feed's Series of Pocket 01 i v/ PHYSIOLOGY; A MANUAL FOR STUDENTS AND PRACTITIONERS, BY HOWARD D. COLLINS, M.D., Assistant to the Attending Surgeon of the Roosevelt Hospital ; Assistant Demonstrator of Anatomy, College of Physicians and Surgeons (Columbia University), New York. WM. H. ROCKWELL, JR., M. D., Assistant Demonstrator of Anatomy, College of Physicians and Surgeons (Columbia University), New York; Member of Association of American Anatomists. SERIES EDITED BY BERN B. GALLAUDET, M.D., Demonstrator of Anatomy and Instructor in Surgery, College of Physicians and Surgeons, Columbia University, New York; Visiting Surgeon, Bellevue Hospital, New York. ILLUSTRATED WITH ONE HUNDRED AND FIFTY-THREE ENGRAVINGS. LEA BROTHERS & CO., PHILADELPHIA AND NEW YORK. Entered according to Act of Congress, in the year 1899, by LEA BROTHERS & CO., In the Office of the Librarian of Congress, at Washington. All rights reserved. WESTCOTT 4 THOMSON, ELECTROTYPFRS. PHILADA. PREFACE. IN preparing the present volume the authors make no claim to original research. Their purpose is, frankly, not only to help the student acquire a knowledge of physiology such as will enable him to read with better understanding the larger works on the subject ; but also to give him actually fuller and more accurate information than can be obtained from the various and popular quiz-compends. Thus, in each section, in addition to the purely physiologi- cal aspects, histological considerations are discussed, such as those of the cell in general and of the neuron in the nervous system ; and, wherever practicable, the application of physio- logical principles to pathological conditions is also dealt with. Acknowledgment as to source of material is made to the larger works of Chapman, Kirke, and Foster, to the manual by Dr. F. A. Manning, and to the lectures delivered at the College of Physicians and Surgeons of New York. HOWARD D. COLLINS, WM. H. KOCKWELL, JR. CONTENTS. PAGE GENERAL CONSIDERATIONS: Physiology; life; living body com- pared to a machine ; metabolism 17,18 CELLS : Structure ; gemmation ; karyokinesis ; development ; amoeba ; classification of tissues 18-24 CHEMISTRY OF THE BODY : Proximate principles ; inorganic ; organic 24-30 THE BLOOD: Gross appearance ; quantity; red corpuscles ; func- tion; globulin; hsematin ; leukocytes; lymphocytes; func- tion ; blood-plates ; plasma ; serum ; gases ; coagulation . . . 30-46 THE LYMPH: Composition; sources; uses 46-48 CIRCULATION OF THE BLOOD: Apparatus; course; heart; action of auricles and ventricles ; capillaries ; veins ; arteries ; speed ; blood-pressure; arterial tension; the blood in circulation; action of heart ; systole ; diastole ; arterial tension ; heart- sounds ; heart-innervation ; vaso-motor nerves ; pulse ; sphyg- mograph 48-70 "CIRCULATION" OF THE LYMPH: Lymphatics; stomata and pseudostomata ; flow of lymph ; sources ; pressure ; lymph- ganglia 70-75 RESPIRATION : Respiratory act ; movements ; tract ; larynx ; trachea ; bronchi ; lungs ; muscles : of inspiration ; of expira- tion ; forced inspiration; forced expiration ; sounds; tidal air; reserve air ; residual air ; complementary air ; expired air ; external respiration ; internal respiration ; nervous mechan- ism ; centres ; section of vagi ; vitiated atmosphere ; effect of respiration on circulation ; special respiratory acts 75-90 5 6 CONTENTS. PAGE DIGESTION: Mouth; mastication; teeth; saliva; ptyalin; deglu- tition; stomach; gastric juice ; peptones; gastric digestion ; vomiting; small intestine; intestinal digestion; intestinal juice; the pancreas ; pancreatic juice ; trypsin ; amylopein; steapsin ; the liver; secretion of the liver; bile; bile-salts; bile-pigments ; cholesterin ; excretion of the liver ; glycogenic function ; large intestine ; defecation . 90-108 ABSORPTION: Dialysis; sites of absorption; villi of intestine; changes in the products of digestion on being absorbed ; des- tination of absorbed food 109-112 SECRETION : Secretions ; excretions ; apparatus ; glands ; dis- charge ; correlation of secretions ; serous; mucous; mammary glands ; milk ; secretions of skin ; structure of skin ; seba- ceous glands ; sweat-glands; hair; nails; sweat; perspiration; absorption by skin; kidneys; structure of kidneys; blood- supply; ureters; bladder; urine; secretion of urine ; compo- sition ; acidity of urine ; course of urine ; amount ; micturi- tion ; urea; uric acid ; vascular glands ; table of secretions . . 113-134 NUTRITION : Sources of income to the body ; expenditures of the body; energy of the body: starvation; diet; overfeeding; nitrogenous equilibrium ; assimilation 134-138 ANIMAL HEAT: Normal temperature; sources of heat; loss of heat ; regulation of heat ; centres for heat-regulation ; limits of body-temperature . 138-140 MUSCLE : Varieties ; microscopic appearances ; general properties ; chemistry; physiology; muscles as levers; oxygen-supply; fatigue ; latent period ; contractility ; stimuli ; electrical cur- rents ; effect of galvanic shock ; electrical state ; excitability and conductivity ; Pfliiger's law of contraction ; ascending and descending currents ; nerve-muscle preparation ; rigor mortis . 140-159 NERVOUS SYSTEM: Fibres and cells; varieties of fibres; nerve- trnnks ; function of fibres ; nerve-cells ; varieties ; nerve-end- ing : nerve-impulse ; neuron; speed of nerve-impulses; degeneration ; regeneration 159-169 Sympathetic System : Fibres; communication with cerebro-spinal >y-trin ; iranylia ; pelvic plexuses ; function; vaso-motor fibres ; relation to secreting glands 169-173 CONTENTS. 7 PAGE Spinal Cord: Gross anatomy; nerves; white substance; gray matter ; nerve-roots ; trophic centres ; functions ; conduction ; motor fibres ; sensory fibres ; transference ; reflex action ; augmentation; automatic acts; coordination; inhibition. . . 174-185 Medulla Oblongata: Anatomy; columns; gray matter ; function; special centres 185-191 Pons, Crura Cerebri, Corpora Quadrigemina: Paralysis following lesions; functions 191,192 Cerebrum : Hemispheres ; fissures; convolutions; regions; lobes; gray matter ; chemistry of brain-tissue ; weight of brain ; course of fibres ; corpora striata ; optic thalami ; functions of cerebrum ; localization of brain -function ; motor areas ; sen- sory areas: paralysis 192-201 Cerebellum: Gray matter ; function 201-203 Tracts in Brain and Cord : Motor tract ; sensory tract ; lesions . 203, 204 Mutilations: Brain; cord; cerebellum; hemispheres 204-206 Cranial Nerves: III nerve; IV nerve; V nerve; VI nerve; VII nerve; IX nerve; X nerve; XI nerve ; XII nerve . . . 206-220 THE SEXSES : Sensations ; special ; common ; pain ; hallucina- tions ; perceptions; judgments 220-222 Touch : Organ ; varieties ; acuteness ; measure of ; pressure-sen- sation ; muscular sense ; temperature-sense 222-225 Taste : Tongue ; papillae ; taste-globlets ; association of smell ; after-taste . . . 225-229 Smell: Odors; olfactory nerves ; acuteness of; sneezing .... 229-232 Hearing : Auditory apparatus ; external ear ; middle ear ; inter- nal ear; membrana tympani ; ossicles; Eustachian tube; semicircular canals ; sense of equilibrium ; cochlea ; canalis cochlearis ; sacculi ; utricle ; organ of Corti ; auditory nerve ; course of sound-waves; localization of hearing; distance; subjective hearing ; musical range 232-241 Voice or Speech : Larynx ; production of voice ; articulate speech ; musical range of the voice . 241-243 8 CONTENTS. PAGE Sight: Visual apparatus; accessory organs of eye; eyelids; lachrymal gland; Meibomian glands; extrinsic muscles of the eye; eyeball; cornea; sclera; aqueous humor; crystalline lens; ciliary muscle; iris; vitreous humor; refraction; accommodation ; retina ; rods and cones ; area of most acute vision ; blind spot ; optic nerve ; optic tracts ; nervous mechan- ism of vision ; binocular vision ; stereoscope ; inversion of the image ; visual sensations and perceptions ; retinal red ; optograms ; achromatism ; after-images ; near-point ; emme- tropic eye; myopia; hypermetropia; presbyopia; astig- matism; diplopia; color-blindness; theories of normal color- perception ; causes of color-blindness ; in sexes ; test for color- blindness 243-265 EMBRYOLOGY: Reproduction; Species; heredity; methods of reproduction ; asexual ; sexual ; theory ; fecundation ; copula- tion ; female organs of generation ; ovaries ; Fallopian tubes ; uterus ; vagina ; ovulatiou ; menstruation ; puberty ; meno- pause; corpus luteum; male generative organs; testicles; spermatozoa ; seminal vesicles ; prostate gland ; penis ; urethra ; impregnation ; coitus ; details of impregnation 267-283 Development: Segmentation; germinal membrane; changes in uterine lining; blastoderm; medullary groove; changes in the mesoblast ; structures derived from : the epiblast, the mesoblast, the hypoblast ;sp lanchnopleure ; umbilical vesicle ; vitelline duct; amnion ; allantois ; chorion ; placenta; foetal and maternal blood ; umbilical cord ; foetal circulation ; vitel- line circulation; placental circulation ; area vasculosa ; forma- tion of heart ; change from foetal to adult circulation ; vertebral column ; cranium ; face ; extremities ; spinal cord ; nerves ; brain; vesicles; folding of brain axis ; eye; auditory appa- ratus ; olfactory apparatus ; alimentary canal ; salivary glands ; pancreas; liver; lungs; Wolffian duct; Miiller's duct ; testicle; ovary; Gartner's duct ; external genitals 283-306 Parturition: Expulsion of the foetus ; uterine contractions ; char- acter of uterine contractions 306, 307 APPENDIX : Table of development of an embryo ; chemical tests used commonly in physiological analysis; metric system . . . 309-311 PHYSIOLOGY. GENERAL CONSIDERATIONS. Physiology, from the Greek yuaiz, and /oyo^, literally means a discourse on nature. At present, however, the word has a more limited significance, meaning that branch of science which treats of the workings of the healthy living body. That portion of physiology discussed in the following pages is the physiology of the healthy living human being. From the very nature of things it is evident that every detail of physiology cannot be studied on the human subject, and the deficiency has been supplied by analogous study on the lower animals. In order properly to understand physiology, it is essential that the student be somewhat familiar with anatomy, or that branch of science which teaches the gross structure of the body. To appreciate the workings of a machine, at least a superficial knowledge of its component parts is necessary. Life : Our definition of physiology assumes the body to be alive ; but what life is, is a very difficult subject to explain. It may be described as the stimulus which keeps active the functions of the body, the characteristic phenomena exhibited by living beings. It is really indefinable ; for while it is readily possible to state the differences between living and dead things, it is impossible to define these differences. Living body compared to a machine : The living body may be compared to any piece of complex machinery, the parts being carefully adjusted to one another and performing their work as long as the proper stimulus is applied. When the stimulus fails and in the case of animals and plants the stimulus is called life the machine stops. 2 Phys. 17 18 CELLS. Still another comparison may be drawn between the body and a piece of machinery say, a steam engine. There is for both the need of fuel that can be oxidized by burning, and of resultant energy, liberated in the form of heat and visible motion. Also, as the parts arc subject to wear and tear, new parts must be supplied. In the case of a steam engine, the fuel is the coal fed to the boiler and oxidized by burning. In man the fuel is the food we eat, burned up in the presence of the oxygen we breathe. But here the analogy ceases, for whereas the coal supplied to a steam boiler never forms an integral part of the boiler, in man the food-stuffs are absorbed and go to form part of the individual himself before they are broken down by the oxidizing agent. Metabolism: The process of building up the food-stuffs, already properly digested, into a part of the body is called anabolism. The breaking down is the kataboiie process, or katabolism. Both together are called metabolism. To make the simile between man and a steam boiler perfect, we should have to imagine a boiler made of coal, instead of iron, which being ignited on its inner surface generated heat which caused the contained water to boil. And further, we should have to imagine that as fast as the inner layers of coal were consumed fresh coal was placed on the outside to make up for the loss. Thus we see that food is digested, absorbed, and incor- porated into the body, only to be broken down again by oxi- dation. This oxidation, or burning, is the source of the <- patho- logical. Thus fatty degeneration of the cells in the breast produces the oil-globules in the secretion of milk. The protoplasm of which all cells are made up has been called the basis of life. Protoplasm is an unstable albuminoid substance of more or less gelatinous consistency. Its reactions are those of albumin (coagulation by heat and mineral acids), and its chemical composition is in varying proportions of the elements C, H, O, N, and S. Protoplasm is living albumin, or proteid. Besides proteid, every cell contains water. The amoeba is a microscopic animal that consists of but one cell ; yet this one cell performs all the animal functions. FIG. 4. Protamoeba (Haeckel). (1) It is capable of motion (Fig. 4), the cell sending out a small branch or arm of the protoplasm. This branch, or pseudopod as it is called, attaches itself to some part ^ of the environment, and then the rest of the cell is drawn forward until the whole cell seems to flow to and into its branch. The process is then repeated, the cell advancing steadily forward. (2) It has the power of food-absorption ; the cell flowing around and completely enveloping any particle of food it may meet, Later on, such of the rood as has not been absorbed is excreted, by a process of flowing away, on the part of the protoplasm. EPITHELIUM. 