To answer this question we must have recourse to the microscope ; and on examining with it the margins of those leaves of Cotyledon calycinum, which have not yet become proliferous, we perceive in each serrature a small rough papilla, projecting on the under disk of the leaf; and in a transverse section the cellular matter at this place is seen to be differently ar- ranged from that in the rest of the leaf; and displaying in its centre a small opaque point, from which a vessel extends to the surface of the 544 CONSERVATIVE ORGANS. [LECT. X. papilla, while another enters the surrounding cel- lular matter. This point is the germ of the new plant ; and the vessel, or rather fascicle of vessels, entering the cellular matter is its umbilical cord. As soon, therefore, as circumstances favourable to its evolution are present, the same vital actions commence in it to promote its growth, and for the formation of new parts, as occur in gems situated on tubers; the leaf, like the tuber, supplying the necessary nutriment, until the roots of the young plants are capable of taking up the nu- tritious fluids of the soil; after which it decays, its assistance being no longer required. In tracing the progress of the germ, we perceive that the radicles shoot out from the surface of the rough papilla ; while the two first leaves of the young plant, imperfectly developed and applied face to face, push out from the edge of the ser- rature on the same plane as that of the surface of the leaf; but the young plant afterwards turns up- wards, so as to rise and stand erect on that surface. These two primordial leaves are always developed, and sometimes expanded, before the radicles make their appearance. But the real proliferous leaf, such as that of Cotyledon calycinum, must not be confounded with leaves which became pro- liferous only when separated from the tree and planted in the ground, under favourable circum- stances; as, for example, those of the Lemon tree ; 2 LECT. X.] ANATOMY OF LEAVES. 545 which are never productive while they remain attached to the parent. These leaves, however, closely resemble the scales of the scaly bulbs; each scale of which produces a young bulb when it is separated from the others on the same caudex, and planted, although one new bulb only is the result of the united functions of all the scales, when they are allowed to remain as aggregated parts of the adult bulb: and the explanation which has been already given (see pages 176-177) of this circumstance, as it occurs in bulbs, is appli- cable to its occurrence in this description of leaves *. * It is rather extraordinary, that although the inhabitants of the Radick islands, lately discovered in the Pacific Ocean (see Kotzebue's Voyage J, are aware of this fact, and constantly raise their Taro, Arum esculentum, by planting the leaves ; yet, it was unknown in Europe until the middle of the seven- teenth century ; when Agostino Mandirola, an Italian minorite of the Franciscan order, first described the art in a small work entitled, Manuale de Giardinieri, published at Venice, in the year 1684?. The leaves he planted were those of the Cedar and the Lemon, and the mode of conducting the operation, which he thus describes, is, I believe, still followed : " Ho " preso un vaso pieno di buonnissima terra sottile e grassa, poi " intorno all orificio vi ho posto le foglie con il gambo sotto terra " tanto che resti meza la folia sopra; posciaho fatto un' orcio- " letto d' acqua che a stilla inaffiasse esse foglie, almo do detto di " sopra, aggiongendovi sempre terra nel scavo dell acqua, ed in " tal modo hanno fatto presa, e gettato fuora le vergelette " in breve tempo." For other particulars on this subject see Beckmans Hist, of Inventions, vol. iii. p. 426. Trans. VOL. I. N N 54(5 CONSERVATIVE ORGANS. [LECT. X. The size of leaves differs very considerably in different species of plants ; but, as has been cor- rectly remarked, " it is not always the largest " plant that has the largest leaf*." The Burdock, Arctium lappa, produces larger leaves than any other British plant ; but these seldom exceed three feet in length, and two and a half in breadth ; whereas the leaves of the Banana, Musa sapien- tium, are sometimes found ten feet in length by two in breadth ; and those of the Talipot Palm, Licuala spinosa, have been known to exceed thirty feet in circumference -^. Having now concluded our examination of the external or physiognomical characters of leaves, we have next to investigate their internal structure or anatomy. To pursue this inquiry, however,. with advantage, some mode of classifying leaves, in reference to their structure, should be adopted. As all leaves have more or less affinity, in point of structure, to the stems on which they are produced, we might adopt the same arrangement that we followed in our inquiries into the nature of stems, and examine them as they belong to acotyledonous, monocotyledonous, and dicotyledonous plants ; but the varieties of structure which each of these * Keith's Syst. of physiological Bot. i. p. 37. t These leaves, which are fan -shaped, when propped up on one side with a pole, serve as a temporary hut or shade to the natives of Ceylon, who sell their merchandise under them. LECT. X.] ANATOMY OF LEAVES. 547 classes present, and the difficulty of pointing out the general features which should be re- garded as the fixed characteristics of each class, present too many difficulties to permit the adop- tion of this arrangement. Equal difficulties occur to render objectionable any classification founded on the form, or the substance, or the arrangement of the parts of leaves. The only plan, therefore, that we can adopt to render our investigation me- thodical, is to discover in what circumstances all leaves agree; and to examine the nature of these, with their modifications in the three great divisions of plants which we have just noticed. In the most cursory examination of the ma- jority of leaves, we perceive that these organs are composed of three distinct parts : one part, firm and apparently ligneous, constitutes the framework or skeleton of the leaf; another, succulent and pulpy, fills up the intermediate spaces of this framework; and a third, thin and expanded, incloses the other two, or forms the covering for both surfaces of the leaf. On a closer examina- tion we find that the first of these parts is vascular, the second cellular, and the third a transparent cuticular pellicle. Admitting, therefore, that these parts are present in every leaf, although we may not be able to discover all of them distinctly, owing to the imperfection of our instruments; we may conduct our inquiries into the structure of NN 2 548 CONSERVATIVE ORGANS. [LECT. X. leaves in reference to their vascular, their cellular, and their cuticular systems. a. Of the vascular system of leaves. Among fallen leaves which have been exposed to the action of the atmosphere in a damp place, or which have dropped into a pond, we generally find some in which the cuticle and pulp are com- pletely destroyed; whereas the ribs or veins, as they are commonly but erroneously termed, being less susceptible of decomposition, remain almost entire, and display the appearance of a beautiful tissue of network, more or less complicated. This is the vascular system of the organ, and the leaf in this state is termed a skeleton leaf. Leaves are frequently artificially made into skeletons by ma- cerating them in water until they begin to putrefy, when the cuticle is easily separated by gently rubbing and pressing them ; and the pulp washed out from between the meshes of the vascular net- work by rinsing in water : and if the operation be carefully performed, the most minute cords of vessels may be preserved * (see fig. 2, Plate 10). * Although skeleton leaves produced by spontaneous de- composition must have been very early observed, yet they were not artificially prepared until 164?5, when Marcus Aurelius Severinus published a figure, with a description of a leaf of Cactus Opuntia reduced to a skeleton. The art, however, was little attended to, until it was revived by Ruysch, in the com- mencement of the eighteenth century. This naturalist at first prepared them, by covering the leaves with insects, which ate away the pulpy part ; but as these anatomists, or satellites, to LKCT. X.] ANATOMY OF LEAVES. 549 These preparations enable us to trace more readily than in the natural leaf, the divisions, subdivisions, and various ramifications of the vascular fasciculi; but beyond this they afford us no information, and we must have recourse to the microscope to obtain a satisfactory knowledge of the vascular structure of leaves. If we commence our investigation with the simplest description of plants, the Lichens and the Mushroom tribe (Fungi) for instance, we perceive, even by the assistance of the best glasses, scarcely any trace of a vascular structure, the whole plants appearing to be little more than an aggregation of cellular substance enclosed in a cuticle. This ap- pearance, however, arises in some degree from the transparency of the vessels preventing them from being distinguished from the cells, and in some de- gree from the simplicity of their structure ; for, as the fluid they convey is not required to be raised to use his own expression, sometimes made sad havoc with the solid parts also, he soon dismissed them from his service, and employed the method I have described in the text (see his Adversariorum Decas tertia, Amst. 1723-40). Fruits were also prepared by the same method ; and the description of the in- terior structure of a pear, by Du Hamel, illustrated with engravings, may be seen in the Memoires de V Academic des Sciences, An. 1730-32. In the Philosophical Trans. 1730, No. ccccxiv. p. 371, Francis Nicholls gives an account of the ske- leton of a Pear leaf, the network of which he split into two equal layers. N N'3 550 CONSERVATIVE ORGANS. [LECT. X. considerable heights, as in the more perfect plants, the conducting tube is consequently more simple. If, however, we take a plant in which the vessels convey a colourless fluid through a coloured cellular structure, as, for example, Mar- chantia polymorpha, we find that the surface of the lobes of the leaf-like frond, when examined by an ordinary lens, is reticulated by depressed lines, within each of which a small nipple-like body rises. When a thin slice (1. c.) of a lobe is placed under the microscope, these lines are discovered to be occasioned by vessels which run immediately under the cuticle, anastomosing with one another, as represented at 1. d. d. d. d.; and the papillary bodies are seen to be cuticular pores surrounded by a slight elevation of the cuticle (1. b. b.) modified as shall be afterwards described. This vascular network is formed by a single porous tube, branching and anastomosing so as to form irregular, lozenge-shaped meshes, which are filled with a dark-green cellular parenchyma. LECT. X.] ANATOMY OF LEAVES. 551 The vessel itself is closely connected with the cellular matter, and when separated (see 2, a highly magnified representation), bears the marks of the cells on its sides. We find nearly the same vascular structure in the Mosses. Thus, if we take a leaf of broad-leaved Bog-Moss, Sphagnum obtusifolium. as a specimen, we find it, when magnified as at 3 (b. shows the natural size of the leaf), consisting of a midrib (a.), on each side of which are many oblong cells, arranged apparently in straight lines : but, under the microscope, these oblong cells are seen to consist of elevated cuticle (as at 4. b. b. b. b. b.) in the meshes of a reticulated vascular system (4. a. a. a. a.) 3 too minute to al- low of the nature of the vessel to be made out by the aid of the highest magnifying powers. It is probably the same as that of the vessels of Marchantia. The leaves of all the Mosses are sessile, although many of them are sheathing; and most of them are furnished with a midrib; but their minuteness prevents any certain informa- tion being obtained as to the manner in which the leaves receive their vessels from the stem, or whether there be a distinct set of returning vessels : they appear to be merely a continuation of the vessels of the cortex of the stem. Sprengel asserts that in the Sphagnum before us, both the cells of the leaf and those on the surface of the N N 4 552 CONSERVATIVE ORGANS. [LECT. X. stem contain a fine spiral fibre; which, however, I have not been able to discover. Proceeding to the next division of plants, those produced from monocotyledonous seeds, we ob- serve the costae or vascular fasciculi distinguishable by the naked eye; of different sizes, and running in gently curved or nearly straight lines, either from the base to the apex, or transversely from the midrib to the margin of the leaf. The former is found chiefly in those leaves which have no de- cided petiole, but spring directly from a bulb or a tuber; the latter in those which are petiolated. We shall examine each kind separately. A bulb leaf of the White Lily, Lilium candi- dum, may be taken as an example of the general distribution and character of the vascular system in the first description, the sessile leaves of mono- cotyledonous plants. On examining it, we find that the vascular framework consists of a distinct midrib, which forms the keel of the leaf, and of less elevated ribs (costce), that extend on each side of the midrib in longitudinal lines, which form a gentle curve, following the shape of the leaf. In the smoother and more succulent leaves of this division, however, these costae are scarcely visible externally, or at least appear merely as striae on the surface of the leaf: and this is the case, also, as far as regards many of the smaller vascular fasciculi, even in those leaves, which have LECT. X.] ANATOMY OF LEAVES. 553 prominent costse. If we now make transverse and longitudinal sections of the Lily leaf, we perceive that the costae are composed of fasciculi of spiral vessels closely accompanied with corresponding fasciculi of proper vessels, and embedded in cel- lular substance; or, that the leaf has a double system of vessels, one for conducting forwards the sap, and the other for returning the proper juice into which the sap has been changed by the functions of this organ. In the transverse sec- tion, these vascular bundles appear like dots upon the divided surface ; and, when magnified in trans- mitted light, display their twofold nature by dif- ference of transparency; the part of each fasci- culus composed of spiral vessels being particularly distinguished by a greater degree of opacity, owing to the spiral thread which composes the coats of these vessels being firmer and more opaque than the coats of the proper vessels. Thus, in the midrib (5. a.), we perceive that the three spots 554 CONSERVATIVE ORGANS. [LECT. X. formed by the vascular fasciculi, which it en- closes, have each one half darker than the other, and the darker half is turned towards the upper disk of the leaf. The same circumstance cha- racterizes the fasciculi enclosed in the costse (5. b. b. b. b.) ; and also those of the intermediate or secondary ribs, which form no prominence on either surface of the leaf. By the aid of more powerful glasses, the distinct character and dis- position of both kinds of vessels are perfectly per- ceptible: and in placing the principal fasciculus of the midrib, detached from the others, under the microscope, this is not only satisfactorily demon- strated, but we become acquainted, also, with the fact, that the spiral vessels (6. a. page 553) of the leaf as well as those of the stem, are found ge- nerally empty, like the arteries of animals; while the proper or returning vessels (6. b.) are always full. I may here remark, that the closer proxi- mity of the spiral vessels to the upper disk of the leaf is common to the majority of leaves ; for, in- dependent of the fact, that the chief function of these organs, namely, the exposure of the sap to the light and air, would lead us, a priori, to conclude that the vessels carrying forwards the sap must, necessarily, be on that side of the leaf most exposed to these agents; the sap- vessels receiving their origin in the stem from the vessels of the albur- num, and the returning vessels terminating in LECT. X.] ANATOMY OF LEAVES. 555 those of the bark, the disposition could not well be otherwise, seeing that the relative position of the upper and under disk of every leaf, to the centre of the stem, is exactly that of the albur- num and the bark. In leaves, however, which stand vertically, or have no distinction of sur- faces, the situation of the spiral vessels is either the reverse, or in the centre of the entire vessels : anatomy thus confirming the idea of the close affinity of such leaves to stems. I have already stated that the bundles and threads of vessels, in leaves belonging to this di- vision of the class of leaves under consideration, run in longitudinal lines. These are not exactly parallel, but approach both at the base and the apex of the leaf; and, also, communicate la- terally in their course by small threads, given off at irregular intervals. This structure is easily de- monstrated by placing a small slice of the Lily leaf (7, page 553), cut immediately withhi the cu- ticle of the upper disk, under the microscope. We perceive that the longitudinal bundles (a. a.) are united by transverse threads (b. b.) ; and this is the case at irregular intervals throughout the whole extent of the leaf. These transverse fasciculi, however, are not threads simply detached from one longitudinal layer, and coming into contact with another, as was supposed by Dr. Grew to be the case in all 556 CONSERVATIVE ORGANS. [LECT. X. leaves*; but are distinct vessels, uniting with the longitudinal bundles in a singular manner, as is apparent (8. a. page 553) in a very highly mag- nified view of one of the angles formed by a transverse fasciculus and a longitudinal vascular cord. One of the vessels appears to belong to the longitudinal fascicle ; but the other termi- nates there, and has its extremity applied to the side of one of the vessels forming a part of the longitudinal fascicle. Whether there is any open- ing directly into the longitudinal vessel on which the extremity of the transverse vessel is applied I have not been able to determine. From these demonstrations we may conclude that the vas- cular system of the sessile leaves of monocoty- ledons, consists of fasciculi composed of spiral vessels accompanied with proper vessels which are not spiral, arranged in longitudinal lines, and connected by smaller transverse threads; the whole forming a reticulated texture with irregular rhomboidal meshes. The longitudinal vessels are a continuation of those which are nearest to the surface, in the root caudex, or the stem from which the leaves immediately spring; and thus * It is bold to dissent from such authority on the subject of vegetable anatomy ; but the improvement of microscopes since Dr. Grew's time, has enabled many Phytologists of very in- ferior abilities to that great man, to point out some errors into which he had been led by the imperfection of his instruments. LECT. X.] ANATOMY OP LEAVES. 