23 (3) It also absorbs oxygen from the surrounding atmos- phere. (4) It is irritable to stimuli; and is capable of (5) repro- duction and (6) growth. Thus we see that the amoeba presents in its single cell all the main functions of the higher animals. But the individual cells of the human body are by no means so well endowed with functions as the amoaba. Differences between plants and animals : The most essential difference lies in the fact that vegetable protoplasm can build up new albuminous compounds out of such chemical bodies as water, carbonic acid gas, and inorganic mineral salts. Animal protoplasm, on the other hand, must have ready- formed albuminous food in order to live. Vegetables exhale oxygen, but inhale carbonic acid gas ; while animals do the reverse. Specialization of cells : As a rule the body-cells are highly specialized for the fulfilment of one or more functions, to the exclusion of all others. Where we find two or more cells of the same kind aggregated together, the mass is called a tissue. Two or more tissues of different kinds are spoken of as organs. By apparatus is meant an association of organs for the per- formance of a common function. Thus the stomach, intes- tines, and pancreas may be spoken of as a collection of organs, forming an apparatus for the common function of digestion. In ordinary parlance an organ is " the instrument of func- tion, the performer of a function." Classification of tissues: The cells in man are, roughly speaking, found to make four main classes of tissues : epithe- lial, connective-tissue, blood-cells, and nerve-cells. Epithelium : The name " epithelium " is given to the cells which cover the skin, and the mucous and serous membranes )f the body ; and which also enter into the formation of the ;lands. Its varieties are (1) simple, a layer of flat (squarn- >us), cubical (spheroidal), or cylindrical (columnar) cells, as in the serous and mucous surfaces ; (2) stratified, when it occurs in layers, as in the skin ; (3) transitional, where it has the characteristics of both in situations where the other two 24 CHEMISTRY OF THE BODY. FIG. 5. forms approach one another, as in the bladder. (4) In the "lands are (\n\i\d functional cell*, which partake of the charac- ter of the epithelium of the surface. They are arranged in groups about the ducts. Such cells are often known as secreting or glandular epithelial cells. Simple epithelium possesses hair-like processes in certain locations, and this is known as viliutctl epithelium (Fig. 5). The hairs are endowed with motion, and wave in such a manner as to throw for- ward small particles which fall upon them. The principal uses of epithelium are: protection, as skin, serous surfaces; motion, ciliated epithelium of air-pas- sages and Fallopian tubes ; secretion, in glands e. ODIES, 33 3 02 C A = -3 *> g tc OQ t-l , > . v~*-^ , * t^ Icj g 0) bio PQ B | d '~c t rr* ,GANIC PRINC g fll .2 I .si il % T3 3 I I 1 5 c ' B aS 'SSs 3sJ milk-cur< n i IMATE I ! "S. = "E i o> . C ^ - .si ^.Sc ^ ^^ 0) s'c'S oteolytic, Y NITROGEN M P4 O 1. Native albumins found in nature, 2. Globulins found in nature, 3. Derived albumins, derived from Class 1 by action of acids, alkalies, or ferments. 4. Fibrin, from Class 1 by ac- tion of " fibrin-ferment," 5. Coagulated albumin, by heat, etc., from Class 1, 6. Peptones, by action of di- gestive ferments on Class 1. f Gelatin, soluble in hot water. Mucin, soluble in weak alkali Elastin, insoluble. Chondrin, soluble in hot wate 1 Keratin, insoluble ; probably ' Crystallizable acids e. g., give Urea and its allies e. q\, kre'at [ Pigments e. g., bile-pigments, Ferments e. g., amylolytic, pi y S.2 ^ S si ^ci |i PROXIMATE PRINCIPLES. 29 ORGANIC BOD] 3 PRINCIPLES. j. ) (melts at 113 F. ^ ( " 140 F. j at closely allied to the - s itl l| ^'"SflS c ^- 3 *o S -rH ** "" 2 *lli'a > d _o ENOUS CIMATI o> J5 g ,Q "o J3 Or^H-u'CSdO S 1 ir g i| a perspira 3 5 IS s E-I 03 "3 ^5~ - .2 J3 .5, o O O 5 fl 0> o g ^3 5^ , - O g-S- - o" " - w o- M - c 1 e . OJ~ S || I H 1 I-H.2 S OQ 3 $& 5 ^-<3 30 THE BLOOD. and magnesium, the phosphates of calcium, and some soluble salts of iron, all of which exist in the earth ; also on the oxygen and carbon dioxide of the air. By various chemical combinations and changes these inorganic compounds are con- verted by the green plant under the influence of sunlight into simple organic substances. Non-chLorophyl-bearinfj plants, such as fungi, have not the property of absorbing inorganic substances and converting them into organic substances, but must receive their food in the form of organic compounds; hence the fungi live on other plants. So too with animals ; they must live on plants and other animals. From these plants and animals we cat we obtain, besides a number of inorganic salts, water, etc., the three great classes of food-stuffs which are eventually to be digested, altered, and assimilated into the body, then " burned " with the liberation of energy, and finally to be cast off as waste-products by means of the sweat, breath, urine, and faeces. These three classes of food-stuffs are protei^, c<>- hydrates, and fats. How these foods are absorbed and assim- ilated will be studied later in the chapter on Digestion. To be sources of energy, the tissues of the body must be oxidized ; this is accomplished by the oxygen conveyed from the lungs to the tissues by the blood. THE BLOOD. General observations : Since the earliest times it has been recognized that the blood is the most important fluid in the human body ; that it is, indeed, indispensable to life. Its loss prostrates and enfeebles the body. With excessive hemorrhage life itself ebbs away. This is readily evident when it is known that the blood, circulating through the sys- tem, carries to the tissues and the cells composing them material for their growth, renewal, and repair; and removes from them that which has become effete. It also tends to equalize more or less the temperature throughout the body. Gross appearance: Freshly drawn blood is a somesvhat viscid opaque fluid of a more or less pronounced red color. This color, which depends entirely on the amount of oxygen present in the blood, varies according as the blood is drawn PHYSICAL CHARACTERISTICS. 31 from the arterial or the venous system. That drawn from the arteries or pulmonary veins is of a bright scarlet, and contains a large amount of oxygen ; while that from the systemic veins is bluish-red to reddish- black, and has far less oxygen. The opacity is due to the fact that blood is not a homogeneous liquid, but consists, as we shall see later, of two elements corpuscles and plasma. These, differing in their refractive power, oppose the transmission of light, thus causing the opacity. FIG. 6. Human blood as seen on the warm stage. Magnified about 1000 diameters, c, c, crenated red corpuscles; p, a finely granular ; g, a coarsely granular pale cor- puscle ; both exhibit two or three vacuoles. In g & nucleus also is visible. r, r, single red corpuscles, flat; r', r', the same on edge; r", r", the same in rouleaux (Quain). Physical characteristics : The specific gravity of the blood at ordinary temperature varies between 1045 and 1062. It is of a slightly alkaline reaction, due to the phosphates and carbonates of the alkaline metals, and has a salty taste and a peculiar characteristic odor. The degree of alkalinity, esti- mated as Na,CO 3 , corresponds in human blood to 0.35 per cent, of this salt. 32 THE BLOOD. If examined under the microscope, the blood will be seen to consist of a fluid, called plasma or liquor sanguinis, in which are suspended small particles called blood-corpuscles. The corpuscles are of three types, known as red corpuscles, white corpuscles or leukocytes, and blood-plates or microcytcx. The temperature of the blood in man is, on the average, 98.9 F. ; but it is very probable that in certain parts of the body it is several degrees higher. Quantity of blood : The total amount of blood is estimated as being one- thirteenth of the body-weight. Thus in an in- dividual weighing one hundred and seventy pounds, the FIG. 7. Red globules of the blood, seen a little beyond the focus of the microscope (Dalton). amount of blood in his body would weigh about thirteen pounds and would measure about six quarts. The corpuscle* weigh about one-third of the total blood- weight. Corpuscles : The corpuscles (Fig. 6) exist in large numbers in the blood, it being estimated that in every cubic millimetre 1 of blood there are 10,000 leukocytes, 250,000 blood-plates, and 5,000,000 red corpuscles. Red corpuscles: Human colored blood-corpuscles (Figs. 7 and 8) are circular, biconcave disks with rounded edp- : in RED CORPUSCLES. 33 diameter they are about s^nj-th in. ; in thickness about in. In water they swell and become flat or convex. When seen singly they appear yellow, but their color is red when seen in groups. Microscopic examination shows that they have no nucleus and no limiting membrane ; but they have an elastic framework, or stroma, which retains an individ- FIG. Red globules of the blood, seen a little within the focus (Dalton). uality for each corpuscle, and allows changes of shape to adapt them for capillary circulation, and brings them back to the original form after such distortion. The color is due to a chemical body, called haemoglobin, which is held within the stroma. The red corpuscles have a specific gravity of 1088, and are the heaviest of the compo- nent parts of the blood. In all mammals, with the exception of the camel family, the c/encrdl character of the corpuscles is the same as in man ; but their size varies in the different animals (Fig. 9). The camel family have the same corpuscles as other mammals, except that the discs are oval instead of circular. In reptiles, fishes (except a few species), and birds the red corpuscles are oval, nucleated, and usually larger than those of mammals. 34 THE BLOOD. Red corpuscles origin : Like all the other cellular elements of the body, there must be a birth of new blood-cells to take Mainnjiils. Birds. Reptiles. Amphibia. the place of those whose life-work is done. The ancestors of the red blood-corpuscles are the large, irregular, polyn->- RED CORPUSCLES. 35 nal marrow-cells. These marrow-cells are found, as their name implies, in the marrow of the long bones. They are found in layers or phalanxes, each phalanx a little more highly developed than the layer underlying it. The most highly developed divide by karyokinesis, and are pushed oif from the mass of cells by the growth of the cells below. The detached cells, after undergoing certain changes in shape, enter the blood-current and become the ordinary red blood-corpus- cle. The marrow-cell is polygonal, nucleated, and only very slightly colored. To become a red blood-cell the marrow- cell loses its nucleus, assumes the biconcave disc-shape, and acquires a greater amount of coloring-matter, or haemoglobin. Such changes have been observed to take place in the mar- row-cells; and also in cases in which individuals have suf- fered severely from hemorrhage, during the recuperation, numbers of only partially transformed marrow-cells have been found circulating in the blood-current, not having had time to be fully altered, as the demand for new cells was so urgent. What becomes of the nuclei which the marrow-cells cast oif is not known. Suggestions have been made that these nuclei may become the blood-plates or else be destroyed by the leukocytes. These are merely suggestions, and not to be accepted as proven. Where the increase in coloring-matter which the marrow-cell acquires comes from, is also a source of speculation. Red corpuscles function: The one important and funda- mental purpose of the red blood-corpuscles is to convey o.i't/f/cii. from the lungs to the tissues. As has been said, the red corpuscles consist of an elastic network or stroma, which holds, probably both mechanically and chemically, a highly complex protcid body called haemoglobin, which, although cry stall izable, is non-diffusible. The form of the crystals of haemoglobin varies in different animals. In man they are prismatic, in the guinea-pig they are tetrahedral, in the squirrel they are hexagonal, and so on. Hamoglobin can be subdivided into two bodies globulin (96 per cent.) and hsematin (4 per cent.). Hcemin (Fig. 10), the hydrochlorate of haematin, is formed 36 THE BLOOD. by adding muriatic acid to haemoglobin. It is of especial iui. rest from u medico-legal point of view, as it can be obtained, by proper manipulation, from a very minute '(//// coat: If we examine under the microscope the blood-clot which has formed under ordinary circumstances, it is seen that the red corpuscles arc caught by the meshes of fibrin and are uniformly distributed through- BL OD-COA G UL UM. 45 out the clot. Many of the leukocytes have made good their escape into the serum. If we take a quantity of blood (Figs. 15 and 16) and retard the time of coagulation (horse's blood coagulates more slowly than human blood), we note that the upper part of the clot is much lighter in color. This is called the " buify coat." The explanation of its formation is very simple. Owing to retardation of the coagulation, the red cor- puscles have had a chance to settle toward the bottom. When seen under the microscope the upper layer, or buify coat, of the clot is found almost free from red corpuscles. FIG. 15. FIG. 16. Vertical section of a recent coagulum, showing the greater accumulation of blood-globules at the bottom (Dalton). Bowl of coagulated blood, showing the clot buffed and cupped. Why blood does not clot in the blood-vessels : As long as the endothelial lining of the blood-vessels is intact the blood does not clot ; but if the endothelium is injured, a clot is formed at the site of the injury. Just what check on coagulation the endothelium plays is unknown, but it is none the less positive. If a vein of some animal, preferably a horse, be carefully ligated in two places some inches apart, the blood within it will not coagulate for a long time, provided the endothelium is uninjured. This is not due to occlusion from the air sim- ply. If such ligated vein with its contained blood be re- moved from the body and carefully opened, the blood may be poured from it into another vein similarly prepared, just as we pour fluid from one test-tube to another. Though the blood is thus exposed to the air, it will not coagulate till the endothelium is injured or loses its vitality. -10 THE LYMPH. If one accepts the theory that the fibrin-ferment is the re- sult of disintegration of leukocytes, it is clear why Mood does not clot within the bloodvc^els, lor then- is no ferment unless a leukocyte disintegrates, the " normal " destruction of leuko- oytes being disregarded as a possible factor, owing to the small Dumber so destroyed at any given time. On the other hand, if the endothelium be wounded, the leukocytes rush to the breach and are broken up; also, if the blood is shed, many leukocytes disintegrate in mass; thus in both instances a large amount of fibrin-ferment is liberated and clotting begins. Blood resume of its uses : The red corpuscles convey oxygen from the lungs to the tissues. The platnna conveys the food, after absorption from the alimentary canal, to the tissues. The pla.