557 the greater number of the circles of the distinct fasciculi, which compose the stems of monocoty- ledons, terminate in leaves until the plant attains its ultimate growth. In order to examine properly the vascular framework or skeleton of the petiolated leaves of monocotyledonous plants, we must arrange them under two subdivisions. 1. Those in which the ribs run longitudinally, or in a direction from the base to the apex of the leaf: and, 2. those in which they run nearly transversely, or in a direction from the midrib to the margin. 1. In this subdivision we perceive that, in the Grasses, the vascular fasciculi resemble, very closely, those of the former division: the ribs being in longitudinal, nearly parallel lines, con- verging towards the apex of the leaf; and united at irregular distances by obliquely transverse threads. If we take a stem leaf of Indian Corn, Zea Mays, as a specimen, we perceive the petiole, which is broad, expanded, and sheathing, deriving its origin from the whole circumference of the knot of the articulation which produces it; dilat- ing gradually as it rises upwards, until its edges become a thin fimbriated membrane, and again contracting, but less gradually, at its upper part, or where it is united to the expansion of the leaf. The vascular bundles, which can be readily traced by the naked eye, are composed of the two distinct 558 CONSERVATIVE ORGANS. [LECT. X. kinds of vessels described in the former division, which appear as dots in a transverse section of the petiole situated almost close to its external sur- face; or as represented at a. a. a. a. (g.), a highly magnified view of the section. The number of the spiral vessels in each fascicle is generally six, three large and three smaller, symmetrically arranged, as may be seen in a transverse section of one of the fasciculi (10. a.) viewed under the microscope ; and the whole surrounded by a mass of much denser cellular matter than the rest of the substance of the petiole. The returning or proper vessels (10. ft.) are much smaller and more numerous than the spiral; and are congregated into a bundle which occupies a space close to the former, between it and the cuticle, and is bounded by a mass of the same dense cellular matter (10. c.) as that which LRCT. X.J ANATOMY OF LEAVES. 559 surrounds the spiral vessels : the object of which is, probably, to give such a degree of firmness to the petiole, as will enable it to sustain, in the erect position, the expansion of the leaf. If we now make a vertical section of the petiole, so as to divide one of the fasciculi longitudinally, in the thickness of the petiole we perceive that the larger vessels are regular spirals (1 1. e.), furnished with diaphragms (h.) at certain distances, the struc- ture of which, however, we shall perhaps never be able to ascertain, owing to the minuteness of the parts ; the diameter of these vessels, although comparatively large, not exceeding ^-^ of an inch. In this section, also, we perceive that the proper vessels (l 1 .f.) are membranous and porous: and that the cells in immediate contact with both sets of vessels are oblong ( u . d.) ; whereas those (g.) which are between the proper vessels and the cuticle of the outer surface of the leaf, and which form the elevated portion of the costa, although they are not oblong, yet, differ both in size and in regularity of structure from those (n. c.) that form the inner substance of the petiole. Close to the cuticle of the upper disk ( 1 1 . a.) is a mass of oblong cells (b.) resembling those in contact with the vessels. Ascending to the expansion of the leaf, which is separated from the petiole by a semitranspa- rent white, condensed, membranous space (see cut 560 CONSERVATIVE ORGANS. [LECT. x. 12. c.), from which the expansion of the leaf spreads out like a shoulder on each side ; we per- ceive that the midrib (12. b.), which is not distin- guishable in the lower part of the petiole (12. a.), becomes very conspicuous on the under disk at this point; forming almost a knob, which passes into a striated ridge, and extends, gradually dimi- nishing in size, to the apex of the leaf. From ten to twelve parallel costae are visible on each side of the midrib, 22 t> .Jill JL:9L which when magni- fied appear like white parallel lines (13. a. a. a.) , running through the green smooth sub- stance of the expan- sion, and taking the curve of its shoulders as if originating in the white semitransparent space (12. c. c.) al- ready described. But between these costse there are several smaller vascular cords, which are scarcely visible on the surface, neither producing elevation nor difference of colour ; and which can be demonstrated only on the dissection of the leaf. One of the more obvious distinctions, therefore, in the structure of the petiole and the expansion 2 LECT. X.] ANATOMY OF LEAVES. 561 in the leaves of the gramineous tribe of plants is, that, in the petiole, the vessels run in distinct fas- ciculi, which are all nearly equal in point of size (14.) ; whereas in the expansion the fasciculi differ considerably in size ( 13.), the larger only being very visible on the surface. In both, there are transverse threads which connect the longitu- dinal bundles, and these are conspicuous even to the naked eye in the more succulent leaves, par- ticularly in those which involve the fructification of the Mays (15.), when viewed by transmitted light. In examining a transverse section of a portion of the expansion, of the leaf of Indian Corn, con- taining one of the visible costae and the interval between it and the next costa; we immediately perceive the difference of structure in the two kinds of fasciculi. The visible costa consists of two large spiral vessels on the same line, and a com- pact fasciculus of proper vessels on each side of the line of spirals, towards both surfaces of the leaf (c. c. fig. 13, Plate 10); while, in the interval, each fasciculus is composed of one small spiral ves- sel only, surrounded with a circle of proper ves- sels, and placed in the heart of the substance of the leaf (d. d. d. fig. 13, Plate 10). But, be- sides these, there is another kind of fasciculi, two of which are generally observed in each space between the visible costse, connected with a pe- VOL. I. 00 562 CONSERVATIVE ORGANS. [LECT. X. culiar cellular apparatus to be afterwards de- scribed. These appear to be modifications of the two vascular fasciculi already noticed ; having the same structure as the obscure or invisible fascicu- lus, and the accompanying compact bundle of pro- per vessels of the visible costa (see e. e. fig. 13, Plate 10). In a section obtained by slicing the leaf, we find all these fasciculi united by trans- verse threads, forming rhomboidal meshes, si- milar to those which have been already described. But although the arrangement of the vascular system of the leaf of Indian Corn, just described, may be taken as a specimen of that peculiar to the leaves of all the Grasses; and to those leaves of monocotyledonous plants which are petiolated, and furnished with longitudinal costse, yet, there must necessarily be many modifications of this arrange- ment. I shall, however, particularly notice one only, that which characterizes very thick and fleshy leaves; such, for instance, as those of the Aloe tribe. On the exterior surface of the leaf of the Aloe, and other leaves of a similar character, we per- ceive no appearance of a vascular system ; but on attempting to tear the leaf across, considerable resistance is opposed to the effort, by a number of tough, longitudinal fibres. These constitute the vascular fasciculi ; and in a transverse section of the leaf, we find that they run through its sub- LECT. X.] ANATOMY OF LEAVES. 563 stance, nearly in the same manner as the fasciculi in solid monocotyledonous stems. Viewing the sec- tion (16.) through a magnifying lens, we perceive that the fascicles are of different sizes, the largest being in the centre: and, in a portion of it (17.) placed under the microscope, we perceive that the central fasciculi (a.) differ from the others (&.), in their structure as well as size, being composed of a dense cord of spiral vessels, accompanied by a very large bundle of proper vessels, which is sepa- rated from it by a thin layer of cellular substance ; whereas the smaller or marginal fascicles appear to be entirely composed of proper vessels. This structure of the central bundles is still more deci- dedly observed, in a very thin longitudinal slice, o o 2 564 CONSERVATIVE ORGANS. [LECT. X. (18.) cut in the thickness of the leaf, and viewed in the microscope by transmitted light. The cord of spiral vessels (a.) is a few proper vessels sur- rounded by , and a layer of cellular matter in- terposed between it and the large fascicle (c.) of proper vessels. In splitting the leaf, we per- ceive that these vascular fasciculi run in parallel lines (19.), and take the same curves as the mar- gin ; but do not enter into the spines (a. a.), which are of a hard ligneous consistence, and supplied by small threads of vessels given off from the fasciculi nearest to them. The vascular cords do not anastomose in any part of the leaf; and very few transverse threads are perceptible, compared with those which occur in the membranous leaves of this natural order. But besides the vascular fasciculi we find, in this leaf, and other monocotyledonous fleshy leaves, a system of tubular cells, which might be mistaken for vessels, situated immediately un- der the cuticle (17. d.) 9 from which they extend in straight contiguous lines to the cellular parenchy- ma, the chief substance of the leaf. These resemble beaded or moniliform vessels (les vaisseaux en chapelet of Mirbel), and appear to be composed of transparent membranous oblong vesicles, united at their extremities; and are either perforated, or contain small granules sparsely spread over their inner surface ( 1 9. a.). It is probable that these cells are part of the absorbing system by which fleshy plants, such as the Aloe, which grow LECT. X.] ANATOMY OF LEAVES. 565 in sandy, arid soils, chiefly derive their nou- rishment : for, as very little moisture is taken up by the roots, these plants are supported, almost entirely, by cuticular absorption. The natural func- tions of these tubular cells, also, being to absorb and to carry fluids towards the centre of the leaf, enable us to understand why the leaves of an Aloe, when separated from the stem, are very long in drying and losing their plumpness ; where- as, if thrown into water, when they are very much shrivelled, they almost immediately regain their original size. The absorbing mouths by which they are supplied will be demonstrated, when we examine the cuticular system of leaves. In the leaves of those monocotyledonous plants the costae of which, instead of being longitudinal, run in transverse parallel lines, forming acute an- gles with the midrib, we find that the arrangement of the vascular framework resembles that of the Grasses in some circumstances ; but differs from it in other respects. Thus the costae are parallel to one another, and communicate by small transverse cords of vessels, so as to form meshes which are rhomboidal or square according to the angles at which these transverse cords are given off from the costae, as in the Grasses. The petioles are., also, in general sheathing, and many of them are furnished with ligulse. But, in almost all of them, the peculiar cartilaginous articulation, which divides the petiole oo3 566 CONSERVATIVE ORGANS. [LECT. X. from the expansion in the Grasses, is not present ; and the petiole assumes a stalk-like aspect before it reaches the expansion. Taking this leaf of Canna Indica (21.), as a specimen of the vascular system in this description of leaves, we perceive, looking at the under disk, that the midrib is much elevated near the base, and gradually diminishes in size, until it appears little more than a mere line at the apex of the leaf. The more elevated costse (a. a. a. a. a.) are the primary vascular fasciculi ; and between these are secondary fasciculi, which are less elevated. To the unassisted eye they all appear to go off from the midrib ; but viewed by a magnifying lens, and with transmitted light, we perceive that all of them do not proceed directly from the fasciculi of the midrib (22. b. b. see page 567), but that some of them (22. a. a.) are branches of the others*. At * In figure 22, the cuticle, except at the space c. d. is taken off, and the midrib pared down, to show the transverse vascular threads, which unite the parallel fascicles ; and which are ren- dered very conspicuous when viewed by transmitted light, ANATOMY OP LEAVES. 567 LECT. X.] the margin they all inosculate, and form, as it were, one fasciculus which, extending from the base to the apex, is the real living boundary of the leaf. Pursuing our examination more closely ; and placing a slice (23.) of the petiole cut transversely near the base of the expansion, under the micro- scope with a glass of a moderate power, we per- ceive that the vessels are arranged in distinct fas- ciculi, which are nearly of the same size in the cen- tre of the section ; alternately larger and smaller (23. a. b. a. b.) near the circumference on the con- vex surface, or that part of the petiole which is towards the under disk of the leaf; and all small (c. c. c. c. c. c.) on the concave surface. The costae (22. e. e. e.) are continuations of those on the con- owing to the dark green colour of the parenchyma in the meshes formed by these threads. * o o 4 568 CONSERVATIVE ORGANS. [LECT. X. cave surface of the midrib, which are curved out- wards in opposite pairs, at different distances be- tween the basis and the apex of the leaf; but the central fasciculi pass on its apex. These vascular fasciculi are embedded in a cellular tissue; besides which the petiole and midrib of this description of leaves contain peculiar pneumatic or air cells (23. d.d. d.) closely resembling those which I shall after- wards demonstrate to you, constitute a great part of the substance of aquatic plants. In a transverse section of a small part of the expansion of the leaf (24. see page 567), we perceive that the vascular cords (a. a. a) run nearly in the centre between the two plates of cuticle, embedded in an opaque green parenchyma; and that, instead of the pneumatic apparatus of the petiole and midrib, there is a transparent layer of large cells (24. b.) immediately under the cuticle of the upper disk. I should, however, inform you that these pneumatic cells are not present in the petiole and midrib of all leaves with transverse costse, belonging to mo- nocotyledonous plants; but the same general ar- rangement of the vascular cords, and, conse- quently, the same structure of the framework, are seen in all of them. With regard to the composition of the fasci- culi; these, as in the other leaves we have ex- amined, consist of spiral and proper vessels; dif- fering, however, in the relative position of the spiral vessels, which in each fasciculus, in these LECT. X.] ANATOMY OF LEAVES. 569 leaves, are placed between two bundles of proper vessels. If we cut a superficial slice from the under disk of the leaf, where any of the fasciculi ra- mify (25. see page 567); and examine it by a powerful microscope, we may rationally conclude from what we perceive, that, although some of the costse are merely continuations of the vas- cular fasciculi of the midrib, separated like threads from a skein of silk, yet that others which also branch from these, and the smaller transverse vascular threads, are actually united by that kind of connexion which, in the vessels of animals, is termed anastomosis. Thus, in the slice under examination, we find that both the fasciculi (a. a.) contain the same number of vessels, which would not be the case, were the one parcel separated from the other ; and although the transverse thread (b.) contains one spiral vessel only, yet this vessel does not appear to be split from the larger fasciculus ; but to be simply united to it, the end of b. being evidently applied to the side of a. if it do not ac- tually open into the cavity of that vessel. Examining, by the same power of the micro- scope, a transverse section of one of the larger fasciculi of the midrib of the leaf of Canna In- dica, we find it to consist of one large, and from three to six smaller spiral vessels, arranged and relatively connected with the proper vessels in a manner closely resembling the arrangement of those in the fasciculi which are found in the 570 CONSERVATIVE ORGANS. [LECT. X. stems of White Bryony (see fig. 4, A. Plate 8). The arrangement of the vessels, however, in the fasciculi of the expansion differ, in some de- gree, from those of the midrib. Placing a trans- verse section (26.) of one of these fasciculi under the microscope, we find that it is composed of two large tangent spiral vessels (a.) surrounded by firm cellular matter ; and of two small isolated vessels (b.), which are spirals also, situated nearer to the upper disk of the leaf; the one before the other, and, in like manner, surrounded by firm cel- lular substance. The proper vessels are collected into two distinct fasciculi ; the largest of which (d.) is on the under disk or back of the leaf, and covered by the cuticle only, while the smallest (c.), which is towards the upper disk, is covered by one layer of cellular substance. In the longitudinal section (27.) made through the centre of the fasciculus, in the thickness of LECT. X.] ANATOMY OP LEAVES. 571 the leaf, e. being the upper and f. the under disk, the relative situation of the vessels is still more clearly demonstrated. We may pause here a few moments to re- mark, that the colour of the juice in the cells of the leaf, is more intense towards the surfaces, while in the centre it is scarcely deeper than that which is contained in the cells of the cutis (e.f.). Is not this probably owing to the sap which is current in the spiral vessels, and which is co- lourless, being given out laterally, and diffusing itself through the cells, in which the change effected by the air and light, that causes the co- lour is produced ? The manner in which the co- lour is diluted as it approaches the limits of the spiral vessels, would authorize such a suppo- sition ; but, wherefore it may be demanded, are the cells immediately under the cutis free from colour ? It is probable, we may reply, that these cells are filled by pure water only, absorbed from the atmosphere; and, consequently, their con- tents are unfitted to undergo those chymical changes to which the sap is subjected in the leaf. But this discussion is premature. From this demonstration of the vascular sys- tem in the leaves of monocotyledonous plants, it is evident that a general character, however variously modified in many instances, pervades the whole. The fasciculi of vessels are distinct; they run in directions parallel to one another; 572 CONSERVATIVE ORGANS. [LECT. X. and the principal bundles are united by smaller transverse cords or fasciculi; which form meshes of a rhomboidal figure, all nearly of the same size in the same leaf. Passing on to the leaves of Dicotyledons, we find the reticulated structure of the vascular framework more complex and varied, than we have found it in the leaves of the two natural di- visions^ which we have already examined. This is made evident to the unassisted eye by holding up between it and the light any newly expanded leaf ; but it is more beautifully demonstrated in the ske- leton of a full-grown leaf, carefully prepared. In a leaf of the Indian Fig, Ficus religiosus, thus pre- pared (fig. 2, Plate 10), we perceive seven princi- pal costae, springing from each side of the midrib (a.), and extending to the margin of the leaf, where they bend towards its apex, and are enarched or inosculate with one another (b. b. b. b.). From each side of these costae, a series of secondary fasciculi spring, which, inosculating, also, with their oppo- nents, form a secondary set of arches (c. c. c. c.) be- tween each pair of the principal costae ; and inclose a tissue of minute and exquisitely beautiful reticu- lations, the result of the numerous ramifications of the vascular threads, as they divide and subdivide as it were out of a greater into a less. But besides this distribution of these vascular fasciculi, we observe a fascicle (d. d.), proceeding from each LECT. X.] ANATOMY OF LEAVfiS. 