-ma also holds in solution the carbon dioxide and water (the result of oxidation) which it receives from the tissues, and carries these products to be eliminated by the skin, lungs, and kidneys. The plasma also holds in solution the urea and other nitrogenous bodies that are carried to and excreted by the liver and kidneys. The leukocytes are protectors against the invasion of bac- teria. Thus it is seen that the various factors of the blood make up a whole that is, a conveyor of supplies, a remover of waste, and a protector from invasion. THE LYMPH. Lymph is a pale straw-colored fluid that bathes all the tiny tissue-spaces of the body and is conveyed by lymph-vessels from the lymph-spaces to the blood. It is slightly alkaline, <>f a salty taste, and has no odor. Composition : Lymph may be compared to a dilute blood free from r<-d corpuscles, but rich in leukocytes. "When the lymph-vessels of the lower animals, such MS tadpole-, are ex- amined under the microscope, it is found that the leukocytes are only present in the larger Trunks; the smallest channels rarely contain particles >r corpuscles of any kind. It is not THE LYMPH. 47 meant that all the constituents of blood are equally reduced when lymph is spoken of as a dilute form of blood. About the same proportions of salts, urea, and sugar are present in blood and lymph. Only about one-half as much serum- albumin and paraglobulin, and about one-quarter as much fibrinogen, are in lymph as are present in blood. Dnrin(/ digestion there is a marked increase in fats and at such times the appearance of the lymph from the intestines is like milk, instead of being straw-colored. This form of lymph, known as chyle, owes its opacity to the distribution of this fatty matter in innumerable particles of very minute though nearly uniform size, measuring about -g-o-jyo^oth ^ an i ncn Each particle is coated over with albumin. Lymph coagulates in the same manner as blood, but more slowly and more feebly ; a difference readily explained by the marked difference in amount of fibrinogen present. This property of coagulation is not possessed by the lymph in the smallest lymph-spaces nor by the chyle in the villi or lacteals near them. It is only developed as the fluids near the tho- racic duct. Lymph sources: Between the individual capillaries of the blood-system there exist small intercellular spaces, into which the fluid portions of the blood are exuded owing to the differences in pressure. This exudate, plus the leukocytes that have left the bloodvessels by emigration, makes up the lymph proper. In addition to the lymph proper, the lymph that fills the lacteals of the intestinal villi absorbs some of the products of digestion, especially the fats. This portion of the lymph that has absorbed the fats is milky in appear- ance, and is called " chyle.' 7 The chyle and lymph proper are carried along in their respective channels and are min- gled together just before the entire fluid is poured into the vein at the root of the neck. Uses: The lymph bathes all portions of the body not reached by the blood. Hence the lymph conveys the nu- trient ingredients of the blood to all cells not directly bathed by the blood. The " chyle/' or lymph of digestion, absorbs nutrient materials (mostly fat) from the intestines and pours this food into the blood-current, to be universally distributed. 48 CIRCULATION OF THK JlLOOh. The lymph gathers up the waste-materials of the cells sur- rounding the lymph-spaces and pours this waste into the blood, to be eliminated by the skin, lungs, and urine. CIRCULATION OF THE BLOOD. The circulation of the blood is the course which the blood, as a transporting medium, follows in taking food and air to the tissues and bringing away the used-up material for ex- cretion, and returning when freshly charged with oxygen and food. FIG. 17. J. V. Subcl. artery. Lungs. Lungs. Heart and lungs of man (Milne Edwards), r. a., right auricle; v., vena mv.i in- ferior; a., right ventricle ; r., aorta ; I. v., left ventricle ; j. v., jugular vein ; c. a., trachea and carotid artery. The circulatory apparatus consists of: (1) The heart, which propels the blood ; (2) The arteries, which convey it from the heart to the different parts of the body ; COURSE OF THE BLOOD. 49 (3) The capillaries, a network of inosculating tubules interwoven with the substance of the tis- sues and bringing the blood into intimate con- tact with it ; (4) The veins, which collect the blood from the capillaries and return it to the heart. Course of the blood : As the motion of the blood is in a circle, it is imma- terial at what part of the vascular system we begin its study. Starting, for convenience, with the blood as it leaves' the lungs, we find that the blood enters into the left auricle from the pulmon- ary veins, thence passing the open mitral valve into the left ventricle (Figs. 17 and 18). Upon con- traction of the ventricle the mitral valve is closed and the aortic valves thrown open, so that the blood is thrown into the aorta, and thence through the systemic arterial cir- culation into capillaries and on into veins, the systemic veins finally joining to fill the venae cavre, and from them the right auricle. From the 4 Phys, 50 CIRCULATION OF THE BLOOD. right auricle the blood passes the tricuspid valve into the right ventricle, whence it is thrown through the pulmonary artery (guarded by the .pulmonary semilunar valves) into the pulmonary capillaries, and thence into the pulmonary veins, whence it started. Thus we have in reality two circulations, the systemic and the pulmonary. Also we see that the blood goes through one set of capil- laries the lungs to be purified of waste-products and re- FIG. 19. Anterior view of heart (Quain). oxygenated ; and another set the systemic capillaries where the oxygen is lost and the waste-products gathered up. Furthermore, the student will take note that the blood which enters the splenic and intestinal arteries from the aorta passes through fim ,sv7x of capillaries before returning to the general venous circulation: the Jirxf *f' the heart it fills up with blood from the veins which has already travelled the circuit. The heart is the prime and important factor in maintaining circulation, and is capable of forcing, unaided, the blood through the entire circuit ; but in addition we find suh^'ulitirii forces assisting the outward flow of blood : (1) Arterial resiliency : At each beat the ventricles inject a fresh quantity of blood into the already filled arteries. To make room for this surcharge of blood the large arteries, owing to their elasticity, are dilated by the increased amount of blood. During the repose of the heart the arteries con- tract by virtue of the recoil of their elastic fibres and drive the blood on. This action of the arteries converts the inter- mittent character of the blood-stream into a steady flow by the time the capillaries are reached. (2) Contraction of muscles: The skeletal muscles, during the ordinary activities of the body, contract, and so compress the veins lying between them. This compression drives the blood out of the veins, but only in the proper direction, as the valves of the veins prevent the reverse flow. (3) Cardiac suction : During the period of repose the open- ing out of the heart-cavity is sufficiently strong to exert a decided suction on the blood in the veins. (4) Thoracic suction: During the expansion of the chest in inspiration a tendency to a vacuum is produced, which ex- erts a suction on the large veins within the thoracic cavity. This fact must always be borne in mind by the surgeon BLOOD-PRESSURE. 57 when operating on the neck. Should a large vein be wounded, it must be instantly closed, lest by reason of this suction air be drawn into the vein. If air should in this way enter the proximal part of a wounded vein, it would be hur- ried along to the heart, and, there expanding, cause serious trouble or even death. (5) A slight rhythmical contraction of the veins. Speed of the blood : In the arteries it has been estimated that the blood in the large arteries travels at the rate of about 300 to 500 mm. per second. As the blood advances in the arteries the speed decreases gradually. In the capillaries, owing to their much larger aggregate diameter (500 : 1), the speed falls very low, and is estimated to be f of a mm. per second. In the veins the speed increases as we approach the heart, and in the venae cavse is found to be from 125 to 200 mm. per second. With these figures as a basis, it has been calculated that a given particle of blood occupies about a half minute of time in going the round of the body. One second of the half min- ute is used in passing the systemic capillaries, another second in traversing the pulmonary capillaries. It has been found by experiment on animals that there is a ratio between the blood-speed and -pressure which may be direct or inverse. Any influence which increases or dimin- ishes the force of the heart will at the same time increase or diminish both the velocity and the pressure of the blood ; while any influence which increases or diminishes the re- sistance to the arterial flow will make the velocity and the blood-pressure vary in an inverse ratio to each other. Other things being equal, both the pressure and the velocity in the large arteries increase markedly during systole and diminish greatly during diastole. Very near the heart the velocity is greatest at the beginning of systole, then drops to almost nothing toward the end of systole, and then, regaining speed, flows at an almost even pace during diastole (Fig. 25). Blood-pressure : By means of manometers the blood-pressure has been determined. It is greatest in the arteries nearest the heart, and gradually diminishes, until in the veins empty- 58 CIRCULATION OF THE BLOOD. ing into the heart it is found to be zero ; or even a slight negative pressure or suction may be present. The figures are as follows : pressure in the large arteries supports a column of mercury 150200 mm. Iiiirh (about four pounds to the square inch). In the capillaries, 3050 mm. of mercury. In the distal veins, 20 mm. of mercury, gradu- ally falling as we approach the heart. 1 234 1 23 4 1 234 Tracings of variations of rapidity and of pressure of blood in the carotid of a horse, obtained by Chauveau and Lortet. The line v represents tin- curve of the ra- pidity of the blood ; and p the curve of arterial pressure. The h'gun < aixl vi -r- tical lines represent corresponding periods in the tracings (McKnidrirk i. The cause of the high pressure in the arteries is due to the force of the blood injected from the heart behind, and the re- sistance of the capillaries in front. Although the blood is pumped into the arteries in intermittent jets, when a *//>"// artery is cut the blood spurts from it in a continuous jet. The reason for this is the elasticity of the arterial walls, these 1 being put on the stretch by the blood forced into them at con- siderable pressure, and contracting again during the period of heart-rest, thus enabling the arteries to keep the blood under elastic compression. The capillaries, though collectively of much greater area than the arteries, by reason of the friction they offer to the blood-stream maintain a less degree of ten- sion. THE BLOOD IN CIRCULATION. 59 Conditions modifying arterial tension : (1) The rate of the heart-beats, by keeping the arteries fuller or less full, will modify the blood-pressure in the arteries. (2) Vaso-motor changes, by increasing or decreasing the friction offered the arterial blood, vary the tension in the arteries. (3) The amount of blood in the system must to a great extent determine the limits of arterial pressure. In great exsanguination the arterial pressure is quite low. (4) Motion of the thoracic walls in breathing necessarily changes the arterial tension by the pumping force exerted by this motion. The blood in circulation : While circulating in the arteries and veins the blood travels at too great a speed to admit of careful examination ; the leukocytes in spherical form and the red corpuscles, bowled along indiscriminately, pass the eye too rapidly for any peculiarities to be noticed. The capillaries of the web of a frog's foot can be easily examined with a microscope. In these the speed of the blood is slow, and ample time is given to study the particular cor- puscles. On entering the capillaries the red cells are seen to occupy the central portion of the stream of plasma. That portion of the plasma in contact with the capillary walls is seen to travel much slower than the central portion, owing to the friction presented by the capillaries. This is known as the " inert layer " of plasma. The corpuscles, being compara- tively heavy, are crowded in the centre of the stream. That the centre of the stream travels faster than the periphery is shown by the fact that if we watch three corpuscles floating abreast, in a moment or two the centre corpuscle forges ahead and leaves its companions behind. Some of the capillaries are so small that the red corpuscles pass in single file ; some may even be so small that the cor- puscle is squeezed up and elongated that it may glide through the constriction ; on entering a vessel of larger calibre the corpuscle, owing to its elasticity, resumes its shape. The leukocytes in the arteries are carried along passively in spherical shape, but on entering the capillaries are crowded 60 CIRCULATION OF THE BLOOD. into the inert layer by the heavier red corpuscles. In the inert layer they change their shape and flatten out against the capillary-wall. They now exhibit their amh<;win^ tin- direction in \\hiHi impressions are conveyed. The ellipse :it tin- upper extremity of the vat-us, looking like a section of the nerre, u intended to represent the va^al nucleus or centre. In this diagram the nerves are incorrectly made to cross, instead of passing behind, the m.rta. pumj^ed with each ventricular contraction is six ounces, in an ordinary man an amount of blood equal to the total blood INNERVATION OF THE HEART. 65 of the body will pass through the heart in about half a minute. The amount of blood poured out by the systole of the ven- tricles is called the " pulse-volume." Innervation of the heart : This matter is somewhat unde- cided at present, for the reason that many of the results must be obtained from experiments upon the hearts of cold- blooded animals. We do know that the mechanism of rhyth- mical contraction is contained within the heart itself. Nerve- ganglia are demonstrated in the frog's heart which are essen- tial to its action ; similar ganglia exist in the human heart. These ganglia are connected with fibres from the pneumo- gastric (or vagus) nerve and with the sympathetic system (Fig. 29). The eardiac inhibitory nerve is a branch from the pneumo- gastric nerve running to the heart. It has an inhibitory or slowing effect upon the heart ; for if we cut the nerve the heart becomes more rapid, and if we stimulate (Fig. 30) the FIG. 30. Effect of stimulation of the pneumogastric nerve upon the action of the heart in a frog. To be read from right to left. peripheral end of the nerve we slow the heart again. This action may be traced to the medulla oblongata, where a cardio- inhibitory centre is located. That the inhibitory influence of the cardiac fibres of the vagus is not directly exerted upon the heart is proven by the length of time elapsing between the application of the stimulus and the appearance of the in- hibitory effect. In some cases even two entire heart-beats occur after a strong stimulus is applied before the heart stops, indicating that some resistance must be overcome, the inhib- itory fibres acting upon inhibitory centres in the heart itself. Stimulation of the vagus not only slows the heart's action, but modifies it. Systole and diastole are lengthened ; the input and output of the ventricle are diminished ; the dias- tole pressure and volume of blood in the ventricles are in- 5 Phys. 66 CIRCULATION OF THE BLOOD. i 1* 8 a II creased, and ventricular become less frequent than auricular ((niti-actions, the latter being often twice as numerous. ( 'crtaiu fibres of the sympathetic from the cervical and up- per dorsal spinal cord pass to the heart. If these fibres are left after all other nerve-connections of the heart are cut away, stimulation of the spinal cord will cause the heart to become rapid. These are known as accelerator nerves. These accelerating nerves act less powerfully than the inhibitory ones. They not only accelerate the heart's action, but in- crease the force of the beat and the out- put (Fig. 31). The cardiac depressor nerve is a cen- tripetal nerve running from the heart to the vaso-motor centre in the medulla. Its purpose is to stimulate the vas- motor centre to dilate the peripheral arteries. The nerve is "stimulating" only when the heart is laboring against too high tension produced by unusually high peripheral resistance. Vaso-motor Nerves. Arterial contraction and dilatation : The muscular coats of the peripheral arteries do not undergo a rhythmical contraction and dilatation, but only con- tract so as to diminish the calibre of the arteries and thus limit the supply of blood to a given part of the body. On the other hand, the muscle-fibres relax and allow the arteries to dilate when the given part demands more blood to satisfy its activities. Hence it is a per- fectly normal condition to have the arteries of one part of the body dilated, while the arteries supplying another part ACTION OF VASO-MOTOR NERVES. 67 are contracted, depending on which part of the body is func- tionating. By tone of the arteries is meant their average normal state of contraction. The nerve-supply of the muscular coats of the arteries is through the " vaso-motor " nerves. They are of two varie- ties according to their function, vaso-constrictor and vaso- dilator. These nerves (vaso-constrictor and vaso-dilator) run together in the same sheath, and are part of the sympathetic system. They have their origin in the vaso-motor centre in the medulla. They pass down the spinal cord to emerge at dif- ferent levels, thence to be distributed to all the different ar- teries. Just before leaving the spinal cord there are second- ary vaso-motor centres in the course of these nerves. Close to the wall of the artery to be supplied there is a third (gan- glionic) centre for vaso-motor action. On emerging from this third centre the vaso-motor nerve breaks up into its terminal filaments which are distributed to each muscle-fibre. Action of vaso-motor nerves : It must not be supposed that contraction and dilatation of the muscle-fibres are active acts on their part depending upon whether the vaso-con- strictor or the vaso-dilator fibres are stimulated. It is true that contraction of the muscle-fibre is an active act depend- ing on stimulation from the vaso-constrictors ; but were this stimulus removed the muscle-fibres would relax and the artery dilate passively, owing to the presence of blood within it. We find that the vaso-constrictors act continuously with more 0r less intensity. The action of the vaso-dilators is only occasionally called into play, when marked dilatation of the bloodvessels is called for. Also the dilators do not directly act upon the muscle-fibres, causing them to dilate ; but act upon and inhibit the action of the vaso-constrictors, thus allowing a passive dilatation of the arteries because the constrictor action is removed. The vaso-motor centre in the medulla is the prime centre, and the only one acting during normal life. The secondary centres in the cord only assume control if the primary centre is destroyed or its connection severed. The third centres 68 CIRCULATION OF THE BLOOD. (close to the artery) only act if the secondary centres are de- stroyed or cut off. Pulse. The pulse is the alternate expansion and contraction of the artery ("excursion of the wall of the artery ") resulting from the fresh injection of blood at each ventricular systole. Of course, the pulse-rate is the same as the number of heart- beats. A number of various terms are used to describe the char- acter of the pulse : Frequent or infrequent^=uumber of beats per minute ; Regular or irregular whether beats follow successively or not; Intermittent beats skipped at regular intervals ; Large or srnall=amplitude of excursion of wall of artery ; Quick or slow^whether wall of artery rises rapidly or not. , ir . ( are synonymous terms ^ f Ir - v ' denoting a high-ten- S aseous > ... Incompr esS ,ble, i g . & S Their "P"*Mi Hard, soft. ^ opposites are J The sphygmograph : While an experienced physician can appreciate slight variations in the character of the pulse, it FIG. 32. Marey's sphygniograph applied to the arm (Marey). is only by means of the graphic method that the different kinds of pulse can be successfully investigated and records kept. The sphygmograph (Fig. 32) is an instrument which VENOUS PULSE. 69 measures the succession of the alternate dilatation and con- traction of an artery, known as the pulse ; and, magnifying these movements, registers them on a surface moving at a uniform rate by clockwork. The resultant tracings are in the form of a wavy line, the irregularities of which show variations of the pulse too slight to be appreciated by the most experienced fingers (Figs. 33 and 34). FIG. 33. Trace of the radial pulse, taken by the sphygmograph. FIG. 34. Dicrotic pulse of typhoid fever (Marey). Extinction of pulse : In health the pulse gradually lessens as we get further from the heart, and is entirely lost where the arteries break up into capillaries. There is no pulse in the capillaries nor a true pulse in the veins. Venous pulse : Were the venous walls rigid, there would be a respiratory pulse in the veins. In deep and infrequent respi- ration, as in partial asphyxiation, the jugular veins may be seen to be distended in expiration and collapsed during inspiration, thus forming an abnormal venous pulse. The cause is the pressure of the atmosphere upon the flaccid vein, it being easier to empty the vein than to fill it under the sudden vacuum produced by the act of inspiration drawing the blood from the surface veins into the thorax. The veins are thus emptied, and, being flaccid, are depressed by atmospheric pressure. This suction power of the inspiratory vacuum in the thorax extends over a small region only ; roughly speaking, the neck and axilla. This is the so-called " dangerous surgical region ;" for if a vein is punctured, air enters the blood-current and is carried to the heart, and death results. But the flaccidity 70 "CIRCULATION" OF THE LYMPH. of the vein naturally causes a collapse of the wall, and air will not rea< lily enter unless the wall remains ri^-id from arti- ficial or abnormal causes. If the vein can collapse, not being held open by connective tissue or fascia, or artificially or as a result of calcareous infiltration, no air will be allowed to enter the vessel, and the only sign will be an expiratory inter- mittent hemorrhage. "CIRCULATION" OF THE LYMPH. The lymphatics : The blood-capillaries as they pass through the various tissues of the body are surrounded by small open spaces, called lymph-spaces. These spaces are the interstices between the various component parts of the body-tissues. The lymph-spaces form the open mouths of microscopic lymph-capillaries, that closely resemble the blood-capillaries in structure. The lymph-capillaries run together and go to form larger channels, known as the "lymphatics n r " lym- phatic vessels"; this arrangement is comparable to the for- mation of veins from blood-capillaries (Fig. 35). These lym- phatic vessels converge finally into two main channels- thoracic duct and right lymphatic duct that pour their con- tents into the venous system at the junction of the internal jugular and subclavian veins on the left and right sides of the neck respectively. The lymphatic radicles (Fig. 36) of the intestine, although not differing in structure from lymphatics elsewhere in the body, are called " lacteals," because of the milk-like appear- ance of the lymph they contain. The lymphatics of the lower extremities, the trunk of the body, and the intestinal tract, go to form a long channel, the thoracic duct, which ascends along the front of the vertebral column 1o the root of the neck on the left side. Here the thoracic duct is joined by lymphatics draining the left side of the head and left upper extremity. The combined channel empties into the left innominate vein at the point of junction of the left internal jugular and left snbelaviau vein. The lymphatics of the right side of the head and right upper extremity join to form the rif/lit lymphatic duct a .-hort STOMATA AND PSEUDOSTOMATA. 11 structure, one and a half inches long, emptying into the right innominate vein. Stomata and pseudostomata : In certain parts of the body FIG. 35. Fro. 36. Lacteals and lymphatics, during digestion. Diagrammatic representation of the origin of the lacteals in a villus. e, central lac- teal ; d, lymph -channels; c, columnar epithelial cells, the attached extremities of which are directly continu- ous with the lymph-chan- nels. there have been found openings, or stomata, by which a direct communi- cation exists between the lymphatic capillaries and certain cavities previously supposed to be entirely closed. We say that absorption takes place through pseudostomata when fluids pass into the lymphatic system through the intercellular cement- substance of the epithe- lium or endothelium covering membranes. Stomata have been found in the peritoneum and pleura. 72 "CIRCULATION" OF THE LYMPH. Flow of lymph : Thus we see that the lymph flows from the tissue-spaces into the veins at the root of FIG. 37. t j ie 1R , e k . 80 it is not justifiable to speak of the " circulation of the lymph," as there is no return-current. The lymph is formed in the lymph-radicles, and ceases to be lymph proper aV soon as it enters the venous blood. But new lymph is constantly formed, and so there is a steady onward flow. Structure of lymphatics : The lymphatic channels closely resemble the veins in struct- ure, but are much thinner walled and pos- sess a much larger number of valves (Fig. 37). Sources of lymph: The lymph present in the lymph-spaces is made up of the exudation of a certain amount of blood-plasma through the walls of the capillaries, together with the leukocytes that have emigrated from the capil- laries. It is probable also that certain por- tions of the body-tissues may be found in the lymph, which, though they have fulfilled their functions and have to be removed, are not entirely waste-products. They may be capa- ble of reorganization in the lymph-glands ;:nd V phatics f (sSpplyr ma y be absorbed by the lymphatics for this purpose. The lymph in the radicles of the intestines is composed of the blood-plasma and leukocytes that have left the intestinal capillaries, plus an innumerable number of microscopic globules of fat that have been absorbed from the digesting food. These particles of fat render the lymph milky in appearance, hence the name "lacteals" for the intestinal lymphatic radicles. This milky fluid is called clii//c. The lymph proper and the chyle mingle in the thoracic duet to be poured into the venous system. I.i/nifili IH'<>I>t raw-colored fluid ; after the ad- mixture of chyle it is milky. Pressure of lymph : The blood in the capillaries has been LYMPH-GANGLIA. 73 stated as being under a pressure equal to 2050 mm. of mer- cury. In exuding through the capillary- walls the blood loses about one-half its pressure ; hence we find that the lymph in the tissue-spaces is under a pressure of 1025 mm. of mer- cury. At the veins of the neck the lymph is under very low pressure, or is at zero, or even under a negative pressure due to the suction of the chest. Factors producing lymph-flow; In some of the lower ani- mals there is a distinct separate lymph-heart, the purpose of which is to act as a force-pump to drive on the lymph, just as the human heart drives on the blood. In man there is no such lymph-heart, but the flow of lymph depends on other causes. They are as follows : (1) The positive pressure (1025 mm. of mercury) in the lymph-spaces, as opposed to the zero or negative pressure at the other end of the lymph-channel, drives the lymph on. (2) The muscular movements of the body compressing the lymphatics force the lymph on in the proper direction, the reverse flow being prevented by the valves. The chyle is aided in its flow by the actions of the muscular fibres of the small intestine, and possibly by the layer of unstriped muscle-fibre found in each intestinal villus. In the small in- testine of the mouse the chyle has been seen to flow with intermittent movements, corresponding to the peristaltic movements. The contractility of the walls of the lymph- vessels themselves, due to the muscle-fibres they contain, probably supplies some of the force. (3) The thoracic aspiration of the chest on inspiration sucks on the lymph in the same manner as it does the venous blood. Lymph-ganglia : The lymph-ganglia, or lymph-nodes, are innumerable small masses, varying in size from 1 mm. to 2 cm. in diameter, interposed in the course of the lymphatics. They are found in great numbers in -the neck, thorax, axilla, groin, and mesentery, and along the great vessels of the ab- domen. A few are found in the popliteal space and in the arm as far as the elbow, but none further down the leg or forearm. The lymph-ganglia consist of a mass of cellular pulp-sub- 74 "CIRCULATION" OF THE LYMPH. Fro. 38. stance, through which run many open channels, the whole being enclosed in a capsule. If a particular lymph- vessel be examined, it will be seen to empty directly at some point into a lymph-ganglion. The lymph flows through the open channels in the substance of the ganglion and emerges on the opposite side, to be again collected into a lymph-vessel, and so on to its destination (Fig. 38). A lymph-ganglion is com- parable to a sponge placed in a snug-fitting rubber bag. The cap- ics, Sim nle lymphatic gland, a, the c suk', with sections of lymph;it rtioii of the gland, in the centre of which the section of a bloodvessel may be seen. The path pursued by the lymph through the medullary portion constitutes the deep or sec- ondary lymph-path of Frey (Car- penter). rubber bag connects on one side with a rubber tube repre- senting the afferent lymph- vessel, and from the opposite side there leads away from the rubber bag another tube repre sc 11 ting the efferent lymph- vessel. The rubber bag is the representative of the capsule of the ganglion, the meshwork of the sponge is comparable to the framework of the ganglion, ;s ml the holes in the sponge to the open channels. The sub- stance of a lymphatic ganglion is adenoid tissue. Purpose of lymphatic ganglia : The lymphatic ganglia serve as filters for the lymph. Also the cell-division of leukocytes occurs in their channels. An important attribute is the way iu which they serve to retard the spread of infection through the body, thus acting as safety-valves. If any portion of the body is infected, the poison is carried by the lymphatics to their especial glands. There its course is stopped till the opposition of the glands is overcome, when it proceeds to the next ones. As a result of this we uet at first en- largement of the glands ; later, if the process continues, they lireak down and are destroyed. If it were not for their action in this way, an infected wound might often MOVEMENTS OF RESPIRATION. 