573 side of the midrib, at the very base of the ex- pansion, and bounding the margin of the leaf. This receives supplies from the arches of the costse, as it passes onwards, until it unites with its fel- low at the apex of the leaf. Such is the general distribution of the vascular fasciculi in the leaves of dicotyledonous plants. It would be incon- sistent with any elementary plan of instruction, to enter largely upon the consideration of the various circumstances which modify the distribution of these fasciculi ; and, therefore, I shall examine the modifications depending upon two states only of dicotyledonous leaves : 1. When the leaf or its ex- pansion is thin or membranaceous ; and, 2. when it is thick QY fleshy. 1. The general distribution of the vascular fasciculi, in thin leaves of dicotyledonous plants, in the majority of instances, resembles that which we find in the skeleton of the leaf of the Indian Fig tree. The vessels of the costse proceed from the principal fasciculus of the midrib, and run between the laminse of cuticle, embedded in the cellular mat- ter, in cords which form visible elevations on the back of the leaf, and corresponding furrows on its face. Each fascicle consists of spiral and proper vessels throughout all its ramifications; and, in whatever manner these vessels are arranged in the fasciculi, the spiral and proper vessels are always associated and, in general, tangent. This arrange- ment is common both to sessile and to petiolated, to 574 CONSERVATIVE ORGANS. [LECT. X. simple and to compound leaves, as far, at least, as respects the expansion. In sessile leaves, how- ever, all the fasciculi do not proceed from the midrib, but some are given off directly from the stem or the branch, and enter the expansion of the leaf at its base, on each side of the midrib. In petiolated leaves, also, the petiole is generally dilated at its point of union with the branch, and at this point the vessels enter the petiole in distinct bundles ; the remains of which are visible in the eschar produced by the falling of the leaves in autumn. Thus in the Apple, the Pear, the Peach, and many other trees, the leaf is attached to the wood by three fasciculi, one of which enters the middle of the petiole, and the others on each side of it. In the Lilac, the attachment is also by three distinct fasciculi ; but there is besides a line of coalesced fasciculi which forms a kind of open crescent; and in the Laurustine the whole of the vessels pass from the wood into the petiole in one fascicle, the transverse section of which is nearly a complete semicircle. In compound leaves, the number of fasciculi passing into the petiole from the wood, is in some instances regulated by the number of the leaflets; in the Elder, we find generally five ; and in the Horse Chesnut, from five to seven or eight. It is, however, the inner part only of these fasciculi, or that which con- 2 LECT. X.] ANATOMY OF LEAVES. 575 veys the sap to the leaf, that is given off from the wood, or rather from the medullary sheath ; for the outer part, which consists of the proper or re- turning vessels, enters the bark, but not the wood. This fact is beautifully illustrated by placing young leafy twigs in coloured fluids. The colour is seen passing tip from the stem into the leaf through the upper portion of each fascicle; while that part which consists of the returning vessels remains perfectly free from colour. Such is the general arrangement of the vascu- lar framework in the thin leaves of dicotyledonous plants ; to examine the intimate structure of the fasciculi, we must have recourse to the same me- thod that we adopted in our examination of them in the leaves of the two natural divisions we have already investigated. Taking the leaf of the Lilac, Syringa vulgaris, as an example of the simple petiolated leaf; and placing a thin transverse slice of the petiole under the microscope, we find that the vessels are arranged in the following man- ner. Close to the upper or channelled surface of the petiole, we find three small distinct fasciculi of spiral vessels (c. c. c. fig. 10, Plate 8), one im- mediately within the cutis, in the hollow of the channel, and one at each side ; but the principal vessels constitute one large compound fasciculus, in the centre of the petiole, which appears of a horseshoe shape, in the transverse section ; and 576 CONSERVATIVE ORGANS. [LECT. X. consists of one fasciculus of spiral vessels, and two fasciculi of proper vessels. The spiral vessels, which form the central fasciculus (d.), are arranged in rays, which are sometimes tangent, at other times separate ; whereas the proper vessels con- stituting the two fasciculi (e. e.), one of which is situated within, and the other without the fasci- culus of spirals, are irregularly embedded in a pulpy parenchyma, and are readily distinguished by their greater transparency. The bark, or true cutis (a.) of the petiole, seems, also, to consist chiefly of several series of the same kind of proper or returning vessels. In the various modifications of this structure of the vascular system, in the petioles of dicotyledonous leaves, the radiated ar- rangement of the spiral vessels is found in all: the petiole in this respect, as well as in the other parts of its structure, closely resembling the stem or the branch from which it springs. In simple leaves, with a few exceptions, although the vascular part forms at first several fasciculi, at the base of the petiole, yet these soon coalesce into one compound fasciculus ; but in compound leaves they remain distinct. If we take a leaf of the Elder, Sambucus nigra, and place a transverse section of the com- mon petiole under the microscope, we perceive ten distinct fasciculi of vessels. Five of these (b. b. b. b. b. fig. 11, Plate 8) are compound fas- LECT. X.] ANATOMY OF LEAVES. 577 cieles, embedded in the cellular substance of the petiole; and five (a. a. a. a. a.) fasciculi of proper vessels occupying the angles of the footstalk, and situated in the bark, or at least exterior to the former and immediately within the cuticle. The compound fasciculi consist, each, of a band of spiral vessels, arranged in rays, and two fasciculi of proper vessels ; one interior, and the other ex- terior to the band of spirals, but both tangent upon it. One of these compound fascicles passes into each leaflet ; and consequently their num- ber, in the common petiole of compound leaves, generally corresponds to that of the leaflets. In compound leaves, however, which have moveable articulations, we perceive that all the separate fas- ciculi are collected into one fasciculus in the arti- culations. Thus, in the common Kidney-bean, Pha- seolus vulgaris, the petiole of which is channelled, with an articulation at the base of the common pe- tiole and, also, at that of each partial petiole, we find that the vascular fasciculi (a.a.b. b. fig. 8, Plate 10) are distinct, and form a circle situated immedi- ately under the bark in the channelled parts of the petiole ; with a considerable portion of lax cellular substance or medulla (c.), enclosed within the circle which they form : whereas, in the articulated parts, there is one central fasciculus only (a. fig. 7, Plate 10), surrounded by a large mass (b.) of very firm cellular matter. The advantage of this change of VOL. u p P 578 CONSERVATIVE ORGANS. [LECT. X. disposition of the vascular bundles, in the articu- lations, is very obvious: for, had the fascicles remained distinct, and surrounding the pith, in the articulations, those on the outside of the flexure, in every considerable motion of the joint, must have described so large a circle, as would have endangered the organization of the vessels by the extension; while those on the inner side would have suffered, equally, by the compression to which they must necessarily have been subjected. But> by the whole of the vessels being situated in the centre of the petiole, the extension and com- pression produced by the flexure, on every part of the fascicle, is not more than can be borne by any individual vessel, whether spiral or entire ; and, thence, the freest and most varied motion of the joint can be exercised with impunity. The neces- sity of such a modification of structure, in the petioles of compound leaves, susceptible of mo- tion, may indeed be inferred from the fact, that articulations are present in all those which per- form certain movements; as, for example, those which fold together their leaflets at night ; those which are endowed with the power of spon- taneously moving their leaflets, as Hedysarum gyrans; and those which fold their leaflets together when touched, as Mimosa sensitiva and pudica. Some simple leaves, those for instance of the Holly Hock, of the Geranium tribe, &c. which have several principal costse diverging from LKCT. X.] ANATOMY OF LEAVKS. 579 the summit of the petiole, and in this respect have an affinity to digitate leaves, present nearly the same vascular structure of the petiole as the compound leaves. The fasciculi are dis- tinct ; and correspond in number to the princi- pal costse of the leaf; each of which may be thus regarded as a kind of midrib, and the leaf as com- posed of a number of conjoined leaflets: so that these leaves, although they are necessarily classed as simple leaves from their external appearance, yet, bear in anatomical structure the same affinity to digitate compound leaves, which the webbed foot of a bird bears to one which is not webbed. A similar structure, also, is found in the petioles of those leaves which are longitudinally ribbed, or nerved as the common expression is, from the base of the expansion, as for instance those of the genus Melastoma; but, when the ribs do not origin- ate from the base, although they are very conspicu- ous, as, for example, in the leaf of the Cinnamon tree, the structure of the vascular system of the petiole is exactly the same as in simple dicotyledo- nous leaves, which are not longitudinally ribbed. If, instead of a transverse section, we place a longitudinal section of any of these leaves under the microscope, we perceive that each fasciculus is composed of spiral and proper vessels, the same as we have already seen constitute the ribs in the leaves of monocotyledons. pp2 580 CONSERVATIVE ORGANS. [LECT. X. Tracing the vascular fasciculi from the petiole into the expansion, in the thin, simple leaves of dicotyledons, we find their divisions, subdivisions, and ultimate ramifications much more diversified and minute than in the leaves of monocotyledons. Whatever may be the origin of these divisions and subdivisions, whether they proceed from one central fasciculus, or from several longitudinal costae, the ramifications become smaller and smaller, owing to a diminution of the number of the vessels which they contain ; but not owing to any diminution of the diameter of the vessels themselves: for, although a principal fasciculus may contain larger and smaller spiral vessels, yet the general comparative magnitude of the vessels, in the smallest fasciculus, is the same as in the largest. This question, therefore, again presents itself: Do the vessels of the leaf inoscu- late and anastomose, or are the smaller fasciculi merely separations from the larger? Dr. Grew, as I have already stated, denied that they ever inosculate or anastomose until they arrive at their final distribution. In appealing to nature for a solution of this disputed point, we find Grew's opinion so far correct, that the vascular fasciculi of the costse, which are given off from the midrib, are separations from the petiolar fasciculi in their progress towards the apex of the leaf; and that the fasciculi forming some of the secondary LECT. X.] ANATOMY OF LEAVES. 581 costae, also, are separated in a similar manner. But in the smaller ramifications, we perceive that many of the fasciculi are connected with each other at nearly right angles; and in these in- stances the vessels are not separations from the larger fasciculi, but are distinct and merely applied in a peculiar manner to the sides of those from which they seem to arise ; as can be readily demon- strated by dissection, with the aid of the micro- scope. In this minute portion of a leaf of the Lilac, sliced from between the cuticles, and examined, by transmitted light, under a very high magni- fying power, we observe that, in the smaller fasci- culus (g. 28.), which is composed of seven spiral vessels, and united nearly at a right angle with the larger fasciculus (h.), three of the vessels (a.) form a curve upwards, and three (b.) a curve downwards, before they unite with the p P 3 582 CONSERVATIVE ORGANS. [LECT. X. larger fasciculus ; while the central vessel (c.) seems to terminate in a straight line on the side of the vessel 3, which is one of those composing the larger fasciculus. In this case it is evident that the smaller fasciculus is not a separation from the larger ; but is joined to it by a species of anasto- mosis ; which; in the central vessel, is effected in a direct manner by the application of its extremity to the side of the vessel on which it terminates; and, in the other six vessels, in a less direct man- ner by the lateral application of a portion near the extremity of each vessel, before it curves outwards to the particular vessel on which it terminates. In other leaves, as in those of the garde'n Lettuce for instance, in which single vessels are often united to large fasciculi (a. b. c. 29. page 581) and to other single vessels (a. b. c. 30. page 581), the nature of the first species of anastomosis, just described, is still more perceptible. One of these examples (28.) demonstrates, also, that the proper or re- turning vessels (d. e.f.f.) unite in the same man- ner as the spiral. Whether the communication of the cavities of these united vessels be direct, as in the vessels of animals, so as to allow the fluids they convey to flow in a uninterrupted stream from the one to the other, I have not been able to determine. It is, however, evident that in the leaves of dicotyle- dons, as in those of monocotyledons, all the vascu- LECT. X;] ANATOMY OF LEAVES. 583 lar ramifications of the foliar expansion are not pro- longations of the vessels forming the petiolar fas- ciculi ; but that many of them are distinct vessels anastomosing with others, although in a different manner from this kind of union as it occurs in ves- sels in animal bodies. It is probable that the in- osculation which occurs in the proper or return- ing vessels, more nearly resembles that which we find in vessels of animals ; for, as the proper ves- sels are simple membranous tubes, any commu- nication between them must be by direct openings, such as are found to exist in the vessels of Mar- chantia (see page 550). 2. The thick and fleshy leaves of dicotyle- donous plants are seldom petiolated ; but when they are so, the arrangement of the vascular fasciculi, both in the petiole and in the expansion, closely resembles that of the thin membranaceous leaves. The sessile leaves of this division are ge- nerally thicker and more succulent than the pe- tiolated. If we take the genus Mesembryan- themum, as affording specimens illustrative of the character of these sessile leaves, we find that the vessels pass from the stem into the leaf in one or more fasciculi, according to the figures of the leaves. Thus in the Hatchet-leaved Mesembryan- themum (M. Dolabriforme) , the leaves of which are connate, the sap- vessels enter the leaf in one bundle, which extends in the direction of its p p 4 584 CONSERVATIVE ORGANS. . [LECT. X. axis, the whole length of the leaf, giving off in its course a few thread-like branches only at consi- derable intervals; and as this vascular fasciculus and its ramifications are situated in what may be termed the pith of the leaf, and are, consequently, imperceptible on its surface; this description of leaves appears to the unassisted eye destitute of vessels. These organs are, indeed, comparatively few in succulent leaves ; and are less necessary than in membranaceous leaves ; for, as succulent leaves either exhale very little moisture, or absorb a considerable quantity from the atmosphere by their surfaces, the nutriment of the plant, in the first case, is sufficient although the fluids taken up by the roots be comparatively scanty ; and, in the second, it is supplied, independent of that which may be furnished by the roots, by cutaneous ab- sorption. In the leaves of the broad-leaved species of Mesembryanthemum, and in similar succulent leaves, the vessels enter the leaf in several distinct fasciculi; which diverging, pass on in nearly straight lines, giving off a few bundles only in their course ; but as they approach the apex of the leaf, whatever its form may be, they divide, subdi- vide, and inosculate as in thin leaves ; and the proper or returning vessels accompany and sur- round the spirals in all their divisions. In the suc- culent leaves of dicotyledonous plants, also, we find the same system of tubular cells, between the LECT. X.] ANATOMY OP LEAVES. 