75 prove rapidly fatal before surgical treatment had time to be of value. EESPIRATION. By the respiratory act we mean the process by which oxy- gen is introduced into the system and by which carbonic ox- ide is excreted. This function is performed by the lungs, and the transfer is effected by the agency of the blood. The respiratory act is subdivided into (1) the "movements of respiration," by which term is meant those movements that cause the thorax alternately to expand and collapse, thus fill- ing and emptying the lungs of air ; and (2) " respiration " used to designate the interchange of oxygen and carbon dioxide between the blood and the air in the lungs (external respiration) ; also the exchange of oxygen and carbon dioxide between the blood and the tissues of the body (internal respi- ration). Movements of respiration : The action by which the lungs are filled with air (inspiration) is as follows : The thoracic cavity is expanded by muscular action ; this expansion draws on the walls of the lungs through the intervention of the pleurae, thus enlarging the cavity of the lungs. To prevent a vacuum in the lungs, air is taken through the nose or mouth, and passes through the pharynx to the larynx ; entering the rima glottidis, it passes through the larynx to the trachea and bronchi. The air is somewhat warmed and moistened in its passage. The trachea and bronchi are lined with ciliated epithelium, which serves to sweep particles of dust and the like out of the air-passages. After the lungs have been filled with air the thoracic wall is made to collapse and the air is expelled from the lungs. The respiratory tract : In all vertebrates and in many in- vertebrates certain specialized structures, either lungs or gills, are specially adapted for allowing the blood to come into close proximity with the air or water which contains air. A lung or gill in its simplest form consists of a thin membrane, on one side of which is a network of thin-walled bloodvessels, while the other side is in contact with the air or water. There is a difference of degree only between the most com- 76 RESPIRATION. plex and the simplest forms. In man the respiratory tract is highly complex, and includes essentially the larynx, trachea, bronchi, and lungs (Fig. 39). FIG. 39. Human larynx, trachea, bronchi, and lungs; showing the ramification of the bronchi and the division of the lungs into lobules (Dalton). The larynx : The air enters the body through the mouth or nares. It first passes through the larynx, the upper part of tin- respiratory passage. This is a short tube, triangular on section, formed of hyaline cartilage, which contains the voral cords. At its lower extremity the larynx joins the trachea. The trachea : The trachea, or windpipe, is a hollow tube, four to four and a half inches long, composed of fibro-elas- MINUTE ANATOMY OF THE LUNG. 77 tic membrane, in which are enclosed a series of from sixteen to twenty cartilaginous rings. These latter serve to stiffen it. At its lower extremity it divides into two bronchi, one for each lung, named respectively the right and the left. The bronchi : These are merely smaller editions of the trachea and of a similar structure. They divide, enter the sub- stance of the lungs, and, subdividing, penetrate every part till they end in the lobules. As they get smaller the walls become thinner and the cartilaginous rings disappear. The lungs : These occupy the larger part of the thorax. They are of a spongy, elastic texture, and on section appear to be solid organs for the greater part. In reality they are hollow. The lungs are two in number, the right and the left, of which the former is partially subdivided into three lobes ; the latter into two. Covering each lung separately is a serous membrane, the pleura, which in the form of a sac envelops the outer sur- FIG. 40. Diagrammatic view of the pleural sacs with the heart and lungs interposed. A, in- ternal mammary artery ; N, phrenic nerve. face of the lung and the inner surface of the chest-wall. The pericardium containing the heart is interposed between the two lungs (Fig. 40). Minute anatomy of the lung : Each lobe of the lung is 78 RESPIRATION. composed of numerous lobules, into each of which a small bronchiole enters, and the minute terminal branches of these bronchioles (infimdibula] widen into a .sort of irregular funnel having pouched or sacculated dilatations, known as air-cc//*. These air-cells are supported by numerous elastic fibres, and are lined with a very thin layer of flat (not ciliated) epi- thelium. Outside the epithelial lining is a very close mesh of capillaries, which are often exposed to air on both sides In- lying in a partition between two of the air-cells. The air- cells or vesicles are about one-sixtieth of an inch in diameter, and the space between the capillaries is often less than the diameter of a capillary (Figs. 41 and 42). The total surface in the lungs bathed by the air is about two hundred square metres. FK;. 41. FIG. 42. Single lohule of human lung, a, ulti- Network of capillary blood-vossi-ls in the mate bronchial tube; h, cavity of pulmonary vesicles .f thr hone, n :>bule; c,, c, c, pulmonary cells, or cavity of vesicle, with capillary plexus : vesicles (Dalton). ft, pulmonary blood-vesaels, supplying capillary plexus (Frey). Muscles of inspiration: During <|iiiet inspiration the cavity of the chest is increased in all three diameters vertical, anteri-jM)>trri.r, and lateral. The upper part of the thoracic ba-Ui-t being fixed, the vertical diameter is increased by the d'-mit of the diaphragm in contracting ( Fig. 4:5). Th<> ex- ternal intercostals, together with other muscles, act on the MUSCLES OF INSPIRATION, 79 ribs so that they turn on their axes, with the result that the sternal ends are raised up and carried forward ; at the same time the peculiar curve that exists in the rib at its angle FIG. 43. TT Si A Diagrammatic sections of the body in inspiration and expiration. A, inspiration; B, expiration ; Tr, trachea ; St, sternum ; D, diaphragm ; Ab, abdominal walls. The shading roughly indicates the stationary air (Huxley). causes an eversion of the lateral aspect of the rib when the sternal end is brought upward and forward. This eversion increases the lateral diameter. The axis of a rib is a line running through the centre of the head (vertebral end) of the rib and through the articula- tion of the rib with the transverse process of the vertebra. The muscles producing these changes in the diameters of the chest are the muscles of " quiet inspiration/' and are as follows : Diaphragm increases the vertical diameter 80 RESPIRATION. Scaleni fix upper part of the thorax ; External intercostals ] and > draw the ribs upward ; Interchondrals J C fix the second rib and raise Serrati postici stiperiores< the third, fourth, and fifth ( ribs; Levatores costarum breves raise the upper ten ribs. It is stated that of the two types of respiration, " thoracic " and " abdominal/' the former is more marked in women and the latter in men. This is true in the sense that women in- crease the antero-posterior and the lateral diameter of the chest more than do men, owing to the functional differences between the sexes, and also due to habits of dress, etc. On FIG. 44. FIG. 45. The changes of the thoracic and abdominal The same in the female (Hutchinson). walls of the male during respiration. tin 1 other hand, adult males and children of both sexes use tin- diaphragm almost exclusively in quiet inspiration (Fiir>. 44 and 45). MUSCLES OF EXPIRATION. 81 During forced inspiration additional muscles are brought into play to permit of a more powerful inspiratory act. Be- side the muscles already enumerated, the muscles of forced inspiration are as follows : Trapezei and rhomboidei fix the shoulders ; Pectorales majores and minores, acting from the fixed shoulders, draw the sternum and ribs upward ; Sterno-mastoidei fix the upper part of the chest ; Erector sphue stiffen the vertebral column ; Serrati postici inferiores. ^ ^ -IT QuadJi lu.bo,,, U a-k^rt- Downward oacro-lum bales, ) Muscles of expiration : At the close of inspiration the va- rious muscles that raised the thorax relax, and by its own weight the thorax drops, thus compressing the lungs and ex- pelling the air. In addition there is an active collapse of the lungs due to the recoil of the elastic tissue in the sub- stance of the lung, the elastic fibres having been put on the stretch during inspiration. Also during inspiration the inter- osseous portions of the internal intercostal muscles were put upon the stretch ; when expiration begins these muscles con- tract, but their contraction is not sufficiently forcible to pull the ribs down, and the only purpose of this contraction seems to be to keep the intercostal tissues tense and thus prevent bulging of the intercostal spaces. Also during inspiration each costal cartilage was twisted in the direction of its long axis, due to the eversion of the rib twisting the rib-end of the cartilage, the sternal end being fixed. During expiration the costal cartilage tends to un- twist itself. So we may justly say there are no muscles for quiet expiration, as the act is performed by the weight of the thorax, elastic recoil of the lungs, and untwisting of the cartilage. Forced expiration is accomplished by the intervention of many muscles as follows : Interosseous internal intercostals act forcibly in drawing down the ribs, when the lower part of the thorax is fixed ; Abdominal muscles fix the lower part of the thorax and press the abdominal contents upward; 6 Phys. 82 RESPIRATION. Levatores ani and perineal muscles, hold the floor of the pelvis rigid against abdominal pressure. Triangtilaris sterni draws the costal cartilages down. Frequency of respiration : In normal adult life, with the body in repose, there are about eighteen respiratory cycles to the minute. Inspiration and expiration alternate without any appreciable pause between the two. The inspiration is some- what shorter than the expiration ; the ratio being about 5 to 6. In infants and invalids the rate is often much more rapid. The ratio to the pulse-rate is about 1 to 4 in the healthy in- dividual. During violent activity both the respiratory- and pulse-rates increase, but the ratio remains about the same. Respiratory sounds : When the ear is placed in contact with the chest-wall, or a stethoscope is used, a respiratory murmur will be heard, fairly marked during inspiration ; short and faint during expiration, in a healthy subject. It varies in diiferent parts of the chest-wall, being loudest over the large bronchi. The changes in these murmurs incident to disease of the respiratory tract are just as characteristic of the differ- ent pathological processes as are changes in the heart- sounds depending on disease of the heart ; and it is upon the recog- nition of these changes that the value of auscultation depends. Associated respiratory movements : With every inspiration (especially if rapid) there is dilatation of the nostrils; and a partial closure during expiration. The rima glottidis is in the same way opened for the ingress of air. This is like the respiratory act, in that the opening is a muscular act during inspiration, and the recoil, elastic, in ordinary breathing. Capacity of the lungs: The lungs, if filled to their utmost, can hold about 4500 c.c. of air. This total of air is divided as follows: (1) tidal air, (2) reserve air, (3) residual air, and (4) complementary air. Tidal air is the ordinary amount of air that passes in and <>ut during each quiet respiration. It seldom goes lower than the large bronchi. Its amount is 500 c.c. Reserve air is the amount of air in addition to the tidal air one can expel from the lungs in a forced expiration. It is ordinarily lodged in the bronchi and bronchioles. Its amount is 1600 c.c. EXPIRED AIR. 83 Residual air is found in the alveoli of the lungs, and remains even after the most violent expirations. Amount, 800 c.c. Complementary air is the air taken in, in addition to the tidal air, in the most forced inspiration. Amount, 1600 c.c. The complementary, tidal, and reserve airs have been called the vital capacity of the lungs. Force of the inspiratory and expiratory muscles : The force of the inspiratory muscles is greatest in people of about five feet seven and a half inches in height, being equivalent, on the average, to a column of mercury three inches high. It diminishes in people above and below this height. The force of expiration is about one-third greater; but the variations are not so regular, as there are many variations due to the fact that the expiratory muscles are used for other purposes, hence becoming stronger. Composition of inspired air : Pure air has a nearly uniform composition : By volume. By weight. Nitrogen, 79 parts 75 parts Oxygen, 21 " 25 " Carbonic acid, 04 part Ammonia and impurities, trace. Expired air : After its sojourn in the lungs there are marked changes in the respired air. Its