585 pulp and the cuticle, which we described in the Aloe (page 564) ; and in the leaf of the Mesem- bryanthemum, under examination, we perceive these tubes commencing immediately under the cutis, and terminating generally in the cells of the central pulp ; but sometimes in follicles, which are both very irregular in form, and of very dif- ferent dimensions. It is probable that part of the fluid taken up from the atmosphere passes at once into the central cells, the contents of which are colourless, while another part remains in the tu- bular cells, and undergoes that change, which is the usual result of the agency of light on the juices of all leaves exposed to its influence. The green colour of the fluids contained in these cells, marks out their limits, in a transverse section of the leaf, even to the naked eye. The structure of the vessels in succulent dico- tyledonous leaves is the same as in all other leaves. The conducting vessels are spiral tubes, of the same diameter at the apex as at the base of the leaf; and the proper or returning vessels are mem- branous, and apparently perforated, although their transparency renders it difficult to determine their real character. The ramifications are all given off at acute angles ; and appear to be merely separa- tions from the caulinar or petiolar cluster, as Dr. Grew supposed was the case in all leaves ; at least 586 CONSERVATIVE ORGANS. [LECT. X. they do not anastomose until, as I have already stated, they approach the apex of the leaf. In this view of the arrangement and struc- ture of the vascular system of the leaf, I have not noticed those cutaneous vessels which Hed- wig has described as lymphatics; but the exist- ence of which has been denied by Mirbel and others. If we admit their existence, these ves- sels must undoubtedly be regarded as forming a part of the vascular system of the leaf, and ought now to be described; but as this involves a knowledge of the entire cuticular system, the consideration of this point must be deferred until we examine that system. II. Cellular system of the leaf. On cutting a thick, succulent leaf transversely, we immediately perceive that it consists chiefly of a pulp ; which, when placed under the microscope, or examined by a good magnifying glass, is evidently com- posed of cellular tissue. Extending our inquiries, we find that this substance forms a large part of the structure of leaves; filling up the meshes of the network formed by the vessels in the thin and very vascular leaves ; and, in all, occupying that space which separates the two cuticular layers, which constitute the upper and the under disks of the leaf. The cellular substance of leaves differs very LECT. X.] ANATOMY OF LEAVES. 587 considerably in density ; but this diversity depends more on the quantity and quality of the juices the cells contain, than on any diversity of structure in the cells. To the same causes, also, may be attributed, in a great degree, the variety of figure which these cells exhibit: for, although they are in some instances globular, or nearly so; and in others triangular ; or more or less regularly hex- agonal ; yet, it is probable, that the majority are originally spheroidal vesicles; and that the va- riations frqm this figure depend on the turgescence of the vesicles, and the consequent compression which must necessarily result from their conti- guity. The hexagonal figure being that which spheroidal vesicles, mutually comprising one an- other, are naturally disposed to assume, we find that a more or less regular hexagon is the most common form of these cells; and this figure is generally more regular in the cells forming the centre of the substance of the leaf, owing to these being there more distended with fluid, than in those towards either of the cuticles. But that the diversity of figure in the cells of leaves does not, altogether, depend on me- chanical compression, is evident from the fact, that those towards the upper disk of the leaf often differ in form from those towards the un- der disk; and yet in both these situations we may suppose the compression to be nearly equal. 588 CONSERVATIVE ORGANS. [LECT. X. This difference, as shall afterwards be explained, is probably necessary for the distinct functions of these two surfaces ; and thence, in every attempt to theorize on the structure of parts, the pro- priety of keeping in view the fact, that plants as well as animals are living beings, and consequently not regulated by those laws which control the configuration of inanimate matter. No plant is better adapted than the Christmas Rose, Helle- borus niger, to demonstrate this diversity of cel- lular structure in the same leaf. If we examine a thin transverse slice of a leaf of this species of Hellebore, with a good magnifying glass, we perceive that, immediately under the cells of the cutis (31. a.), which are large and oval, there is a range of tubular cells (b.) terminating in the true pulp or parenchyma of the leaf (c.) ; which con- sists of irregular hexagonal cells, and occupies the whole of the space between the tubular cells and the cuticle (d.) of the under disk of the leaf. Under the highest power of the microscope, we find that these tubular cells (32. b.) have apparently a di- rect communication with the cells (a.) of the cutis ; and also with the range of cells (c.) im- mediately beneath them: but in neither are the mouths open, for, a membrane bounds the oblong cell in every direction. In the trans- verse section of some leaves, as for instance those of Calla j&thiopica (33.), which is an aquatic LKCT. X.] ANATOMY OF LEAVES. 589 plant, we find several ranges of tubular cells (b.) commencing under the cutis (a.) of the su- perior disk. Some of these terminate in the hex- agonal cells (d.), and others in empty cavities (f*f.) y such as are present in the leaves and the petioles of all aquatic plants, and which, being filled with air, seem intended for enabling them to rise above or to float on the surface of the water. Under the microscope these tubular cells (34. b.) in the leaves of Calla, resemble a chain of short vessels, with valvular partitions; and were, in- deed, erroneously regarded as such by Malpighi and Leuwenhoek, who observed them in the stem. But these cells are not furnished with open mouths, nor with valves, neither where they originate in the cuticular cells (34. a.), nor at their union with each other, nor where they termi- nate (d.) in the common pulp of the leaf; and this is the case, also, in all leaves furnished with tubular cells. When the tubular cells are cut transversely, they appear to be of an hexagonal 590 CONSERVATIVE ORGANS. [LECT. X. figure (35.), and not round 3 as might be sus- pected from their longitudinal aspect. Sprengel and some others have stated that the cells in all , leaves are elongated near the upper surface ; but the simple inspection of many leaves is sufficient to refute this statement. Among the succulent leaves, even, in which the tubular cells are more frequently met with than in membranaceous leaves, they are not always present ; as, for in- stance, in the leaf of Hoya carnosa, the only dif- ference between the upper and under disk of which is in the structure of the cutis, which on the su- perior surface (a. fig. 16, Plate 10) is a simple transparent pellicle, while on the inferior (b. ibid.) it is cellular ; and in the cells near the upper disk being filled with a greener and more opaque juice than those near the under disk. With regard to the individual structure of the cells constituting the parenchyma of leaves, we find it is the same as that of the cells in the other parts of the plant. Each cell appears to be a dis- tinct, transparent, membranous vesicle, formed into the figure it displays by the pressure of the contiguous cells, and thence, the partition sepa- rating each cell, must be a double membrane. This is more evident in the microscopic examina- tion of the cellular substance of some leaves than of others; thus, in this minute portion (36. page 2 LECT. X.J ANATOMY OF LEAVES. 591 589), taken from the leaf of Iris Germanica, we perceive that not only the cut edges (c.) of the cells appear double; but that where some of the cells deviate from the hexagonal figure, there are evident interstitial spaces (a. d.) be- tween them, which if the cells were not distinct- vesicles would not occur. These interstices have been noticed by Leuwenhoek, Treviranus, Com- paretti, and M. Kieser; the last of whom sup- poses them to exist at every angle of every cell, and by their conjunction to form canals which surround it ; and to be the only passages by which the fluids are conveyed through the cellular sub- stance*. It is unnecessary to point out the im- probability of this opinion ; and it is sufficient foi our purpose to demonstrate the existence of these interstices, to prove the double nature of the in- tercellular partitions. This is very evident in the elongated cells, which we have seen exist near the upper disk of many leaves ; and in the spaces formed by the conjunction of these only, have I been able to perceive any resemblance to the in- tercellular canals of M. Kieser. A question arises in consequence of the sup- position that each vesicle is a distinct sac: in what manner do the cells communicate with each other, and with the vessels which they surround ? * Mem. sur I* Organisation des Plantes, p. 20. 592 CONSERVATIVE ORGANS. [LECT. x. Malpighi, aware of the necessity of an explanation of this point, maintains that a small tubular pro- duction (a. a. 37.) issues from each cell or vesicle, by which it communicates with the contiguous cells (b. b.), and with the vascular system of the leaf (c. c.); and has given a microscopic repre- sentation of this cellular structure in the Cactus (37.) to illustrate his position. A similar idea was entertained, also, by M. de Saussure, who describes the cellular part of the leaf as a con- geries of minute transparent vessels, which are so dilated between their junctions as to assume the appearance of cells or vesicles. But, notwith- standing the high authority advancing this opi- nion, my observations prevent me from according with it: since in no leaf, which I have examined, have I been able to detect these communications. In the Cactus, if we select a small portion of a vessel with some cellular matter adhering to it, as in the minute morsel (38.), under the microscope, before us, we perceive that the cells (b. b. b.) are LKCT. X.] ANATOMY OF LEAVES. 593 in close contact with the vessel (a. .), but neither spring out of it, nor appear to have any direct communication with it; and that neither the elon- gated cells (b.b.b.b.b.), nor the spheroidal (d.d. d.), appear to communicate with one another by any tubular production. The transparency of the cellu- lar membrane produces an appearance (c. c. e.e.e ) at the points of contact of the cells, which might be mistaken for small tubes, but which arises from the impression of the contiguous cells upon one another. Even in that peculiar modification of the cellular structure, which is found immediately with- in the cutis of the inferior disk of some leaves; and in which the cells assume the appearance of ana- stomosing tubes (3Q.), none of the tubular connect- ing processes, described by Malpighi, are perceived; nor do these cells appear to communicate directly with the vessels which they surround. As this is the most curious modification of the cellular structure of leaves, I shall demonstrate it to you as it appears in the floral leaf of Helleborus viridis, which I select for this purpose, from its being almost devoid of colour. In the minute portion before us, as it appears under the microscope, we perceive the range of cells e. e. (39) closely ap- plied to the proper vessels d. d. but not communi- . eating with it: the cells c. c. c. c. assume various forms, but all are apparently tubular ; and, where they are more transparent than common, we dis- VOL. i. Q Q 594 CONSERVATIVE ORGANS. [LECT. X. tiuctly perceive the nature of their conjunction (b.b.); but no projecting tubular processes. The spaces, such as a. a. are filled with air in the same manner as the vacuities in the leaves of aquatic plants. In what manner then do the cells commu- nicate? To answer this question properly, we ought to understand the structure of the intercellular membrane. But here our instruments fail, if they do not mislead us ; and, under glass.es of the highest power, this membrane appears different under different circumstances : by transmitted light, it seems a simple, unorganized, transparent pel- licle ; but, by reflected light, is evidently porous. I have already stated my belief that the cells of the stem communicate by pores, and I see no reason for altering this opinion with regard to those of the leaf: although I do not concur in opinion with M. Mirbel and Sprengel, that the form, position, magnitude, and number of these pores can be determined. An opinion has been advanced, that the fluids may be transmitted from cell to cell, " consistently with the integrity of the cellular structure," by the exercise of the alternate functions of secretion and absorp- tion;" but these functions imply the existence of either glands or vessels connected with the absorb- ing and secreting surface, which are, however, even less demonstrable than the pores. Upon the whole, the question is still unanswered; and all LECT. X.] ANATOMY OF LEAVES. 595 that we certainly know of the subject is, that the fluids are transmitted from cell to cell, through every part of the vegetable system, although the structure by which this is accomplished remains undiscovered. Whatever may be the mode in which the cells communicate with one another, their contents are more or less fluid or solid, according to their situ- ation in the thickness of the leaf. Thus, in thin leaves the cells near the inferior disk are more trans- parent, owing to their contents being more fluid than those near the upper disk ; but in both we per- ceive a number of granules, which are more opaque and of a deeper green, as the cells containing them approach the upper disk. In succulent leaves, and those which maintain a vertical position, the opacity and green colour of the granules, are the same towards every face of the leaf; but they are generally colourless in its centre. In the cells, also, of some leaves, regular crystallized salts are found; and in others the fluids are tinged of dif- ferent hues besides green; in which cases the leaves themselves display the same hues on one or both surfaces. The size of the cells varies in different leaves ; in some, even when examined under the most powerful glasses, they appear like the smallest vesicles ; while, in others, they are so large as to be perceptible to the unassisted eye. QQ2 596 CONSERVATIVE ORGANS. [LECT. X. I have already demonstrated to you the exist- ence of large vacuities in the foliar parenchyma of the leaves of aquatic plants. Mirbel regards these as accidental productions, rents or defects in the cellular texture; an opinion, however,, which is instantly refuted on a minute examination of these parts. If we place a portion of the petiole, or any of the larger costse of an aquatic, this thin trans- verse slice, for example, of the petiole of Trapa natans (fig. 12, Plate 10), under the microscope, we perceive that the vacuities have a symme- trical arrangement around the centre (c.), which is vascular and consequently more opaque than the rest of the slice; and that some of the va- cuities are open (a. a.), while others (b. b.) are closed. The membrane which covers some of the vacuities as they appear in the slice (fig. 12.) before us, is a diaphragm, which, as it forms the roof of one cavity, is also the floor of another ; and it is owing to these diaphragms not being all on the same plane that some of the vacuities appear de- void of them, in the transverse slice of the petiole of the leaf of any aquatic plant. The intimate structure of these diaphragms is seen in a highly magnified view of one of them, as represented at fig. 13, Plate 10 ; in which b. shows that the dia- phragm consists of regular hexagonal framework, with the intervening membrane either perforated or studded with small transparent, amylaceous gra- LECT. X.] ANATOMY OF LEAVES. 597 miles; and a. that the lateral partitions between the cavities, apparently consist of square cells, when transversely divided, as in the figure before us, although they are hexagonal, when viewed laterally, as in the diaphragm. Each cavity is lined on every side with a thin pellucid pellicle, closely resembling the external cuticle ; and frequently hairs, knobs, and similar cuticular productions are found projecting into these cavi- ties. Kieser, who is the only author who has no- ticed these bodies, remarks that he had sometimes observed, in the cavities of Calla -^Ethiopica, small globular, pedunculated bodies ; which, springing from the sides, project towards the centre of the cavities ; but, according to my observations, they are more common in those of Typha, Equisetum, and Nymphcea, in the latter of which they closely resemble the branched hairs (see Plate 9, fig. 14.), which form the tufted pubescence on the under disks of some leaves. In closing this view of the anatomy of the cellular system of leaves, I have to remark, that although it embraces the more common varieties that are met with, yet it is probable that, in the vast range of the vegetable kingdom, many other diversities of structure of the cellular matter exist. From these inquiries into the structure of the vascular and cellular systems of leaves, the affinity which exists between the stem and the leaf is very 598 CONSERVATIVE ORGANS. [LECT. X. obvious. In the stems of monocotyledons, the vessels run nearly in straight lines in distinct fas- ciculi, embedded in a cellular pulp; and a simi- lar vascular arrangement presents itself in the leaves of this tribe of plants. In dicotyledons, on the other hand, the vascular fasciculi of the stem are not distinct, but form a reticular tissue which covers the whole circle of the stem ; and., in like manner, in the leaves, the vessels ramify in every direction, forming a most complicated and beau- tiful network, the interstices of which are filled with the cellular pulp. The leaf, therefore, may be regarded., in some respects, as a mere expansion of the stem; and, consequently, in aphyllous plants, we perceive that the stem is adapted to perform all the functions of the leaf. The internal structure of the floral leaves or bractece, and of those more temporary foliar appendages, which are termed stipulce, is nearly the same as that of the real leaf; even the scales that envelop buds, and which are always described as deriving their origin from the cortical part only of the stem, and consisting chiefly of cellular matter, have in every respect the same structure as leaves, as far, at least, as relates to their vascular and cellular systems. LECT. XI.] ANATOMY OF LKAVES. 599 LECTURE XL OF THE CUTICULAR SYSTEM OF LEAVES! USE OF THE CUTICULAR APERTURES. OF THE APPEND- AGES OF THE STEM AND LEAVES PUBESCENCE THORNS PRICKLES GLANDS PROPS: USES OF THESE APPENDAGES. EVERY leaf is covered with a real skin or epi- dermis, which not only guards the vascular and the cellular matter from external injury; but is the medium by which it performs the important functions of absorption and exhalation. In the majority of leaves, the epidermis can be separated from the parts it covers: and appears to be a compound organ, or to consist of two distinct layers; the exterior of which is a fine, transparent, apparently unorganized pellicle, and the interior vascular and cellular. But the opinions and de- scriptions of phytologists are at variance on this subject. Grew*, Malpighi "f~, Du Hamel J, Des- fontaines, Mr. Keith ||, M. Kieser f , M. Mirbel **, * Anal, of Plants, p. 62. t Anat. Plantarum, p. 212. J Phys.des Arbres, i. 8. Mem. de Tlnst. Nat. i.4-81. || Syst. of Phys. Bot. i. p. 313. 5f Mem. sur F Organ, des Plantes t p. 141. ** Elcm. de Phys. veg. i. 36.' 600 CONSERVATIVE ORGANS. [LECT. XI. and some others, have described the cuticular co- vering of leaves as a simple body ; while M. de Saussure*, Mr. Francis Bauer -}-, and M. De- eandolle;}:, concur in the opinion which I main- tain of its compound nature. Let us examine the proofs upon which this is founded. The true epidermis, or the delicate pellicle which forms the outermost covering of the leaf, can be readily demonstrated in any small portion of the cuticular covering carefully raised by the point of a lancet, and placed in a drop of water under a powerful microscope. In this small portion, taken from a leaf of Dianthus Caryophyllus,\t isseen (a.40. page 602) extending beyond the area of the meshes (b. b.) of the interior cuticular layer, which are seen through it, and is evidently a simple pellicle. But Mr. Keith , who admits that it may be seen in this manner, supposes that its individuality is not proved by such a demonstration, as the meshes, the intervals of which it might originally have filled up, may be accidentally obliterated ; but, although there is some plausibility in this objection, yet, when we take this appearance in conjunction with the double character observed in the trans- verse section of the cuticular covering of every * Encyclop. Method, i. 67. f Tracts relative to Botany. I Mem. de I'Inst. Nat. i. 351. Syst. of Physiol Bot. vol. i. p. 313. LECT. XI.] ANATOMY OF LEAyES. 601 leaf which we have examined, there is sufficient reason for believing that it is as much a distinct layer as the cuticle in the human body, although, in the leaf, it cannot be readily detached from the interior layer of cutis. It is described by Saussure as being perforated by the slits or pores which, I shall soon demonstrate to you, are found on one or both of the surfaces of every leaf; but we shall find, on minutely examining these, that it is not perforated by them, but enters into them, as well as into every gland opening on the surface of a leaf, as a lining membrane ; and is, in fact, the covering of every part of the vegetable texture, which would otherwise come in contact with the air. If, however, it cover every part of the sur- face of the leaf, and is an imperforated mem- brane, by what means, it may be asked, does the fluid which exhales so freely from the leaves es- cape? It is certainly not easy to answer this ques- tion ; but as we can scarcely form an idea of a membrane perfectly free from pores, even in a living body, transmitting fluids ; we may conclude that, although no pores are visible in this mem- brane, even when it is examined under the mi- croscope, yet, it does not follow that no pores exist ; and, in accounting for the transudation of the fluids, which the leaf throws off, we must always bear in mind, that the functions of living bodies are influenced by different powers from 602 CONSERVATIVE ORGANS. [LECT. XI. those which regulate the operations connected with inert matter. The second or interior cuticular layer is seen through the epidermis, and consists of a vascular network resting upon a layer or layers * of cells. Taking the same portion of the cutis of Dian- thus Caryophyllus (40.) to demonstrate the super- ficial structure of this layer; we find that, except within the boundary of the detached epidermis (a. a.), the whole is spread with a network of ir- regular hexagons formed by lines which appear double, and terminate in a ring surrounding a slit or oblong pore (c. ,*.), which occupies the centre of one of the longest bounding lines of almost every alternate hexagon. The same appearance of the interior layer of cutis is seen on both surfaces of the leaf of Dianthus ; but, as I shall afterwards demonstrate, this is not the case in the majority of leaves-}-. If we now place a very thin transverse * Mr. F. Bauer describes the cuticle of a species of Hae- manthus as composed of several layers of cells. See Tracts relative to Botany. Lond. 1805. I In Plate 10, figure 3 represents a portion of cuticle LECT. XI.] ANATOMY OF LEAVES. 603 slice of the leaf of Dianthus under the micro- scope, to examine this layer in its thickness, we can readily distinguish it (b. 4 1 .), by the form of its cells (c.) from the parenchyma (d. d.) upon which it rests; and, also, from the epidermis (a.) which covers it; for, in this leaf, the cells of the parenchyma assume an oblong form im- mediately under the cutis, whereas the cells of the cutis are irregular spheroids, and the simple layer of epidermis is remarkably distinct. Whether these cuticular cells have any direct communication with the oblong cells beneath them, I have not been able to determine; but in separating the cutis, by tearing, some of the ob- long cells always adhere to it; and, when viewed through it, by transmitted light, appear like a smaller cuticular network (d. d. 40.), filling up the larger meshes. These might be mistaken for the cuticular cells; but the transparency of the cutis prevents its cells (e. 4 1 .) from being visible in a superficial view of the organ. In this trans- verse section of the leaf of Dianthus (41.) the dis- tinct nature of the epidermis (a.) is perfectly evi- dent, both as a covering to the true cutis (b.) and as lining the slits (e.), one of which is here di- vided lengthways. taken from the under disk of the leaf of Hoya carnosa; and figure 4-, a portion from the upper disk of the same leaf. Both are magnified about 300 times. 604 CONSERVATIVE ORGANS. [LECT. XI. The lines forming the meshes which thus cha- racterize the cutis of leaves, were first described by Hedwig as vessels, originating in the circum- ference of the pores; an opinion which is sup- ported by the elder M. De Saussure and M. Kieser ; and which is confirmed by the microsco- pical examination of a portion of the cutis of any leaf. Remarking the facility with which this part of the cuticular structure can be demonstrated, our surprise is excited that Sprengel, Link, Mirbel, Jurine, Krocker, and others, should have ad- vanced the opinion, that these reticulations form no part of the real structure of the cutis, but are merely the adherent fragments of the sides of the subjacent cells : for, as I have already demon- strated, the parenchymal cells are much smaller than the cuticular meshes ; and, when the cutis is sufficiently transparent, they are seen through it very distinctly (4O. b. .), but not at all coin- ciding with the sides of these meshes. Admit- ting, therefore, that these lines are lymphatic vessels, it is not improbable, as M. Kieser has asserted, that they terminate by one extremity in the larger vascular fasciculi ; but on this point I have not been able to satisfy myself. The meshes, which they form, differ very much, both in form and size, in different leaves*. In al- * Mirbel, reasoning from the false opinion which he had formed of the structure of the cutis, observes, " les differences 2 LECT. XI.] ANATOMY OF LEAVES. 605 most all the monocotyledons, in the Grasses, and in every plant the leaves of which have parallel costae, the meshes are nearly irregular parallelo- grams ; but, in forming these, the vessels some- times run in straight lines, as in common Meadow grass, Poa trivialis (45.) ; sometimes in slightly un- dulated lines, as in the White Lily, Liliurn candi- dum (44.); and sometimes zig-zag, as in Indian Corn, Zea Mays (fig. 14, Plate 10). In some of the fleshy leaves they are nearly regular hexagons, as on the upper disk of Hoya carnosa (fig. 4, Plate 10), and on both surfaces of the leaves of Aloe verrucosa (43.) : but, in the majority of di- cotyledons, they assume very irregular figures -j~. WK " qu'elle presente viennent de la forme des cellules dont elle ** faisait partie." " Les parois cellulaires restant attachees a " l'piderme, y dessinent de petits compartimens dont la forme " indique celle du tissu cellulaire lui-meme. Tantot ce sont des " parallelogrammes plus ou moins reguliers, tantot des hex- " agones, tantot des polygones divers, dont les cotes sont on- dules.' 1 Elem. de Phys. veg. 1 partie, p. 36. f This irregularity is well exemplified in common Sorrel, Rumex acetosa. In Plate 10, fig. 9 represents a minute portion 606 CONSERVATIVE ORGANS. [LECT. XI. Whatever may be the figures which they pre- sent in the cutis covering the spaces between the vascular ramifications of the leaf, they in- variably appear as irregular parallelograms ( an d in the Agave tribe (59- p. 6li), and a few other families of plants, they are quadrilateral. In almost all leaves they are surrounded by a border, in which the vessels forming the cuticular meshes appear to terminate. Placing minute portions of the cu- ticle of different leaves under the microscope, we can readily ascertain the superficial form of these pores : the following are the principal diver- sities of form which I have observed *. 1. A simple slit, more open in the middle than at either end, bisecting an oval shield ; which may, there- fore, be termed the oval scutiform aperture, oscu- lum scutiforme ovatum, as exemplified on the lower disk of the leaves of Sage (48. p. 610), of Lactuca quercina (4Q.), Dandelion, Leontodon Taraxacum, Sweet-scented Coltsfoot, Tussilagofragrans, many of the Grasses, the common Bean, Vicia Faba, &c. 2. A simple slit, bisecting an oval shield enclosed * In naming the different kinds of pores, I have assumed the fact, that they are respiratory organs, or apertures resembling in their functions the spiracula of insects ; but, on account of the simplicity of their structure, I have preferred the term osculum to that of spiraculum. VOL. 1. R R 610 CONSERVATIVE ORGANS. [LECT. XI. within a ring, which I name the annulated scu- tiform aperture, osculum scutiforme annulare, as seen on the inferior disk of the leaves of Helle- borus fcetidus (51.), the Violet tribe (52.), the Coffee plant, Coffea Arabica (53.), the White Lily, Lilium candidum (44. p. 605), Gardenia la- tifblia (46. p. 606) ; the common Cabbage, Bras- sica oleracea ; the outside of the tubular leaf of Saracenia (54.), &c. 3, An open slit, or an oblong- pore, enclosed by a simple oval ring, which may thence be termed the annulated aperture, osculum annulare; as on the inferior disk of the leaf of Hibbertia scandens (50.), Ivy, Hedera Helix (55.), Jacquinia ruscifolia (56.), Dianthus Caryophyllus (40. p. 602), Laurus Canarlensis, Wolfsbane, Aconitum neomontanum, Privet, Ligustrum vul- gare, &c. 4. A circular pore in the centre of a circular shield, which we may name the circular aperture, osculum circular e ; as on the inferior LKCT. XI.] ANATOMY OF LEAVES. 611 disk of the leaf of common Sorrel, Rumex acetosa (a. b. fig. 10. Plate 10), of the Primrose, Primula (57.), and, very beautiful, on both surfaces of Cactus opuntia (38.). 5. A quadrilateral pore, osculum quadrilaterale, surrounded by an ele- vated margin, as on both surfaces of the leaves of Agave Americana (59-), and of all the other species of the succulent tribe to which it belongs. I cannot avoid remarking, in this place, the na- tural separation, which may be traced by the form of these apertures, between the Aloes and the Agaves. In the former the pores are always circular (60.), and in the latter they are inva- riably quadrilateral (59.) 61 It is impossible to notice all the modifications of these different species of the cuticular aper- ture; I shall, therefore, only remark, that among the varieties of the annulated aperture, we some- times find the space between the pore, or the shield and the enclosing ring, divided into distinct por- R R 2 612 CONSERVATIVE ORGANS. [LECT. XI. tions ; and occasionally a double ring, with the in- tervening space, also, divided into four or more equal parts: examples of the first variety are found on the lower disk of the leaves of Lilac (61. p. 611), of Acuba Japonica (6^.), Cussonia thyrsi- flora (63.), Hoya carnosa (fig. 3, Plate 10) ; and on the upper disk of the leaf of Rumex ace- tosa (fig. 9, Plate 10). The upper disk of the leaves of Viola odorata affords an excellent illus- tration of the double ring (64.). But the most re- markable form of the cuticular pore, which I have yet observed, is found on the back of the leaves of common Oleander, Nerium Oleander. It ap- pears, on a superficial view, a simple oval aper- ture without any shield, but guarded by hairs which cross it in different directions (65. a. .); and is comparatively much larger than any of the other kinds of pores. Some writers have ventured to assert, that no apertures are found in those plants which are generally regarded as composed almost entirely of cellular matter, among which Marchantia is usually reckoned ; but the incorrectness of this opinion can be easily proved, by placing a small LECT. XI.] ANATOMY OF LEAVES. 613 slice of the cutis of Marchantia under the microscope. The aper- tures are oval, and placed in the centre of a slight elevation, as re- presented in the marginal cut (*). In respect of size, pores differ considerably in different plants ; but on the leaves of the same plant their size is nearly uniform. The largest, as far as rny obser- vations extend, are those found on the leaves of Oleander; and the smallest on those of the genus Myrteae. Sprengel says, that in the Coronariae " their longitudinal diameter is from -rV to ^ " part of a geometrical line, and their diameter, " in the cross direction, is from -^V to ^V part;" but in " the Myrteae, Rosaceae, Leguminosae, " and Caryophylleae, two hundred of them, at " least, might lie upon a geometrical line-f~." In number the foliar apertures vary, also, in dif- ferent plants. The more minute they are, the more numerous. On the lower surface of the leaf of Gardenia latifolia, we find an aperture in almost every mesh ; but in the Aloe tribe scarcely one pore for twenty meshes, and on the leaf of Oleander, one among sixty. With regard to position, these aper- tures are in some instances arranged in lines from the base to the apex of the leaf, and have the same t Elements of the Phil, of Plants, fyc. by A. P. DecandoUe and K. Sprengel, 310. RR3 614 CONSERVATIVE ORGANS. [LECT. XI. direction throughout; but in the majority of leaves they have no regular arrangement, and as- sume different directions. In herbaceous plants we generally find them on both surfaces of the leaves; but in ligneous plants they are scarcely ever seen on the upper surface. They are never situated on the costae, nor on the edges of the leaf. But these demonstrations make us acquainted with the superficial aspect only of the foliar aper- tures, beyond which it is surprising that no Phy- tologist has yet attempted to push his investiga- tions; although it is by no means difficult to determine their structure by the aid of the micro- scope. Placing a very thin vertical slice of a leaf of the Clove Pink, Dianthus Caryophyllus, cut in the direction of the axis of the leaf, under the microscope, we find that the aperture (e. 41. p. 602) which is thus divided in its longitudinal diameter, is a short cylindrical tube penetrating completely through the cutis, and terminating in a cul de sac, which is impressed into a vesicle (f.) that appears to communicate with the oblong cells (d.) immediately beneath the cutis. But al- though the aperture penetrates the cutis, there is no opening through the epidermis (.), which, on the contrary, enters into the tubular part of the pore and lines it throughout. In another slice of the same leaf, cut so as to divide one of the aper- tures in its cross diameter (42. p. 602), we per- LECT. XI.] ANATOMY OF LEAVES. 615 ceive that the vesicle (/*.) appears to be double ; from which it is probable that it is this vesicle, seen through the transparent substance of the cutis, which gives the appearance of the shield in the superficial view of the aperture. As we find that, in the superficial view of these apertures, the character varies considerably in different plants, so this form of the tube and the vesicle is also variously modified ; but the general charac- ter is nearly the same, with a very few exceptions, throughout the vegetable kingdom, as far, at least, as my observations extend. Thus in the Agave (59- p. 611), the superficial form of the aperture is very different from that in Dianthus (40. p. 602) ; but if we examine it in a transverse section of the leaf, although we find the general structure of both agreeing, inasmuch as the aper- tures of both are lined with the epidermis, pene- trate the cutis, and have at their bottom the vesi- cular ring ; yet we perceive that that of the Agave differs from that of Dianthus, in terminating in a large dilated cell (66. f.), which is found always R R 4 616 CONSERVATIVE ORGANS. [LECT. XI. filled with air, although it is closely surrounded by the oblong cells (e.) of the parenchyma, turgid with green juice. In the oblong section, also, of this aperture (67.) the vesicular ring (f.) does not appear to be simple as in Dianthus, but is divided by a duct, in which the aperture (a.) seems to terminate, and which apparently opens into the air cell (f. 66.) ; which is cut away in this section (6?.)- In the leaf of Oleander the aperture (d. 68. p. 615) expands into a kind of sac where it pene- trates into the substance of the parenchyma (c.) ; and it is throughout lined with the same kind of hairs which guard its orifice ; but I have not been able to determine whether its lining membrane, which is a production of the epidermis (a.), be porous ; although I have examined it by glasses of the highest powers. I may here remark that the section of this leaf displays an example of a cutis consisting of four layers of cells (b.). Decandolle considers that the cuticular apertures are con- nected with the ultimate ramifications of the ves- sels of the leaf* ; and, if it be true, that the cu- ticular meshes are formed by lymphatic vessels, which terminate on one hand in the larger vessels of the leaf, and on the other, in the vesicular circles surrounding the fundus of the aperture, this opinion must be correct. * Journ. de Phys. iii. p. 130. LECT. XI.] ANATOMY OF LEAVES. 617 The information we have thus obtained of the structure of these pores, -induces me to believe that they are the respiratory organs of plants. But Phytologists have considered them intended for the functions of absorption and exhalation*; and even Sprengel, who hints something regarding their analogy to the breathing spiracula of insects, rejects the idea of their being respiratory organs, because "they are not in immediate contact with the spiral vessels-f~." If, however, it be admitted, as I have endeavoured to prove, that the spiral vessels are not air-tubes, but sap-vessels, this ob- jection must fall to the ground. The idea that they are absorbing organs, is supposed to gain support from the circumstance, first ascertained by Bon- net J, that leaves absorb more powerfully with their lower than with their upper surface; and Sprengel remarks that " the slits are more nume- " rous in juicy plants, which are nourished more " by the surface of the leaves than by the roots**." But 1 reply, that although leaves absorb chiefly iby their inferior surface, yet, this does not prove that these apertures are the absorbing organs ; for we find none of them on the lower side of the leaves of Nymphsea and other aquatics, which * Among those holding this opinion are MM. Bonnet, Decandolle, Sprengel, Mirbel, Mr. Ellis, and Mr. Keith. f Elements of the Phil, of Plants, j 31 1 . ** Ibid. I. c. t Recherches sur I' Usage des Feuilles, p. 20. 