temperature is changed to nearly that of the body, regardless of the tempera- ture of the atmosphere. The oxygen is decreased, carbon dioxide and aqueous vapor are increased ; also there are present many minute particles of volatile organic bodies. Nitrogen is unchanged. The composition of expired air is about as follows : By volume. Nitrogen, 79 parts Oxygen, 16 " Carbon dioxide, 4 " Water increased (from 6 to 30 ounces per day). Volatile organic bodies are added. The total weight of expired air is somewhat heavier than 84 RESPIRATION. that of inspired air, because the amount of carbon dioxide gained weighs more than the amount of oxygen lost. In temperate atmospheres the volume of expired air is greater than that of the inspired air, because the air has been expanded by being heated. If, however, the expired air lie reduced to the temperature of that inspired, the volume of expired air will be found less than that inspired, as the car- bon dioxide gained is less bulky than the oxygen lost. Value of nitrogen: The nitrogen in the atmosphere serves merely as a diluent ; pure oxygen would be too powerful to serve for respiration. We do find a little nitrogen in solu- tion in the blood-plasma, but it probably serves no purpose in the body economy. External respiration : By external respiration is meant the interchange of gases in the respired air and the gases of the blood while in the pulmonary capillaries. It is true the ordinary tidal air of respiration, lodged as it is in the upper air-passages, does not come into actual contact with the pul- monary capillaries, but by virtue of the physical laws of diffu- sion of gases the oxygen in the tidal air is diffused into the reserve air, and carbon dioxide and other wastes of the re- serve air are diffused into the tidal air. A similar exchange takes place between the reserve air and the residual air. Thus it comes about that the residual air, while never leaving the air-cells of the lungs, is constantly revivified with fresh oxygen and constantly gets rid of its carbon dioxide and other impurities. The blood coming to the pulmonary capil- laries through the pulmonary arteries is rich in carbon diox- ide and other wastes, but poor in oxygen ; on the other hand, the air in the alveoli is rich in oxygen, but poor in carbon dioxide. Hence the oxygen in the air is under high tension as compared with the oxygen in the blood; and likewise the carbon dioxide of the blood is under higher tension than the carbon dioxide of the air. Owing to these ditlcivnces in pressures, interchanges of the gases readily take place, so as to equal i/e the pressures. In the blood oxygen is in loose chemical combination with the haemoglobin (<>x\ -h \ moglobin). The amount of oxygen in the residual air of the pulmonary alveoli is estimated at NERVOUS MECHANISM OF RESPIRATION. 85 about 10 per cent., that of expired air being 16 per cent. It is found that unless there is present about 4 per cent, of oxy- gen there is no tendency for the blood of the pulmonary arteries (venous) to take up fresh oxygen ; or, the tension of the oxygen in the reduced hemoglobin of venous blood is about 4 per cent., and unless the oxygen-tension in the lungs is greater there is no absorption of oxygen. But, as we have seen, the amount of oxygen amounts to at least 10 per cent., and therefore the excess is sufficient to exceed the demands, and the exchange is readily made by diffusion through the thin capillary walls of the alveoli. On the other hand, the tension of the carbonic acid in the pulmonary arteries is much higher than in the alveoli, and hence the extrusion of this gas by diffusion is accomplished. The following changes in the blood are noted after passing through the pulmonary capillaries : (1) Color, deep purple to bright scarlet by oxidization of the reduced haemoglobin i. e., from venous becoming arterial blood ; (2) gains oxygen ; (3) loses carbonic acid ; (4) becomes cooler ; (5) coagulates more readily. Internal respiration : Internal respiration means the ex- change of gases between the arterial blood and the tissues of the body. This interchange takes place through the walls of the systemic capillaries, and depends on the same physical laws (differences in pressure) as already described for the pulmonary exchange. The arterial blood, rich in oxygen, gives oxygen to the needy tissues, and in return relieves the tissues of their burden of carbon dioxide and other wastes. In the systemic capillaries the blood changes from arterial to venous. Nervous mechanism of respiration : Respiration is described as one of the so-called automatic acts, but is profoundly modi- fied by the reflexes and partially controlled by the will. Throughout our lives we continue to breathe unconscious of the fact, unless our attention is called to it. A dash of cold water on the skin or a disagreeable mental or visual pict- ure will produce an uncontrollable gasp ; these are examples of reflex influences on respiration. One may so exert will as to increase or decrease the rate of respiration, but only up to 86 RESPIRATION. certain limits. It is impossible to commit suicide by " hold- ing the breath." Centres for respiration : In the medulla oblongata arc cen- tres for each half of the body, from which arise automatically the rhythmical impulses for the inspiratory act*. These so- called automatic impulses are generated by the presence or absence of oxygen in the blood (not presence of carbon dioxide) bathing the medulla. If the blood in the medulla is rich in oxygen, there is no impulse for an inspiratory act ; but as soon as the oxygen is diminished to a given point the respiratory centre is stimulated to make an inspiratory effort. The nerves which convey to the muscles of inspiration their impulse are the phrenics (to the diaphragm) and the intercos- tals (to the intercostal muscles). In addition to the auto- matic impulses from the centres, there are certain regular reflex impulses (other than the unusual ones mentioned above cutaneous reflexes, etc.) that are conveyed to the centres by the pneumogastric nerves. The pneumogastric nerves arising in the medulla scud branches to the lungs that terminate in filaments distributed to the walls of the air-cells. These fibres are afferent, and are stimulated by two factors : (1) The lack of oxygen in the air of the alveoli after expiration ; (2) During the partial col- lapse of the lung at the end of expiration these filaments are mechanically pinched by the walls of the air-cells. The stimuli thus generated in the terminal filaments of the pulmonary branches of the pneumogastric nerves convey their impulses to the centres of respiration, producing an ad- ditional impulse for inspiration. The condition (amount of oxygen) of the blood bathing the respiratory centre is, however, the prime factor, the afferent impulses of the pneumogastrics being only secondary. While there are distinct expiratory centres in the me- dulla for expiration, they act to produce forced expiration only when properly stimulated by afferent nerves as, for example, if the superior laryngeal nerve be stimulated, the result is a forced explosive expiration, or cough. Under <>/-(f innri/ oircurngtance* when the inspiratory act is accomplished, the condition of the blood no longer calls for SECTION OF VAGI. 87 more oxygen, and, the inspiratory impulse ceasing, the mus- cles of inspiration relax and expiration is a purely passive act. In dyspnoea, however, the expiratory centres are stimu- lated. Synchronism of respiratory centres : Although there is a respiratory centre for each half of the body, nevertheless, owing to their association-fibres, the two centres work syn- chronously. By making a median section of the medulla, the centres are disassociated, so that, if proper artificial stimuli be applied, one half of the chest may be made to inspire, while the other half expires. Lack of oxygen in the blood : If the need be only moderate, there will be increased effort of both expiration and inspira- tion, and the respirations will be rapid a condition known as hyperpnoea. As the oxygen becomes less and less abundant the symptoms become more severe, and the condition is known as dyspnoea. The dyspnoea increasing, the respiratory efforts become very violent, and the condition of asphyxia is seen. In this the face is blue, eyes staring, face anxious, and respir- ations very rapid and strident. Then follows a convulsive condition which is brief, the convulsions being very violent and involving the whole body. After this the patient lapses into a state of exhaustion, in which the respirations are slow and very feeble, and the general condition is one of collapse. Death ensues very soon. Apncea is absence of breathing. Section of vagi : If one pneurnogastric nerve be divided below the offshoot of the inferior laryngeal nerve, there is a temporary excitation of the respiratory rate, due to the mechan- ical stimulus of cutting ; but soon the respirations return to normal, .the other vagus taking up the work of both. If both vagi be divided below the inferior laryngeal nerve, there is an excitation of respiration due to the stimulus of cutting, soon followed by slower and deeper inspirations and active expirations. Stimulation of the central ends of cut vagi will bring the rate and character back to the normal. If the superior laryngeals be divided, a cough or expiratory sniff follows the act of cutting ; later on the subject dies of 88 RESPIRATION. ' foreign-body pneumonia," because the sensation of the lar- VMX being lost foreign particles pass the. larynx without being coughed out, enter the lungs, and give rise to a fatal pneu- monia. Section of the inferior laryngeals gives the same result ; for although sensations in the larynx are present if the superior laryngeals are intact, all motion in the larynx is lost and coughing is ineffectual. Vitiated atmosphere : In ill-ventilated rooms the air of the room is used repeatedly, and, besides becoming partial!} de- prived of its oxygen, is charged with carbonic acid and with putrescible nitrogenous organic matters. This gives rise to an atmosphere which is intolerable to one who enters from fresh air. That such a condition is unsanitary needs no argument. The mere presence of the excess of carbonic acid is not in itself injurious; but the amount of carbon dioxide is indica- tive of the amount of organic matter present, and it is the latter that are highly poisonous. It is generally accepted as a fact that about 1000 cubic feet of air-space per head must be allowed in sleeping-quarters, and sufficient facilities for exchange of air to allow complete change in each hour. This ventilation must be accomplished without exposure to draughts. Effect of respiration on the circulation : As the heart and its great vessels are contained together with the lungs in the thorax, an air-tight cavity, there naturally results a certain alteration in pressure on the heart when the capacity of the thorax changes. Though the expansion of the lungs during inspiration tends to compensate for this increased capacity, it never does so fully, as part of the atmospheric pressure is expended in overcoming the elasticity of the lungs. There- fore, during inspiration the pressure on the heart become.- considerably less than the atmospheric pressure exerted on the bloodvessels outside of the thoracic cavity. This differ- ence varies from 5 to 8 mm. of mercury during the pause, up to 25 or 30 mm. at the end of a deep inspiration. The result of this is to draw more blood through the veins into the heart during inspiration, and consequently to increase ar- terial tension. The relative values of intrathoracic press- SPECIAL RESPIRATORY ACTS. 89 ure and arterial tension may be graphically expressed as in Fig. 46. Special respiratory acts : There are a number of involun- tary and voluntary special respiratory acts, largely reflex, which are dependent upon modifications of inspiration and FIG. 46. i e i J Comparison of blood-pressure curve with curve of intrathoracic pressure. To be read from left to right, a is the blood-pressure curve, with its respiratory undu- lations, the slower beats on the descent being very marked ; bis the curve of intrathoracic pressure obtained by connecting one limb of a manometer with the pleura! cavity. Inspiration begins at i, expirati9n at e. The intrathoracic pressure rises very rapidly after the cessation of the inspiratory effort, and then slowly falls as the air issues from the chest ; at the beginning of the inspiratory effort the fall becomes more rapid (Foster). expiration e. g., sighing, hiccough, cough, sneezing, speaking, singing, sniffing, sobbing, laughing, yawning. Sighing : This results from a prolonged inspiration, the air passing noiselessly through the larynx and being rather sud- denly expelled. Hiccough: This resembles sighing, but the inspiration is sudden, due to spasmodic action of the diaphragm. Cough : This results from a deep inspiration followed by a forced and sudden expiration, during which the glottis is momentarily closed by spasmodic action of the vocal cords. Sneezing : In this case, following on a deep inspiration, by a sudden and forced expiration the air is directed through the nasal passages. Speaking : In this case there is a voluntary expiration, and the vocal cords, being rendered tense by their muscles, vibrate as the air passes over them, producing sound. 90 DIGESTION. Singing : This only varies from speaking in the differing tension and position of the vocal cords and the consequently different sounds produced, Sniffing: This results from rapidly repeated but incom- plete nasal inspirations. Snhltiny: This consists of a series of convulsive inspira- tions, during which the glottis is more or less closed. Laughing: This results from a series of short and rapid expirations. Yawning : This is an act of inspiration, more or less in- voluntary, accompanied by a stretching of various facial muscles. Sucking: This is not really a respiratory act. It is caused chiefly by the depressor muscle of the os hyoides, which, by drawing down and back the floor of the mouth, produce a partial vacuum in it. DIGESTION. By digestion we indicate the process by which food is in- troduced into the body and prepared in such way that it becomes suitable for absorption and tissue-nutrition. The process may be divided logically and conveniently into masti- cation, insalivation, deglutition, stomach digestion, intestinal digestion, and defecation. The mouth : The first portion of the alimentary tract is the mouth. In this is received the food destined to support the body. It is a cavity contained between the jaws and bounded laterally by the cheeks. Its roof is formed by the hard and soft palate, and its floor by the tongue. It is lined through- out with mucous membrane, which is provided with numer- ous mucous glands, and into it open the ducts of the salivary glands. The tongue is both a prehensile organ and also the chief seat of the sensation of taste. In this cavity occurs the process of mastication. Mastication : When a mass of food enters the mouth it is caught by the tongue and moved to a position such that it may he erushed and ground between the upper and lower teeth. This process is favored by the action of the tongue and of the cheeks, which not only crush the softer food- MASTICATION. 