618 CONSERVATIVE ORGANS. [LECT. XI. have floating leaves ; although these leaves ab- sorb powerfully by their lower surface, and exhale by their upper, which is covered with these aper- tures. Decandolle and Sprengel's remarks regard- ing succulent and fleshy leaves, are also incor- rect ; for, in the genus Aloe, which is supported chiefly by absorption, the apertures are compara- tively few ; and the function of absorption in these plants seems to be connected with a small papilla, which rises in the centre of every mesh ; and pro- bably acts in the same manner as radicles on roots. It is still more difficult to accord with the opinion that the same foliar apertures perform such oppo- site functions as those of absorption and exhalation ; although there is nothing incongruous in supposing that they are both exhalant and respiratory organs. That they exhale, was first rendered probable by the experiments of Treviranus, who found that plates of glass applied to the lower disks of leaves were soon covered with drops of water, while they were not at all bedewed when they were affixed to the upper disks ; and Decandolle afterwards proved that the aqueous transpiration is greatest in those plants which are supplied with the greatest num- ber of apertures. The opinion, however, still prevailed, that leaves exhale also by the upper disk, although the majority of leaves have no apertures on that disk; but before yielding im- plicit credence to the assertion, that these aper- LECT. XI.] ANATOMY OF LEAVES. 619 tures are the foliar exhalants, it was necessary to prove that no exhalation takes place when they are obstructed; and to determine this point I made the following experiment: Two twigs of Laurustine, each having four leaves nearly of the same size, were cut from the plant, and brought to the same weight by being placed in opposite scales. The lower disks of all the leaves on one twig were next brushed over with a composition of mucilage of gum arable and a small proportion of Traga- canth; and when this was dry, each twig was placed under a cylindrical glass jar containing air, and immersed in a saucer of water. In a short time the sides of the jar containing the twig in its natural state, were covered with drops of water ; but, at the end of two days, not the smallest quantity of mois- ture appeared on the sides of the jar containing the twig, the apertures of the leaves of which had been obstructed by the mucilage. The conclusion to be drawn from this experiment is, that that surface only on which apertures exist exhales, and conse- quently that these apertures are the exhaling or- gans. This experiment leads us, also, to draw a conclusion from the experiments of M. Bonnet, dif- ferent from that drawn from them by that author. He concluded that, because the leaves of trees which were laid with their lower disk upon water remained longer green and fresh than those which had the upper disk applied to the water, this fluid 620 CONSERVATIVE ORGANS. [LECT. XI, was absorbed by the under surface; but if, by thus placing such leaves, we obstruct the exhaling pores, we can readily see why the leaves will be longer withering, since no exhalation takes place ; while in those placed on the opposite disk, the ex- halants are free to perform their functions, and thus empty the foliar cells of their aqueous con- tents more quickly than the absorbents of the upper disk can supply them. In stating this ar- gument, I do not feel bound to point out by what media leaves absorb : although I have already hinted my opinion, that this function in the suc- culent plants of arid soils is performed by a dis- tinct set of organs. A further proof of the truth of this conclusion is, that the leaves of herbs, as Bonnet himself observed, remain fresh nearly the same length of time when placed on either sur- face; for these apertures are present on both surfaces; and, besides, these leaves sooner wither, whichever surface is in contact with the water, than the leaves of trees which are laid with their inferior surface upon that fluid. Were I now to enter fully into the proofs, that the foliar apertures are, also, respiratory organs, I should be anticipating the arguments which must be again detailed, when we come to the consider- ation of the general functions of the leaves ; and, therefore, I shall at this time, as briefly as possible, examine those proofs only which are connected with the structure of these organs. LKCT. XI.] ANATOMY OF LEAVES. 621 All animals that require the presence of air for their existence, have some peculiar apparatus for producing that change in the blood which has been termed its oxygenizement ; and the change, so termed, is said to be the result of respiration, whether it be performed by lungs or by spira- cula. Plants, also, require the presence of air; vitiating it, under certain circumstances, in the same manner as animals, but, under others, in- creasing the proportion of its oxygen : hence plants may properly be said to respire, and the question arises, by what organs is this function performed? Phytologists have generally agreed, that the leaves are the lungs of plants ; but still it may be inquired, does the whole of the leaf act, or in what part of it are the respiratory organs si^ tuated ? My answer is, that the foliar apertures are the actual breathing organs of the plant. In support of this position I refer to the situation of these apertures, which are never seen on leaves that are not exposed to the air ; for the leaves of sub- mersed aquatics are devoid of them; even the leaves of plants which are not naturally aquatics, if they be submersed, soon lose them ; and al- though some plants of the higher classes, which grow in the air, have no leaves, yet, these have apertures on the stem, which, in such instances, perform the respiratory function. But the most perfect plants are furnished with leaves which, 622 CONSERVATIVE ORGANS. [LECT. XI. from being membranous and from the nature of their attachments, are moveable in the air, and thence have constantly a fresh atmosphere of that fluid applied to their breathing apertures; this mobility of the leaf supplying, in some degree, the motion of the thorax and the diaphragm in the more perfect animals. The plants which have very thick and immoveable leaves, on the contrary, or which are devoid, of leaves, as they resemble the cold-blooded and slow-moving animals in their tenacity of life, like them, also, require a smaller supply of air, and consequently, as we have al- ready seen, are less amply supplied with breathing apertures. I have not been able to ascertain whe- ther the apertures themselves have the power of opening and shutting; but from the appearance of the orifices of these organs, as they are seen on the leaf of Indian Corn, when very highly magnified (see Plate 10, fig. 15), it is not improbable that some degree of dilatation and" contraction takes place, although we cannot determine the fact. In structure these organs seem well adapted for the purposes of vegetable respiration, when we con- sider that the changes effected by this function in the sap of vegetables in the leaf are not required to be so quickly produced as those in the blood of animals ; even of insects of the lowest description. The air is admitted through the funnel-shaped pore, whiqh perforates the cutis, into a vesicle si- LECT. XI.] ANATOMY OF LEAVES. 623 tuated under it ; and which probably communi- cates with the cuticular cells, as these are in ge- neral found filled with air. The aqueous contents of the cells that form the parenchyma of the leaf, are thus brought into immediate contact with the atmosphere. It is not easy to assign a rea- son why these apertures are found on the under disk only of the leaves of trees, while they ap- pear on both disks of herbaceous leaves; there being lymphatics on both disks of the former as well as of the latter description of leaves. If any connexion could be traced between the re- turning vessels and the apertures, the difficulty would be diminished, the situation of these vessels being on the lower disk of the leaves of trees. With regard to the origin of these apertures, M. De Saussure's and M. Kieser's observations would lead us to believe, that they are merely the terminations of numerous vascular processes from the larger fasciculi, which, gradually pene- trating the cuticle, are thus enabled to discharge their fluids. This opinion, however, is altogether hypothetical. They are so far essential that they are found on every leaf in contact with the at- mosphere; their structural characters, position, and situation, are the same on the leaves of every plant of the same species ; and their existence seems to be influenced by no conditional circum- stance except the presence of air; for I have al- 624 CONSERVATIVE ORGANS, [LECT. XI. ready demonstrated the incorrectness of the as- sertion, that etiolated leaves are devoid of aper- tures, or at least the remark is not universally applicable. With regard to the fact, that they are not found on submersed leaves, even of land plants which are made to grow in the water, I may merely observe, that the leaves produced on such plants differ from those which are natural to them, not in the absence of apertures only, but in form, structure, and functions. The knowledge of the structure of leaves en- ables us to form a correct idea of the importance of these organs in the economy of the plants. We find the vessels which convey the sap from the roots terminating in the leaf, and spreading out their contents through its cells, to undergo certain chemical changes which are essentially influenced by the action of the air and light : we find, also, a new system of vessels commencing here, which take up again the sap thus converted into proper juice and conduct it downwards, depositing in their course the various secretions formed from it, either in the stem or in the roots, as the nature of the plant requires; and, in aid of these operations, a cuticular system admirably adapted by its trans- parency to transmit the rays of light into the foliar cells, and by its organic apertures to admit the air, and at the same time favour the exhalation of the superabundant water, which the ascending 2 LECT. XI.] ANATOMY OF LEAVES. 625 sap necessarily contains. But, besides fitting the sap for yielding the secretions found in the bark, wood, and roots of plants, the leaf itself is a secerning organ, and contains in its cells and fol- licles many secretions useful, undoubtedly, to the plant itself; but, independent of that, of the first importance in medicine and the arts; and in supplying food for the support of animal life. Thus the leaves of Henbane, Hyosciamus niger; Deadly Nightshade, Atropa Belladonna; Wolfsbane, Aco- riitum napellus and neomontanum; Hemlock. Co- nium maculatum ; Fox-glove, Digitalis purpurea ; the whole genus Tobacco, Nicotiana ; Wild and Garden Lettuce, Lactuca virosa and saiiva; the genus Thorn Apple, Datura ; Yellow-flowered Rhododendron, R. Chrysanthum; the Poison Oak, Rhus Toxicodendron, and many other plants, con- tain alkaline principles, which produce very powerful sedative and narcotic effects on the ani- mal economy ; and the Prussic acid, a still more direct sedative, is present as a secretion in the leaves of the Laurel Cherry, Prunus Lauro-Cera- sus. Bitter and tonic principles are found in the leaves of the genus Wormwood, Artemisia; Centaury, Chironia Centaurium ; Horehound, Marrubium vulgare; Marsh Trefoil, Menyanthus trifoliata; Garden Angelica, A. Archangelica ; Hyssop, Hyssopus officinalis, &c. : astringent in those of the Oak genus, Quercus; Bears Whortle VOL. i. s s 626 CONSERVATIVE ORGANS. [LECT. XI. Berry, Arbutus uvce Ursi; almost the whole of the genus Sumach, Rhus; the Tea, Thea; and many other genera: and emetic in Asarabacca, Asarum Europceum, &c. The purgative proper- ties of the leaves of Senna, Cassia senna, and of Hedge Hyssop, Gratiola officinalis ; and the diaphoretic of Sage, Salvia officinalis, are well known. The secretions of some leaves are so acrid as to inflame and blister the skin when applied to it; as those of many of the species of the genus Ranunculus*; and of Savine, Juniperus Sabina: on the other hand, some leaves, as those of Marsh Mallow, Althea officinalis, and of Com- mon Mallow, Malva sylvesiris, afford bland mu- cilages; others, as those of Sorrel, Rumex acetosa, and of Wood-Sorrel, Oxalis acetosella, contain cooling acids: and some again, as those of the Mints, Menthce; Balm, Melissa officinalis; Rue, Ruta graveolens; the Cajuputi tree, Melaleuca Leucadendron, &c. secrete essential oils, which rouse and stimulate the animal system when taken into the stomach, or even when applied to the skin. As food, men employ a great variety of leaves, which yield a bland fecula and saccharine * The leaves of R.jlammula are used for raising a blister in the Hebrides. They are chopped and rubbed between hot stones immediately before being applied ; and generally raise a blister in an hour and a half. The leaves of R. sceleratus have the same effect ; but often occasion an irritable sore, which cannot readily be healed. LECT. XI.] ANATOMY OF LEAVES. 627 matter, or in which the acrid secretions can be easily destroyed by cooking; and the number of those which might be used for this purpose, but are still neglected, is very considerable*. No distinct secerning organs have yet been dis- covered in leaves, except as relates to some sac- charine and resinous exudations and to the essen- tial oils; and the organs producing these have been termed glands. The exudations give various cha- racters to the surfaces of leaves; but the oils are preserved in distinct follicles; which, in many in- stances, open by excretory pores, that are rea- dily distinguished from the common foliar aper- tures ; and through these they are exhaled, pro- ducing the odours for which many leaves are distinguished. I should now endeavour to de- monstrate the forms and structure of these glan- dular organs; but as they are found sometimes on the stem as well as on the leaf, I shall not con- fine myself to the examination of them as con- nected with the leaf only; but as part of the ge- neral vegetable appendages. The term appendage is applied to certain or- * The leaves of the common Dock, Rumex patientia, were eaten by the ancients under the name of Lapathum; and are still eaten in some places of Germany under the name of Eng- lish Spinage. The leaves of R. scutatus and of many species of the genus Atriplex, were also formerly used as pot-herbs, and, indeed, afford very palatable and nutritious food. ss2 628 CONSERVATIVE ORGANS. [LECT. XI. gans which are occasionally, but not invariably, found connected with the universal vegetable organs. They are never all present on the same species of plant ; but which of them soever is found on any individual is general to the species, and invariably present. All of them are import- ant to the plants on which they are found ; and a knowledge of them, besides throwing great light on vegetable physiology, is of utility to the practical Botanist, in affording characters for specific distinctions. The caullnar and foliar ap- pendages, or those connected with the stem, branches, and leaves, to which we have now to direct our attention, may be classed under the six following heads : Glands, Pubescence, Spines, Prickles, Props, Foliaceous appendages, and ano- malies *. 1. GLANDS, Glandulce. Linnaeus has defined the vegetable gland to be " a little tumour excreting " an humour -f^;" but this definition comprehends those glands only which are external and elevated above the cutis ; and, as we find many minute or- gans of a structure distinct from the common tex- ture of the part on which they are situated, and se- * I have adopted this term from Mr. Keith, because I can suggest no better for the classification of the objects intended to be described under it. f " Glandula est papilla humorem excernens." Phil. Bol, *.*, LECT. XI.] CAULINAR AND FOLIAR APPENDAGES. 629 parating a peculiar fluid, embedded in the cellular substance, or half sunk in the cutis, and, if elevated above its surface, displaying great diversities of form, its exceptionable character is evident. In rejecting, however, this definition, the difficulty of forming an appropriate one must be acknow- ledged. A definition formed altogether on the existence of the secretory function, would occasion us erroneously to regard as glandular any part where the presence of a fluid, distinct from the common juices of the plant, might lead us to sus- pect the operation of that function, although the part should display no organic peculiarity sufficient to authorize the application of the term gland to it. On the other hand, a definition founded on struc- ture alone, or on figure or position, would lead us as far astray. Perhaps we shall arrive nearer the truth if we take into consideration both structure and function, and say that a vegetable gland is a minute organ, differing in structure from the com- mon texture of the part where it is situated, and separating some peculiar matter from the ordinary v eg eta b le fluids. Guided by this definition, we find glands on the stem and leaves, situated both under the cutis and on its surface. In describing these organs some arrangement is necessary ; and in looking into books to know what has been done in this way, we find that the arrangement proposed by Guet- ss3 630 CONSERVATIVE ORGANS. [LECT. XI. tard, a French Phytologist, who first attempted the subject, has been adopted by almost every succeeding writer. He describes seven species of vegetable glands : the miliary, the vesicular, the scaly, the globular, the lenticular, the utricular, and the cup-shaped; but as the cuticular aper- tures, which he mistook for glands, constitute his first species ; and the thin scales which cover the fructification of the Ferns his third, there is sufficient reason for rejecting this arrangement. In attempting another, we must first take into consideration the situation of these minute organs as far as relates to their being under the cutis, or exterior to it; and, consequently, I propose to divide them into two classes, internal and external, and to subdivide these into genera and species. A. The INTERNAL caulinar and foliar glands are probably of various kinds, but one only, the follicular, has yet been detected. 1. The follicular gland, glandulafollicularis, is in the form of a small sac or follicle. It is ge- nerally found in the substance of leaves, and is furnished with an excretory duct which opens upon the under disk of the leaf. It is readily disco- vered by its transparency, which gives the leaf a punctured appearance when it is held up between the eye and the light ; as exemplified in the leaves of Perforated Saint John's Wort, Hypericum per- foratum ; All Spice, Myrtus Pimenta ; the genus LECT. XI.] CAULINAR AND FOLIAR APPENDAGES. 