91 masses, but bring the less tractable portions repeatedly under the action of the teeth. There are during life two sets of teeth, temporary and per- manent. In the first set are twenty teeth , and in the second thirty-two. The permanent set are arranged as follows : Upper. Lower. Molar. Bicuspid. Canine. Incisor. Canine. Bicuspid. Molar. 3214123 A tooth may be roughly described as consisting of a crown, neck, and fang or fangs. The crown is the portion which pro- jects above the margin of the gum ; the neck is the con- stricted part below the crown which is covered by the free FIG. 47. Section of human molar tooth, magnified (Owen). edges of the gum ; while the fang or root includes all below this. On making a section of a tooth, for instance a molar (Fig. 47), it will be seen that there is a cavity within the 92 DIGESTION. crown which extends into and through the fangs, opening by a small aperture at their apices. This is the pu/jt-rdrify, and contains in the living tooth the pulp. A portion of the tooth- structure resembles bone, anatomically as well as chemically. The crusta petrosa, or cement, is true bone, and covers in the r<>- teids. When distended it measures about fifteen inches from end to end, and about five inches antero-posteriorly. Glands of the stomach : If one looks closely at the mucous surface of the stomach, it is seen to present a sort of reticu- lated (Fig. 49) appearance, the meshes being larger at the FIG. 49. Free surface of the gastric mucous membrane, viewed from above, from pig's stomach ; cardiac portion ; moderately magnified. pyloric than at the cardiac end of the stomach. It is in the interstices of this mesh that the glands open. The openings are smaller at the cardiac than at the pyloric end, and the character of the glands changes: we therefore speak of \\\<> varieties of gastric glands (1) peptic ; (2) pyloric. (1) The peptic glands are arranged in groups throughout the stomach, but not so abundantly at the pyloric end. They often consist of a simple tube dipping into the surface and lined with columnar epithelium (Fig. 50), but they may be GASTRIC JUICE. 97 branched^, e., several glands may empty into a common duct. The columnar epithe- lium in the deeper portion Fro. 50. of the gland contains large, almost globular, cells, which are known as peptic cells. (2) The pyloric glands, or mucous glands, like the peptic, may be simple or compound. The ducts are larger, and the large cells are wanting (Fig. 51). During digestion the cells of both varieties of glands become swollen, and in them are found granules which are supposed to be pep- sin, or that from which pepsin is formed. Gastric juice : When the stomach is not at work it con- tains no gastric juice, but is bathed in an alkaline mucus. As soon as food enters the organ, however, it immediately begins to secrete considerable nnantitios nf nn 'irirl fliiirl Compound gastric follicle, from the car- 1Cl > diac Dortion of the human stomach. which soaks into and mingles with the food. The cele- brated case of Alexis St. Mar- tin, who had a gunshot-wound resulting in gastric fistula, enabled Beaumont, surgeon U. S. A., to investigate accu- rately its composition. It is a limpid, colorless fluid of specific gravity 10011010 and acid reaction. It contains about \ per cent, solid matter. Its composition is nearly Water 99.50 Pepsin .25 Hydrochloric acid .05 Salts (alkaline chlorides and phosphates) .20 7-Phys. 100.00 diac portion of the human stomach. 1, excretory tubes leading to the sur- face ; 2, tubular follicles containing spheroidal cells (Kolliker). 98 DIGESTION. This composition is not constant, as the proportions vary considerably, HC1, for example, being present much more abundantly in some cases. Secretion of gastric juice: The stomach secretes about fifteen pints of gastric juice per diem. The hydrochloric acid is probably secreted by the cubical parietal cells of the peptic glands. Very little seems to be formed by the pyloric glands. FIG. 51. Tubular follicles, from pyloric portion of pig's stomach, showing their csecal ex- tremities and epithelial lining; at a is the torn end of a follicle, showing its cavity more highly magnified. The pepsin comes from the globular cells in the peptic glands. These cells are supposed to form a subst:mr<- called pepsinogen, from which pepsin is derived. Pepsin is derived for commercial or for experimental purposes from fresh stomachs by scraping the surface and dissolving out the ferment with cold water, or by mincing the mucous mem- brane and extracting the ferment with glycerin after dehydrat- ing with alcohol. Functions of the gastric juice : The principal function of the gastric juice is the transforming of proteids into peptones. CHARACTERISTICS OF PEPTONES. 99 This action depends upon the presence of both pepsin and acid. The first change which occurs is the formation of acid albumin, but as the action of the ferment continues the acid albumin is transformed to peptone. The presence of acid al- bumin is demonstrated by the addition of an alkali, which precipitates it. The action of the pepsin in converting proteids to peptones is called a proteolytic action, and chemically its action is to cause a hydration of the proteid molecules. Milk is curdled in the stomach by a ferment, aside from pepsin, which is derived from t the gastric juice. This action takes place in the absence of hydrochloric acid. Rennet (de- rived from the fourth stomach of calves) is used for this pur- pose in cheese manufacture. Fats are unaffected, except that the albuminous capsules of fat-cells in adipose tissues are digested and the oil set free in globules. Carbohydrates are unaffected. In conclusion, therefore, it can be said that gastric diges- tion is of a preparatory character, fitting the food for further digestion in the intestines. Not only the carbohydrates and fats pass the stomach practically unchanged, but part of the proteids, the hemipeptones, are not completely digested. That it is not indispensable is proven by the fact that re- cently the entire stomach was removed from a woman, yet she showed no ill effects, but lived for a year or more, and finally died from another cause. Characteristics of peptones: (1) They are diffusible i. e., have the property of osmosis, or passing through an animal membrane. This is of great importance in digestion, for if this property were absent no animal food could be absorbed from the intestines. (2) They are very freely soluble in water and neutral solutions. (3) They do not respond to the chemical tests for other proteid substances. They are not precipitated by heat and the mineral acids, but are precipi- tated by tannic and picric acids and by the bichloride of mercury. Muscular action of the stomach : The stomach is elastic, and is supplied with circular and longitudinal muscles in its 100 DIGESTION. middle coat. These muscular fibres are capable of producing peristaltic movements of the organ, which turn the food over and over during the process of digestion. This elastic pouch is closed at each end by strong, sphincter-like, circular bands of muscles at the cardiac and pyloric openings, and until the stomach digestion is well advanced none of the contents escapes; but as the peptone-making advances the pyloric opening permits the escape of the digested food, and this is aided by strong peristaltic efforts on the part of the stomach at its pyloric end. Toward the end of digestion the pylorus permits the escape of undigested as well as of digested mat- ter. The circulation of the stomach-contents is circumfer- entially toward the pylorus, but centrally toward the cardiac opening. The digested food as it leaves the stomach is called chyme. Time required for digestion : The time varies with the kind and amount of food from one to five or six hours. Digestion is favored by rest of the stomach before eating, by gentle ex- ercise of the mind or body, by an undisturbed mental condi- tion, and by a healthy condition of the body. Conditions favoring gastric digestion : The temperature of the body is most favorable, and the presence of acid prefer- ably HC1 is essential. For the best results the percentage of HC1 should be about -^th of 1 percent. Excess of acid, or neutralization, will stop the process. The movements of the stomach assist digestion by thoroughly mixing the food and the gastric juice. Digestive secretion does not continue except during the presence of food. Also the removal of completed products assists in the conversion of the re- mainder. Capacity of the stomach : About a quart in the adult, but its muscular walls enable it to contract so as to fit its con- tents if much or little. When empty the stomach is tightly contracted. Nervous mechanism of gastric digestion: The pncumo-as- trie and sympathetic (splanchnic from the solar plexus) are the nerves which supply the stomach, and besides then- there lerous ganglia in the stomach-walls. The ordinary INTESTINAL DIGESTION. 101 motion-stimulus of the organ lies in the intrinsic ganglia. Irritation of the pneumogastric nerve causes contraction ; its division, cessation of peristalsis. But, further than this, the vagus has control to a considerable degree over secretion in the stomach. Digestion of the stomach after death : When death occurs at a time when the stomach contains food and gastric diges- tion is going on the walls of the stomach are often partially digested, even to such an extent that a perforation occurs. This condition is often found in post-mortems on the human body. Many reasons have been brought forward as to why this does not occur during life, but no entirely satisfactory theory has yet been propounded. Some have thought that- gastric ulcers were merely an example of localized self-di- gestion. Vomiting: The regurgitation of food from the stomach through the cardiac orifice, and thence through the mouth, may occur when the cardiac opening is free and the pylorus is closed. This is usually a reflex act, and is performed by the contraction of the stomach, aided by the pressure of the abdominal muscles opposing the fixed diaphragm. It may be described as a reversed peristalsis. The stimuli which ex- cite the reflex may be either local in the stomach or periphe- ral. Violent irritation of the gastric mucous membrane will excite it; also mental impulses, from ocular, auditory, or olfactory sources; injury or irritation of the testis, ovary, kidney, etc.; unusual motion, as swinging ; certain diseases ; and effort of will in some is sufficient. There is a vomiting- centre in the medulla acting through the pneumogastric nerve. Absorption from the stomach: Although the stomach is not designed as an organ of absorption, nevertheless probably some water and peptones are absorbed by the capillaries of the stomach-wall. Small Intestine. Intestinal ( digestion : By the peristaltic action of the gut the food is carried on through the length of the organ, but its 102 DIGESTION. progress is more or less impeded by the vahruke conniventes, which are folds of the mucous membrane extending trans- versely across the intestine at right angles to its long axis, and occupying usually one-third or one-half of the circum- ference, but sometimes extending all the way around. They commence close to the stomach, and are well developed in the upper two-fifths of the small intestine. They then gradually diminish in size and number, and finally disappear at the commencement of the lower fourth. These folds of the mu- cous coat not only retard the too rapid advance of food, but cause it to be thoroughly exposed to the action of the diges- tive fluids (Fig. 52). Soon after passing the pylorus food FIG. 52. Portion of small intestine laid open to show valvulse conniventes (Brinton). comes in contact with the alkaline secretions of the small in- testine and of the liver and pancreas. In the small intestine the food is still further prepared for absorption, and from this part of the alimentary tract the digested food is taken up for body-nutrition. The peristaltic action is controlled by Un- sympathetic system of nerves. Auerbach's plexus lies be- tween the circular and longitudinal muscular coats. It is also known as the plexus mesentericus. The blood-supply is also controlled by the sympathetic system. Meissner's plexus lies beneath the mucous coat, and is regarded as the source of control of the blood-supply and of the function of absorp- tion. Glands of the small intestine: There are three types of irland- peculiar to the small intestine: (1) Lieberk film's, (2) Brunner's, and (3) Fever's glands. GLANDS OF THE SMALL INTESTINE. 103 FIG. Glands of Lieberkiihn : These glands (or follicles or crypts) are thickly distributed over the whole surface of the small and large intestine, being larger in the large intes- tine. They are simply tu- bular depressions in the mucous membrane, lined with columnar epithelium, which contains occasional large " goblet "-cells. Brunner's glands are found in the duodenum alone, and are situated in the submucous tissue. They resemble the pylo- ric glands of the stomach, and, like them, are usually compound glands. The duct of the gland passes up through the mucous membrane and opens at its surface (Fig. 53). Peyer's glands: These are of two varieties (1) solitary and (2) agminate. (1) The solitary glands consist of a rounded mass of whit- ish adenoid tissue about ^-th to y^th in. in diameter, situated in the submucous tissue, but often projecting to the surface of the intestine. Each lymphoid mass is surrounded by Lieber- kuhn's follicles (Fig. 54). FIG. 54. A vertical section of the dvwdenum highly magnified. 1, a fold-like villas; 2, epithe- lium of the mucous membrane ; 3, orifices of the tubular glands ; 4-5, orifice of a duo- denal gland ; 6-7, two vesicles of the latter, more highly magnified, exhibiting the epi- thelial cells lining their internal surface (Leidy). Portions of the mucous membrane from the ileum. moderately magnified, exhibit- ing the villi on its free surface, and between them the orifices of the tubular glands. 1, portion of an agminated gland ; 2, a solitary gland ; 3, fibrous tissue (Leidy). 