631 Citrus, &c. A beautiful variety of this gland is observed on the leaf of Coffea AraUca, si- tuated in the angles formed by the parting of the larger costse from the midrib : its follicle, which occupies a space rather greater than the thick- ness of the leaf, forms a small elevation on the upper disk, and opens on the lower by a large ex- cretory pore, guarded by stiff hairs inclining over it. The real glandular part of the follicular gland is cellular, and forms the parietes of the fol- licle, which is the receptacle only of the secre- tion; and this, in general, is an essential oil. As the odours of leaves depend chiefly on the exhala- tion of their essential oil, they are often regu- lated by circumstances affecting the excretory ducts of these follicles. Thus the duct being closed by the pressure of the cells turgid with sap, in the fresh stem and leaf of Sweet-scented Vernal Grass, Anthoxanthum odoratum, no odour is per- ceived ; but it opens, when these cells shrink, as the grass dries, and, then, the agreeable perfume which is peculiar to new hay is exhaled. The odour is permanent in some plants, as Mint, Sage, &c. but very evanescent in others, as, for example, in the leaves of Gaultheria odorata, which yield a very agreeable odour when fresh, but become scentless a few hours after their separation from the tree. B. The EXTERNAL caulinar and foliar glands S 8 4 632 CONSERVATIVE ORGANS. [LECT. XI. are more easily detected than the internal, and appear to be more numerous. They may be ar- ranged under two genera, the sessile and the pe- diculated; and each of these subdivided into species. a. The sessile gland lies on the surface of the stem, or of the leaf, or is slightly depressed in the cutis. It comprehends three species : the simple papillary, the compound papillary, and the scaly glands. 1. The simple papillary gland, glandula pa- pillaris simplex, is usually situated on the lower disk of leaves; and, in many instances, it appears, to the unassisted eye, a mere pore, exuding a drop of viscous or oily fluid ; but is, in fact, a small tubercular elevation. Thus on the back of the leaf of Crescent-leaved Passion-flower, Passiflora la- nata, the dark spots seen by the naked eye on each side of the midrib, are found to be slightly elevated, circular, papillary glands, with an apparent pore in the centre (Plate 9, fig. 27. a.) when viewed with a good glass ; but when the gland is placed under the microscope, the pore is discovered to be a de- pression only, covered with a very thin transparent epidermis, which extends over the whole surface of the gland. In general, however, the papillary glands are more conspicuous ; and appear, even to the unassisted eye, small elevated bodies, with a broad base, placed, in some instances, as it were, in a socket. Their structure is cellular; but the LECT. XT.] CAULINAR AND FOLIAR APPENDAGES. 633 cells are smaller, more regular than those of the substance of the leaf, and arranged in circles. I have not been able to detect any vessels passing into these glands, nor to discover their excretory pores ; except in the sting of the nettle, which is the excretory duct of a papillary gland. A va- riety of this gland, generally described as a dis- tinct species, under the name lenticular, glandula lenticular is, is found on the surface of the stems of Stripe-flowered Psoralea, P. glandulosa, and of many other dicotyledons. It is a small follicle, which generally contains an oily or a resinous fluid ; and differs from the internal follicular gland only in its situation on the surface. 2. The compound papillary gland, glandula papillarum composita, is best exemplified as it is found on the leaf of the Stone Pine, Pinus pinea. If we examine the surface of this leaf, with a good magnifying glass, we find that it is streaked with whitish lines, on which are seen small black spots arranged in a regular series (Plate 9, fig. 28. A.). Under the microscope each of these spots is discovered to be the excretory pore of a com- pound gland, composed of six distinct glandules, forming a ring or collar (fig. 28. B. a.) around the pore (&.), which generally appears obstructed by minute, dark-coloured, terebinthinous par- ticles ; and is seen to be really a pore, only when the cutis is very carefully raised. The glandules 634 CONSERVATIVE ORGANS. [LECT. XI. resemble the papillary glands in some degree, each being a slight elevation with a depression in the centre; and it is probable that these are the secret- ing organs, and, severally, communicating with the excretory pore, pour the fluid they secrete into it; whence it is exuded: and acquires its dark colour by exposure to the atmosphere. 3. The scaly gland, glandula squamom*, re- sembles a minute scale attached to the surfaces of the leaves on which it is found, as, for instance, on that of Rhododendron punctatum. The under surface of this leaf is covered with glands of this species, which appear, when viewed with a good glass, like small brilliant detachable scales, white and shining round the edge and dark in the centre (Plate 9, fig. 29. A. a.). When one of these scales is placed under the microscope, the white border is found to be beautifully fluted (fig. 29. B. b. c.), and lying flat upon, but not attached to the surface of the leaf ; and to be the loose margin of the scale covering the gland, which appears to be a slightly elevated papilla, discharging its secretion by se- veral excretory pores which open upon the surface. The real shape of the gland, however, is that of an inverted cone, of which the scale is the base. * I am surprised that the error of Guettard, who gave this name to the thin scale which covers the fructification of Ferns, has been copied by Mr. Keith in his System of Physio- logical Botany. See vol. .i. p. 69. LECT. XI.] CAULINAR AND FOLIAR APPENDAGES. 635 It is seated in a depression of the cutis, and com- municates with the interior of the leaf by means of a duct, which passing through the apex to the base of the cone penetrates the cutis. The texture of this gland is cellular ; and its secretion is an essential oil. The leaves of the Sea-side Balsam, Croton Eluteria, furnish examples, also, of the scaly gland *. b. The pediculated gland is elevated from the surfaces on which it is found, by an interposed pedicle or stalk. It comprehends four species, the cup-shaped, the knob-like, the stipitate, and the branched glands. 1. The cup-shaped gland, glandula cyathifor- mis 9 as its name imports, resembles a shallow cup or saucer, supported on a thick, short, footstalk (Plate 9, fig. 30). It is found on the petioles of some leaves, as, for instance, those of the Nec- tarine, Amygdalus Persica, and the Passion- flower ; and in the serratures of others, as those of the Bay-leaved Willow, Salix pentandra, &c. The secretion is generally of a resinous character, and exudes from the hollow part of the gland, which is devoid of epidermis. This species of gland, like the majority of the external glands, is cellular, and we can distinctly trace into it a cord of both * Sprengel has figured these in his Elements of the Phi- losophy of Plants (P.-vi. fig. 8); but regards them merely a* scales. 636 CONSERVATIVE ORGANS. [LECT. XI. spiral and proper vessels, which, apparently, ter- minates in the substance of the gland. 2. The knob-shaped gland, glandula clavifor- mis, resembles a knob or a small nail, which, in the language of the artisan, has not been driven home. The head, which is the glandular part, is slightly convex on the upper surface; and displays a rough striated border, encircling a round flat spot devoid of epidermis (Plate 9, fig. 31), from which the secreted fluid is discharged. The in- ternal structure of this gland closely resembles that of the cup-shaped gland. It is generally found either on the stem or the petiole of the leaf, or on both in the same plant, as for instance in the Castor-oil plant, Ricinus communis. 3. The stipitate gland, glandula stipitata, is so named from being supported on a long slender stalk (Plate 9, fig. 15), and is, in fact, a stalked variety of the cyathiform gland. It is the smallest of the external glands, and is situated either on the margin of leaves, as in Croton penicillatum ; or on the disk, as in Sun-dew, Drosera rotundifolia. 4. The branched pediculated gland, glandula pediculata ramosa, is a small hemispherical gland supported on a branched stalk (Plate 9, fig. 32). The appearance of moss on the stem of the Moss Rose is produced by glands of this description. The individual glands are cellular, devoid of epi- dermis, and each is furnished with vessels, which are branches of a fasciculus, which enters the LECT. XJ.] CAULINAR AND FOLIAR APPENDAGES. 637 gland-bearing stalk at its base. These glands ge- nerally secrete a viscous resinous matter, which is also sometimes odorous. The minuteness of the vegetable glands pre- vents me from attempting to offer you any anato- mical demonstration of their structure. They have, indeed, been arranged into cellular and vas- cular glands by writers who have endeavoured to trace their structure, and who have stated some of them to be entirely cellular, and others chiefly vascular ; but as we can scarcely suppose that any part, completely devoid of vessels, possesses the function of secretion, we must receive these state- ments with caution. With regard to the use of glands to the plant, at least as far as relates to the conservative organs, we know nothing. They, probably, do not play so important a part in the vegetable as in the animal economy ; but we can hazard nothing more than conjecture on the sub- ject. ii. PUBESCENCE, Pubescentia. Under this term are included all the kinds of down, hairs, and bristle-like bodies, found on the surfaces of the conservative organs. They differ very considerably in form and texture, and on these differences I shall attempt such an arrangement of them as may facilitate your knowledge of the subject. All vegetable pubescence consists of small, slender bodies, which are either soft and yielding to the slightest impression, or *rigid and compa- 638 CONSERVATIVE ORGANS. [LECT. XI. ratively unyielding; the former are, properly speaking, hairs (pili), the latter bristles (setce) ; and, therefore, under these two heads every kind of pubescence may be arranged. A. HAIRS, Pili, are fine, slender, cylindrical, flexible bodies found on the surfaces of the herba- ceous parts of plants. Some of them are the excre- tory ducts of glands, a fact which was first de- tected by Guettard ; and on which Linnaeus too hastily formed his definition of the hair, which he describes generally to be > we have a view of the costa which terminates in the hinge. LECT. XI.J CAULINAR AND FOLIAR APPENDAGES. 679 the air is charged with moisture, the scroll swells, and by unrolling itself enlarges the thickness of the ring, and thus produces a pressure upwards and backwards, by which the lid is forced open. I have found, from experiment, that the en- largement of the ring one sixteenth of an inch, would bring the lid nearly into the erect position ; and it is evident that the opening of the lid must follow such an enlargement of the ring, unless we suppose that the hinge length- ens synchronously with the unrolling of the scroll, which is not probable. In advancing this explana- tion, I must allow that it is purely hypothetical, and requires to be confirmed by observations on the habits of the plant in its living state. With regard to the fluid found in these vege- table pitchers, the most probable opinion is, that it is obtained from the atmosphere, and is intended for the nourishment of the plant; for we can scarcely suppose that so large a quantity of mois- ture can be thrown out as a excretion, in a plant growing in the dry, sterile situations where Ne- penthes is found, and for the sole purpose of drown- ing a few insects*. Rumphius indeed observes, that the insects which crawl into the pitcher all die, * Sir E. J. Smith supposes these pitchers to be merely fly- traps; and, speaking of the source of the fluid, says it is " certainly secreted through the footstalk of the leaf." Introd. to Phys. and Syst. Bot, p. 197. x x 4 680 CONSERVATIVE ORGANS. [LECT. XI. " except a small squilla or shrimp, with a gibbous " back, sometimes met with, which lives there;" but it does not appear that putrefaction goes on in these pitchers, and the constant renewal of the water is, certainly, not favourable to this process. . Another pitcher is the tubular leaf of Sarrace- nia, for it can scarcely be termed an appendage, seeing that it does not issue from a leaf, but originates, directly, from the collar (collet) of the plant, which is stemless. In the species before us, S. purpurea, the hollow part of the leaf (see 4, page 491) is an infundibuliform, ventricose cavity, almost tubular below, and having a wide open mouth, bounded in the fore part by a revolute lip, and backwards by a broad dilated margin, auricled on each side, and somewhat resembling the spout of an ewer. In two of the species, however, S.jlava and S. adunca, this expanded part of the lip projects forwards over the mouth of the cavity, and constitutes a kind of lid, al- though not moveable. The lower part of the tube terminates in a thick spongy cord, that ul- timately expands to form the part which attaches it to the plant ; and the remainder of the leaf is a thin expansion, extending along the whole front of the tubular part. Dividing the funnel or pitcher longitudinally, we perceive that its narrower or lower half is devoid of any proper cutis, and lined LKCT. XI.J CAUL1NAR AND FOLIAR APPENDAGES. 681 with a reticulated epidermis only, which is studded with long hairs all pointing downwards, and co- vers a number of glands, the excretory pores of which open upon its surface. The ventricose part of the funnel is smooth, shining, and lined with a cutis closely resembling that of the outside of the leaf; this terminates, however, by a well-defined line a little below the lip ; and at the faux, if it can be so termed/ of the funnel, we again find a band of the same reticulated, glandular epidermis as at its bottom, except that it is devoid of hairs, and exudes a sweet, viscid secretion. The whole of the interior surface of the dilated lip or hood, is co- vered with short stiff hairs, all pointing down- wards. The pitchers of Sarracenia vary consi- derably in size; some, particularly those of S. flava, being often more than three feet in height. All of them contain some fluid. The intimate structure of the whole is the same as that of the leaf. If the use of the pitchers in Cephalotus and Nepenthes be problematical, there is sufficient evi- dence for asserting, that those of Sarracenia are chiefly intended for fly-traps; and it is very pro- bable that the gases, and the other results of the putrefactive process, produced by the bodies of the insects which enter them and die there, may be essential to the healthy economy of the plant. According to some observations on the power of Sarracenia to entrap insects, made by Dr. J * 682 CONSERVATIVE ORGANS. [LECT. XI. Macbride, of South Carolina, it appears that the flies, which are attracted to these leaves, first alight upon the faux of the pitcher, and appear to sip, with eagerness, the sweet secretion exuded from the glandular band, which you have seen is situ- ated just below the margin. " In this position they " linger ; but at length, allured as it would seem " by the pleasure of taste, they enter the tubes. " The fly which has thus changed its situation, will " be seen to stand unsteadily, it totters for a few " seconds, slips, and falls to the bottom of the tube, " where it is either drowned, or attempts in vain to " escape against the points of the hairs*." Dr. Macbride attributes the fall of the fly to the down- ward or inverted position of the short attenuated hairs at the faux, on which the fly is unable to take a hold sufficiently strong to support itself; but I am inclined to think that a kind of intoxication is produced in the insect, either by the secretion it sips, or by some exhalation within the pitcher ; for I have seen flies, which had fallen, take wing within some of the large pitchers, and again drop before they had reached the faux : and we may infer that this frequently occurs from the humming noise within the pitcher, which is never heard, but when a fly is using the wings. That this seldom occurs in S. adunca, the species on which Dr. Macbride made his observations, is probable, be- * Linnean Transactions, vol. xii. p. 4-8. LECT. XI.] CAUL1NAR AND FOLIAR APPENDAGES. (583 cause the faux in this species is covered by the hood, and the light excluded. If the fly attempts to crawl up, the inverted position of the hairs is a sufficient obstacle to its escape. Spiders, a small species of Phalaena, and some other insects which enter these tlibes, appear, however, to ascend without difficulty. I have, already, stated my accordance in the opinion that these putrefying masses may be be- neficial to the growth of the plant ; but this conjecture, Mr. Keith observes *, " cannot be re- " garded as quite satisfactory till such time as it " shall be shown that the health of the plant is " injured when insects are prevented from ap- " proaching it." It is curious to observe, that these fly-traps become serviceable to some indi- viduals belonging to the very division of the animal creation which they serve to destroy. Dr. Macbride says, " in the putrid masses of " insects thus collected, are always to be seen " one or two maggots in a very active state." He was unable, for some time, to ascertain the in- sect to which these belonged ; " but while watch- " ing attentively some tall tubes of Sarracenia " flava growing in their natural situation, a large V' he remarks, " caught my attention: it * Syst. of Physiol. Bot. vol. ii. p. 286. f This viviparous musca was more than double the size of the common house-fly, had a reddish head, and the body hairy, and streaked greyish. Lin. Trans. I. c 684 CONSERVATIVE ORGANS. [LECT. XI. " passed rapidly from one tube to another, delay- " ing scarcely a moment at the faux of each, " until it found, as it should seem, one suitable " to its purpose; then, hanging its posterior ex- " tremity over the margin, it ejected, on the in- " ternal surface of the tube, a larva with a black " head, which immediately proceeded downwards " with a brisk vermicular motion." Sir J.E. Smith, also, notices an observation made at the Botanic Garden at Liverpool, which shows that, even in this country, the Ichneumon fly employs the pitchers of these plants for a similar purpose. An Ichneu- mon was, one day, observed by one of the gar- deners forcing large flies into the tubular cavity of a leaf of Sarracenia adunca; and it is probable, that it had previously deposited its eggs in their carcasses, the eggs of the Ichneumon being often deposited, and the larvae hatched, in the carcasses of other flies or their larvae*. c. Peduncular ascidium. This description of organ is rarely found on the peduncle. On Surubea Guianemis -f~, however, we find a small hollow body, connected with a singular forked projection, which rides as it were across the flower-stalk: and a similar body, but without the fork, is observed on the peduncle of Ruyschia * Some Ichneumons deposit their eggs in the aurelia of moths and butterflies. t Aubl. Boyl. Meyer, Fl. Essequtb. p. 120. LECT. XI.] CAULINAR AND FOLIAR APPENDAGES. 685 clusicefolia *, and that of Marcgrafia. In these instances, however, the name ascidium can apply to the form only of these organs, as they do not appear intended to hold fluids ; and perhaps they may rather be regarded as ascidiform bracteae than asascidia-f~. vii. Anomalies. The only appendage which I shall notice under this head > belongs to a North American plant, Venus' Fly-trap, Dionoea musci- pula ; and it is certainly the most extraordinary production of the vegetable kingdom. The leaf of this plant is radical, sessile, and nearly spa- tulate in figure (Plate 10, fig. 5. c.) : the midrib, however, is produced beyond the apex of the leaf, and supports ^an appendage which has some re- semblance to a steel trap. It consists of two lobes, almost elliptical (fig. 5. b.), connected to- gether by a whitish, cartilaginous costa, which is, apparently, a production of the midrib of the leaf. The lobes resemble the leaf in colour and con- sistence, but their margin is somewhat cartilagi- nous, and furnished with long setaceous teeth, placed at the distance of the tenth of an inch from one another. The superior disk of each lobe is studded with minute glands, and furnished with erect little spines (fig. 5. a.), placed so as to form an equilateral triangle, with the apex pointing * Jacq. Amer. Tab. 51. fig. 2. f They have been termed Anthocerynium. 686 CONSERVATIVE ORGANS. [LECT. XI. towards the midrib which unites the lobes. To the naked eye, these spines appear like simple Tmstles ; but when they are examined under the microscope, each spine is found to consist of two distinct parts ; the one (fig. 6. a.), a small cellular papilla; and the other (&.), which is supported on its apex, a firm tapering pointed body, resembling a small, in- verted bodkin. This appendage is endowed with so much ir- ritability, that as soon as a fly or other insect alights upon the upper disk of either of the lobes, so as to touch any of the spines, the lobes, if the plant be in a healthy condition., immediately close upon it; and the spines either empale the little animal, or the teeth on the edges of the lobes, crossing one another, prevent its escape, and detain it until it dies. The anatomical examination of these lobes does not elucidate the phenomenon connected with their functions. They resemble a leaf in their cellular and vascular structure: the vessels being given off from the midrib in arching fascicles, which anastomose, towards the outer margin of each lobe. The epidermis is glandular, and is probably the seat of the irritability of the ap- pendage; but, I have not had an opportunity of examining it with sufficient care, to enable me to determine in what particulars it differs from that of the leaf. LECT. XI.] CAULINAR AND FOLIAR APPENDAGES. 687 Mr. Ellis, who first described this curious appendage*, imagined that the glands were the ir- ritable parts; and that, as soon as an insect touched any of these, the motion of the lobes was produced. Every other succeeding writer has stated, generally, that it is merely necessary to touch the upper surface of the lobes to excite their action. From a variety of experiments, however, I am convinced that no motion is produced until one or other of the spines be touched; and, from the manner in which these are affixed to the pa- pillae, and the connexion of the latter with the cuticle of the lobes, I am induced to suppose, that the touching the spines communicates a thrill or tremor to every part of the surface on which they are situated, which excites into action the irri- tability on which their motion depends. It may be stated as an objection to this opinion, that it is altogether hypothetical ; but no hint that can tend to explain so interesting a phenomenon should be withheld. As the intention of this appendage is evidently * " The Dionoea was first brought to this country, in the " summer of 1768, by Mr. Young, gardener to the Queen; " and Mr. Ellis described it, and had a drawing and a plate " engraved from a plant which flowered in his chambers in the " following August. It was from this plate and his characters of " the plant, that Linnaeus's description was drawn up for his " Mantissa. '* Vide Smith's Selection of the Correspondence of Linn&us, vol. i. p. 235, and vol. ii. p. 72. 688 CONSERVATIVE ORGANS. [LKCT. Xf. to intrap flies and other insects; this question presents itself what benefit can result to the plant from such a function? We may reply, that there is reason for supposing that the plant derives some advantage either from the putrefac- tion of the dead insects, or from something which can be obtained from animal matter; a suppo- sition, the probability of which is much strength- ened by an experiment made by a Mr. Knight *. Having laid fine filaments of raw beef upon the appendages of a plant of Dioncea, he found that this plant was more luxuriant than any other in the same place, although they were all treated alike, with the exception of the supply of beef. That some principle, therefore, is evolved during the decomposition of animal matter, peculiarly favourable to the growth of this plant, is pro- bable, and to secure this, may be the intention of its singular appendages. An appendage resembling that of the Dionoea, in miniature, is found upon the glume of another North American plant, the Leersia lenticularis. * Mr. Knight was, at the time of making the experiment, gardener to George Hibbert, Esq. and is now a respectable nursery-man in the King's Road, Chelsea. INDEX FIRST; CONSERVATIVE ORGANS METHODICALLY ARRANGED. The Numbers indicate the Page. THE CONSERVATIVE ORGANS comprehend, A. The root; B. The stein and branches ; C. The leaves; D. Appendages, A. THE ROOT, Radix, consists of Caudex, caudex, 128; Rootlets, radiculee, 129; fibrils, fibrillce, 129. In situation, roots are subterraneous, subterraneee, 189: aerial, aerece, 190: floating, natantes, 191 : parasitical, parasiticts, 189. In direction, perpendicular, perpendicular ~es, 192 : horizontal, horizontales, 192 : oblique, obliques, 195. In duration, annual, annute, 208: biennial, biennes, 208 : perennial, pcrennes, 208. In substance, woody, lignosee, 199: fleshy, carnosce, 190. Inform and composition, simple, simplices, 130: branched, ramosee, 1 39 : articulated, articulate*, 140. The simple root is Sp. 1. conical, conica, 130; var. a. spindle-shaped, fusi- formis, 131 : b. truncated, preemorsa, 132: VOL. r. Sp. 2. subglobular, subrotuitda, 134. var. a. turnip-shaped, rapifor- mis, 134. 6. flattened, placentifor- mis, 134. 3. fibrous, fibrosa, 136. var. a. filiform, > /?/(/brmtJ, 138. 6. capillary, capillarif,}38. c. cord-like, funiliformis, 139. The branched is Sp. 1. branched, ramota, 140 : 2. toothed, dentata, 140. The articulated is Sp. 1. jointed, articulata, 141 : 2. kneed, geniculata, 141: var. a. contorted, contoi'ta, 142: i. necklace-like, monilifm- mis, 1 42. The appendages of the roots are scales, squamee, 144: suckers, stolones, 145: tubers, tubera, 146 : bulbs, 6u/6t, 164. The fier is /i. closely attached, conjunctum ; comprehending, Sp. 1. the conjoined ovate, conju- gatum ovatum, 149. 2. conjoined club-shaped, clavceforme conjttnctum, 150. 3. fingered, digit atum, 151. 4. palmated, palmatum, 151. 5. prehending, INDEX FIRST. Sp. 1. the solitary, solitarium, 153. S. congregated, congrega- tum t 154. The bulb is a. solid, solidus ; comprehending Sp. 1. the superincumbent, soli- dus superpositus, 1 66 : 2. lateral, laterally, 169: 3. inclosing, includens, 171. 6. scaly, squamosus, Sp. 1. the squamous, squamnsus, 172: 2. granulated, granulatus, 178. c. Laminated, tunicosus, Sp. 1 . Concentric, concentricus, 179: 2. nestling, nidulans, 186. B. THE STEM comprehends The trunk, truncus, 279. stalk, caulis, 280. straw, culmus t 281. scape, scapus, 281. stipe, stipes, 282. In composition, the stem is simple, simplex, 254. divided, ramosus, 255. In direction, 1. Erect, erectus, 244. var. a. straight, strictus, 245. 4. fiexuose, flexuosus, 245. c. tortuous, toi'tuosus, 245. ^/t transversis intertinctus, 287. In/orw, Sp. 1. round, teres, cylindricus, 272. 2. half round, semiteres, 273. 3. compressed, compressus, 273. var. a. two-edged, anceps, 273. 5. niembranaceous, ^Ay/- loidcus, 273. 4. angled, engulatus, 273. var. a. obtuse, obtuse angula- tus, 273. ub-var. a. three-corner- ed, trig-onus, 274. sub-var. 3 four-corner- ed, tetragonus, 274. sub-var. /"> 492. two-edged, adcipitcs, 492. spherical, spfueroidea, 492. ovoid, ovoidea, 492. coccoon-shaped, fusina, 492. club-shaped, clavata, 492. hooked, nncinata, 493. lenticular, lenticularia, 493. The opej; of the leaf is acute, acutusy 493. sharpish, acutiusculus, 493. spine-pointed, cuspidatus t 493. muc ron ate, mucronatus, 493. awned, aristatus, 493. cirrhose, 'cirrosus, 494. obtuse, obtusiis, 494. var. obtusus cum acumine, 496. thickened, incrassatus, 496. retuse, retusus, 496. emarginate, emarginatus, 496. truncated, truncatus, 496. , jagged, prasmorsus, 49S. tridentate, tridentatus y 496'. In respect to 6awe, the general figure gives its character; but a leaf is also unequal, iiuequale, 497. The margin of the leaf is * entire ; ** indented; *** bordered; **** rolled. * entire, integerrima y 497. # sinuatcd sinuata, 497. gnawed, erosa, 497. toothed, dentatay 497. var. a. equally, tequaliter, 498, 6. unequally, ineequalitery 498. c. deeply, profundcy 498. J. verticillato-pinnnta, 523. vertcbrated, vertebrata, 596. INDEX FIRST. The doubly compounded are twice paired, bigeminata, 523. thrice paired, tergeminata, 524. twice ternate, biter nata, 524. doubly pinnate, bipinnata, 524. conjugated and pinnate, conjugata- pinnata, 524. ternate and pinnate, ternata-pin- nata, 524. digitated and pinnate, digitata-pin- nata, ib. The much compounded are thrice ternate, triternata, 525. triply pinnate, tripinnata, 526'. In substance the leaf is membranous, membranaceum> 510. chaff-like, scariosum, 511. chartaceous, chartaceum, 511. leathery, coriaceum, 511. rigid, rigidum, 511. fleshy, carnosum, 511. succulent, succulentum, 511. In productions it is spiue-bearing, spiniferum, 541. leaf-bearing, foliifcrum, 541. flower-bearing, fl&riferum, 541. plant-bearing, proliferum, 541. D. GENERAL APPENDAGES: these comprehend, i. Glands, glandules, 628. ii. Pubescence, pubescentia, 637. iii. Thorns, spince, 647. iv. Prickles, aew/a, 649. v. Props, fulcra, 652. vi. Foliaceous appendages, appendi- culifoliacei. vii. Anomalies, anomnli. i. Glands, glandula, are internal, external. Internal glands are, Sp. 1. the follicular gland, g-. /b//t'-' cularis, 630. External glands are sessile, pediculated. * The sessile^ comprehend Sp. 1. the simple papillary gland, #. papillaris simplex, 632. var. glandula lenticular is, 633. 2. compound papillary, papilla- rum composita, 633. 3. scaly, squamosa, 634. ** The pediculated. Sp. 1. the cup-shaped, cyathiformis, 635. 2. knob-shaped, clavifurmis, 636. 3. stipitate, stipitata, 636. 4. branched, ramosa, 636. ii. Pubescence, pubescentia, com- prehends hairs, />i7t, 638. bristles, jeftp, 638. A hair is, Sp. 1. Simple, pilus simplex, 639. var. a. awl-shaped, subulatus, 640. 6. hooked, hamosus, ib. c. gland-bearing, glandulosu*, ib. 2 . compoun d , /w7e/,y compositus, 640. var. a. feathery, plumosus, ib. 6. branched, ramosuy, ib. c. starlike, stellatus, ib. The bristle is, Sp. 1. Simple, simplex, C41. var. a. awl-shaped, subulata, 642. i. spindle-shaped, fusifor- mis, 643. 2. compound, *e/ composita, 643. var. . forked, furcata, 644. 6. fasciculated, fascicu- lata, ib. iii. Thorns, spinee, are, Sp. 1. Simple, spina simplex, 647. var. solitary, solitaria, ib. 2. compound, spina composita, ib. var. a. forked, bipartita, 648. 6. three-pronged, tripar- tita, ib. c. branched, ramosa, ib. In situation thorns are terminal, terminales, 648. axillary, axillares, ib. superaxillary, super axillares, ib. subaxillary, infer axillares, ib. iv. Prickles, aculei, 649. Sp. 1. straight, ratft', 660. 2. curved, eww, ib. var. a. incurvi, ib. 6. recurvi, ib. 3. spiral, circinnati, ib. In composition they are in pairs, geminati, 650. palmated, palmati, ib. v. Props, fulcra, comprehend * tendrils, cirrhi, 653. ** claws, claviculi, 656. Y Y 4 INDEX FIRST, *** bladders, ampulla, 661. **** hooks, hamiy 662. The tendril is, Sp. 1. Simple, cirrhus simplex, 653. 2. compound, compositus, ib. var. a. bifid, bifidus, ib. i. trifid, trifidus, ib. c. branched, ramosus, ib. In situation the tendril is axillary, axillaris, 653. subaxillary, subaxillaris, ib. lateral, laterally, tb. opposite, oppositifolius, ib. petiolar, petiolaris, ib. foliar, foliar is, ib. In direction : convolute, convolutus, 656. revolute, revolutus, ib. ** The c/rtw is, Sp. 1. cirrhal, cirriformis, 656. 2. radicular, radiciformis, ib. *** the bladder, ampulla, 661. **** the hook, hamus, 662. vi. Foliaceotts appendages are, Sp. 1. the stipule, stiptila, 662. In duration it is fugacious, fugaz, 663. deciduous, decidua, 663. persistent, persistens, ib. In number : solitary, solitaria, 664. twin, gemina, ib. In substance : leafy, foliacea, 664. membranous, membranacea, ib. chaff-like, scariosa, ib. In situation : lateral, lateralis, 665. intermediate, intermedia, ib. embracing, amplexans, ib. sheathing, vaginosa, ib. var. hypocrateriformis, ib. marginal, marginalia, ib. interfoliaceous, intrafoleacea, ib. extrafoliaceous, extrafoliacea, ib. 2. The bracte, bractea, 667. var. a. spathe, spatha, ib. A. involucre, involucrum, ib. 3. The pitcher, ascidium, 669. Sp. 1. caulinar, caulinum, ib. 2. foliar, epiphyllum, 672. 3. peduncular, pedunculare, 684. vii. anomalous, anormcs, 685. INDEX SECOND. A. AERIAL, application of the term to leaves, 628. u?Esculus Hippocastaminiy structure of its gems, 463. Agave Americana, its foliar apertures, 611, 615. Agostino Ma7idirola, extract from his treatise, 545. Air, a component of soils, 218. vitiated by vegetating roots, 219. kind of, contained in hollow stems, 424 . Alburnurft, its character, 336. changes into hard wood, 337. " hardens when a tree is barked, 337. simple hardening of, does not constitute wood, 338, formation of the, 340. dies when exposed to the air, 354. is a reservoir of proper juice in winter, 355. ^ loses its irritability on becoming- wood, 856. Algae are propagated by the propago, 450. Alienated leaves, their character, 528. Aloje, its leaf has few pneumatic apertures, 618. soccotrina, its proper juice affords a dye, 118. mode of its growth, 205. structure of its leaf, 562. verrucosa, form of its cuticular meshes, 605. Althaea officinalix t structure of its root, 439. Annual herbaceous roots, structure of, 431. i zones of wood, origin of, 338. Annular vessels, their characters, 82. Anthoxanthum odoratum is odorous only when dried, 631. Apertures, foliar, 608. their varieties, 6o9. their size and number, 613. are the exhaling organs of leaves, 610, 619. are the respiratory organs, 62 1 . Aphis, its lateral propagation, 457. Appendages, enumeration of, 628. Aquatic plants, structure- of their foliar parenchyma, 596. . the leaves have no pneumatic slits, 608. Arctium lappa, structure of its root, 433. - has the largest leaf of any British plant, 546. Argillaceous earth, its effects in soils, 214. Ascidium of Cephalotus/oWicM/ara, 670. Nepenthes Distillaloria, 672. Atropa Btlladonna, structure of its root, 439. 2 INDEX SECOND. B. Banyan tree, a remarkable one, 198. Barclay, Dr. his opinion respecting the cells of honeycomb, 373, Bark, structure of the, 317. reproduction of the, 848. Barley indicates a poor soil, when it has long roots, 224. Bauer, Mr. Francis, his opinion of the cuticle, 600. Bauhin, Casper, first described the potatoe, 27. Beauvois, M. de, his opinion respecting the medullary tube, 867. Biennial herbaceous roots, structure of, 343. Boerhaave, his definition of a plant, 42. Bonnet, his remarks on absorption by leaves, 617. Botany, derivation of the term, 2. its antiquity, 6. its utility to medicine, 19. its utility to agriculture, 28. its importance as a branch of general education, 29. Bradley, Mr. his theory of grafting, 397. Branch, origin of the, 384. organization of its germ in the bud, 386. is fully formed when protruded from the bud, 388. . origin of the adventitious, 391. buds, when formed, 395. remain latent, 395. * progress of its germ, when it is not evolved, 403. Branching, its theory illustrated by a diagram, 404. . new theory of, 391 405. Brunfels, the restorer of the science of Botany in Europe, 10. Bryonia alba, structure of its stem, 414. its stem contains three kinds of vessels, 419. Buds, supposed by Darwin to have a sensorium, 376. origin of, 485 395. contain the germs of future branches, 386. theory of their formation, 391. do not proceed from the alburnous vessels, 393. branch, when formed, 395. are distinct individuals, 395. stem, when formed, 395. effects of temperature on, 399. Bulbs, how to distinguish from tubers, 165. -- - are reservoirs of nutriment, ib, . .. are hybernacula, 165. are the lateral progeny of the plant, 165. are exhausted in producing the stem and leave*, 169. are subterraneous buds, 1 87. scales of, can produce new bulbs, 170. flowering, the last of the natural series, 187. their vessels may be injected, 181. their spiral vessels are easily detected, 184. new leaf, are the lateral progeny of the old bulbs, 185. new, sometimes formed within the old, 181. cauliriar, structure of, 453. erroneously regarded as gems by Linnaeus, 453. . produce leaves only, 455. Bulliard detected the seeds of fungi, 452. INDEX SECOND. C. Cactus, structure of its foliar cells, 592. Ceesalpinus first classed plants, 1 1 . Calcareous earth, its use in soils, 213. too much, injurious in soils, 214. Calla jtEthiopica, structure of its leaf, 588. Caloric, a part in soils, 221. Cambium, 341. Camcrarius first proved the sexuality of plants, 1 2. Canna Indica, structure of its leaf, 566. Carbonic acid gas, present in soils, 220. Carbonated hydrogen gas, formed in soils, 220. Carex arenaria, utility of, 1.93. Caterpillar, its tenacity of life, 62. frozen, lives after it is thawed, 476. Caudex, a reservoir of nutriment, 129. structure of, in dicotyledons, 406. Cells, how those of the pith are formed, 372. symmetrical arrangement of, in some annual roots, 432. some have transverse slits in their sides, 432. longitudinal, in some roots, are valvular, 433. lateral communication of, 595. their sides are double, 324. Cellular integument, its character, 321. is the seat of colour, 322. texture, its nature, 71. - pores, their existence doubtful, 826. Cephalotus/o//tctt/rt4man,.ffusyt.fiees, Orme, Fit?. 2. A. Fy.Z , Hiuyt, Rets, Ortne,, &yll& m Fiq 2 B fy3 ' ,iri,f Fig 2 *v?.a fy W.I ., Surst.Reet . 0-rme. Mg. Iff. Kriifr-i,: . 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