104 DIGESTION. (2) The agminate glands (Peyer's patches) consist of groups of these adenoid masses, making " patches " in the mucous membrane \ to 3 in. long and about \ in. wide. Intestinal juice: The intestinal juice, or succus cuff-rim*, is the secretion of the intestinal glands. This secretion is yellow in color and is markedly alkaline. Its effect upon diges- tion is not fully understood, but it probably has some effect upon saccharose. However, its chief function seems to be to supply loss of fluid L e., to take the place of that which is absorbed as digestion progresses. At any rate, the contents of the small intestine as they enter the colon are about as fluid as when they leave the stomach. Other digestive secretions: The most important digestive fluids that act upon the food while in the intestines are not derived from the intestines, but are poured into the small in- testine near its beginning. These secretions are two in num- ber, and are formed respectively by the pancreas and the liver. The Pancreas. The pancreas is an organ lying in the upper part of the abdomen in contact with the duodenum : in length it is about six inches, and is thicker at its right or duodenal end. It is a conglomerate gland, resembling in structure the salivary glands. During digestion it is active, but is quiescent iu the intervals. Its secretion, pancreatic fluid, is discharged into a main duct which receives branches from the lobes of the gland, and is emptied with the bile through a common open- ing about two or three inches beyond the pylorus. During digestion the cells of the organ become granular, and the granules are thought to consist of the substance from which the ferments of the pancreas are derived, -i/ntoyen, rather than of the ferments themselves. Pancreatic juice : The pancreatic juice is a clear, colorless fluid, having an alkaline reaction and a notably viscid con- sistency. It coagulates with heat, and is made quite gelat- inous by cold. Specific gravity, 1015. Its composition varies. DIFFERENCE IN ACTION OF TRYPSIN AND PEPSIN. 105 Composition of pancreatic juice : Water 90 Organic matter: Ferments, ") Serum-albumin, I Alkali-albumin, ( Fats, soaps, etc.,J Inorganic salts (chiefly sodium chloride) . . 1 100 Pancreatic ferments: (1) Trypsin, a peptone-forming (pro- teolytic) ferment, which continues the digestion of proteids begun in the stomach. It forms a peptone which resembles the stomach peptone in its reactions. This ferment, unlike pepsin, only acts in an alkaline medium. It acts less vigor- ously upon gelatins and other nitrogenous bodies. (2) Amylopsin, a starch-changing (amylolytic) ferment, by which starch is converted to maltose, as by the ptyalin in the saliva. (3) Steapsin, a ferment by which fats are broken up from the large globules and emulsified or saponified in alkaline media. It is claimed by some that this is not a ferment- action, but is the result of the action of the alkaline intestinal contents upon the fat. Of these processes the emulsifi,cation, or breaking the fat- globules into minute particles, is by far the more important, as it allows this form of food to be absorbed from the gut. Milk is an excellent example of a natural emulsion. Sapon i fixation (or soap-making) results from the fatty acid combining with an alkali, forming the corresponding salt and glycerin e. g. : Stearin 4- Potassium hydrate = Potassium stearate + Glycerin. Difference in action of trypsin and pepsin : These two fer- ments act on proteids in different ways. Under the influence of pepsin the proteid first swells, then gradually becomes softer, changes in color, and breaks down into a grumous 106 DIGESTION. mass. When exposed to the action of trypsin, however, pro- teids do not swell up, but are eroded or oaten away, seeming at times as though they were full of worm-holes. lYp.Mu cmi verts proteids iiitohemi-albumose and anti-albumose, which it later converts into hemipeptone and antipeptone. Tryp- sin splits the proteids at once into deutero-albumose, and alni carries the process a step further than pepsin, splitting the hemipeptones into leucin and ty rosin. Function of pancreatic juice : It is most active during diges- tion, and is peculiar in having an effect upon all forms of food which require preparation for absorption upon proteids, starches, and fats. Conditions favoring pancreatic digestion : Moderate heat (100 F.), an alkaline medium, and the removal of the prod- ucts of the ferment-action as soon as the change is com- pleted. The Liver. The liver is the largest gland in the body, and is situated in the upper part of the abdominal cavity. It secretes a fluid known as the Itilc or gall, which is stored in a bladder lying attached to its lower surface. The functions of the organ are : (1) secretion, (2) excretion, (3) glycogenic function, and (4) elaboration of urea. Secretion of the liver: The secretion of the liver is stored in the f/df/-Mile, but the former is characteristic of the bile of herbivora, and the lat- ter, bilirubin, of the bile of carnivora. The pigments are FIG. 56. Cholesterin from the contents of an encysted tumor. orygtallizable, and are insoluble in water. The crystals have the "Teen ; in /.- add fuming nitric (nitroso-nitric) acid, and there results a play of colors which is best seen when the bile solution is in thin layer on a white plate. The presence of the bile-pig- ments is shown also by absorption-bands in the spectrum. Bilirubin is derived from haemoglobin, and biliverdin from the bilirubin, as they are chemically closely allied. GLYCOGENIC FUNCTION OF THE LIVER. 109 In considering the death of the red blood-corpuscle, it has been already stated in the chapter on the Blood that the haemo- globin from the disintegrated red corpuscle is eliminated by the liver. This is the haemoglobin that makes the bile- pigment. Cholesterin is a crystallizable, insoluble substance which belongs to the alcohol group in chemical composition. Best recognized by microscopic appearance of the crystals (Fig. 56), though it may be tested chemically by the addition of sulphuric acid, which gives a red reaction. Use of bile in digestion: (1) The alkaline reaction of the bile aids the pancreatic and checks the pepsin digestion ; it aids in the emulsion of the fats, and is probably very active in this process. (2) It moistens the mucous membrane and favors the absorption of digested food. (3) It acts as a natural purgative and as a natural antiseptic, and in this way is very essential to the proper performance of the diges- tive process. As a purgative bile acts by stimulating peri- stalsis. Excretion of the liver : The bile for the most part, in nor- mal conditions, is a sort of circulating fluid : it is secreted by the liver, poured into the intestines, and reabsorbed from them, to be returned through the portal vein to the liver for recirculation. There is, however, a small proportion of biliary matter, about one-sixteenth, which is not absorbed, and this consists chiefly of the pigments of the bile. The salts are nearly all reabsorbed in the assimilation. Further than this, the liver is found, so to speak, to filter materials which would be poisonous if circulating in the general system, and either to reject them at once, or to store them up and return them slowly back to the intestine. The excrementitious material from the liver is known as stercobilin. Stercorin is found in the fa3ces, and is thought to be an excretion of the liver : it closely resembles cholesterin, and is supposed to be a modifi- cation of cholesterin by digestion. Whether or not the ster- corin (cholesterin) is an excretion of the liver corresponding to the urea of the kidney is somewhat uncertain. Glycogenic function of *the liver : The liver normally forms a substance resembling starch in its chemical composition. 110 DIGESTION. This is known as glycogen, and is formed from glucose taken up by the portal circulation. Its chemical formula is that of starch (C 6 H 10 O 5 ), and it is derived from glucose (C 6 H 12 O 6 ) by dehydration, and is rapidly changed by diastatic ferments to glucose. This process is known as the glycogenic function of the liver : its use is the storage of a fund of carbohydrate mail-rial (an "animal starch") to maintain a steady supply to the system. Elaboration of urea : This subject will be discussed under the Urine. Large Intestine. Structure : While the villi and valvulse are absent, yet throughout its whole extent there are found tubular and soli- tary glands which closely resemble FIG. 57. those of the small intestine. The tubular glands, however, are more numerous, longer, and more closely set together (Fig. 57). Digestion : The chyme which enters the large intestine still continues under the influence of the ferments, and the process of digestion continues. section of the mucous mem- The food ma 7 "ndergo acid fermon- brane of the colon, i, free tation here, but there is no new diges- surface exhibiting the onh- . . ~, ces of the tubular elands; tive action. Inat the large intestine m^Sr' 88 "^' 1 ' may have the power of acting upon food is shown by the absorption of fats, proteids, etc., which are taken in nutrient enemata. Defecation : The expulsion of the refuse of digestion from the intestine is partly a voluntary act, but more especially re- flex. The voluntary act is the pressure of the abdominal muscles upon the contained viscera, while the reflex is an in- creased peristalsis in the sigmoid flexure and rectum and the relaxation of the sphincter. The centre which governs this act, so far as it is reflex, lies in the lumbar region of the spinal cord. FACTORS FAVORING ABSORPTION. Ill ABSORPTION. Absorption is the process by which the digested food is taken from the intestines and carried into the blood, whence it is taken to nourish the cells. The same term is applied to the removal of worn-out material from the tissues. Chyme is the name given to food after digestion. By digestion, the proteids, starches, arid fats, which were not dialyz- able, have become peptones, sugars, and emulsified fat. All these products of digestion are readily capable of di- alysis, and therefore ready for absorption. The absorption takes place through the bloodvessels and lymphatics of the intestine. By dialysis we mean the property of fluids which enables them to pass through animal membranes osmosis. This we have seen is possessed in a high degree by the ingredi- ents of chyle. The reverse process may occur, and fluids (serum) from the blood may similarly be drawn into the intestinal canal, as is seen when the salines are used as purga- tives. Too much importance must not, however, be attached to osmosis as accounting for the phenomena of absorption. Under certain circumstances substances are absorbed inde- pendently of this principle. For example, when tested ex- perimentally outside the body, sugar is less diffusible than sodium sulphate, yet it is absorbed more rapidly from the intestine. Certain coloring-matters are not absorbed at all, the cells appearing to exert some selective action. It is also difficult to explain by osmosis the absorption of emulsified fats. It is probable that the protoplasm of the living cells on the walls of the intestine has some specific action. Sites of absorption : In the stomach and large intestine the absorption is very much less than in the small intestine, but there is reason to think that there is considerable ac- tivity to absorption from the entire gut so long as digestion continues. Factors favoring absorption: (1) The valvulse conniventes greatly increase the area of the intestinal surface, and by their 112 ABSORPTION. shelf-like formation delay the advance of chyme. (2) The villi of the intestine not only increase the area, but are the special organs of the function of absorption. (3) The con- traction of the intestine upon its fluid contents also favors, mechanically, the filtration of the contents through its walls. Villi of the intestine: The villi are almost innumerable, minute, teat-like projections from the surface of the wall of the intestine. They are very numerous in the small intestine, but none is found in the large gut. Each villus is covered by an epithelial layer, and within, FIG. 58. supported by areolar tissue, is a delicate capillary network of bloodvessels, a muscular layer (muscularis mucosse), and a more or less branched ending of a lac- teal vessel (Figs. 58 and 59). FIG. 59. a An intestinal villus. a, layer of cylindrical epi- thelium, with its exter- nal transparent striated portion ; hb. blood vr>sHs enterlngftnd leaving the villus ; c, lymphatic ves- sels, occupying its cen- tral axis (Leydig). Patch of Peyer's plands, tVom the lower part of the ileuin, showing villi (mag- nified). The ileo-Cffical valve slio\v> tlie absr\s a more Complicated form of tnhnlar inland in that it is convoluted and tortuous. ("2) Compound tulmtnr of nllimale glandular elements grouped ahoul the main tulmle, which acts duct. This form of inland is found in kidncN . testlS. -nli \arv and mammarx j-land-. lirnnner's o-lands ( Ki^-. and in maiix of ihe other ^lan dnlar structures. glands, in which 1 lie ^landnlar structure is diviled inlo lol> ulcs, or tifiin'. These "-hinds in i\ he regarded as a relineinenl of the compound luhiilar VarietV, and example^ are found in (he sali\ar\ -dand and in the Mi-ihoiniaii follicles Processes of secretion : ( 1 i P.y /*A//.v/V^/ procc e \ iz., Ultra ti"ii and dialysis the cells are aide to separate from the .( un,. o| Itiiiniii-r's II.MII IMIIIIMM ml. vim,. v . vs u N N U7 . n(-. xxhu-h uuk Up u-mW (hat (ho t\uw of dialysis may uv> ami K tH>itsH|iioittlYft twt & I 1 * . m^a^ain, i ho taot that tho oomlitmii ot iogfftdtatl makos din i MU (ho jwnnanonoy uf tht> ,i wmlil I'vx .'At*mi\M/ iwnnvsst- ill,- vUs of glamls -.nui MI iiu- hi.s-ii pltimt Mid tidd u il i jmHHsss^ av jnvuli in iiu- iniiuMii produdnf f laodi idivtiy) ptptlo^ tod \*\\\- Dlsohrtrgo of soaottons : Tlu> Mvifiions max tVMi ilu- i'l.iu.l-. M loon M ilu-x .uv' t,MMi-l. as iu tho '1,1-. riu-x in.i\ IT lottg ^^aimnl in au x-I,|>,-l (.Mm ui du- >,. - d ill---' -l.inJ u KOM M i'^ ^ M>U ,sv.i i.Mi.il , ,-r in .1 fiilK ( lr\rl,.|u-,l furm in thoir ;n :!. in.l- , -n,li .1- liir Li, lux m.l 1 , \\ hioh hdiUI^ ( !,-(> small iu>Miu(-,. IMK inoinni^ jini.Mitn \\ lirn itimukted Oonditloiui ffiotUif flMidulMr Activity : (M) I f tin* Minono/ */' A/MM/ MtwiHtf M^H//A (I f/All(/ IT i n, I r.i- r, I , (li,-ir will |>o |Q. I MX in Crf & lun, u, MI . .iii,l, , ,MIX, i , U , .i-. .1 ml,-. during functional iotiv)^j it"- |knd \\iii Inoi^Me in vas- rul.MKv I iu- - 1, .in. i, li. I,M r\.iin|.l t -, ilnrin^ tli^vstiou || .jinh- .MI- -,.i ; -, ,1, IMK \\ lirn ulh- i-. ii|.plir,l \\ ill. inn. li 1, blood . ' i i'x Hi.- eoloi l--|iMiinu dnnn^its \wv\w\ uf 'M (A) An luwwtw m M^ md^id/ i^w*i wA*VA M*^ //Aim/ uo^ inuul.Kr ili,- ; .|.iM.l (o gmitvr .'il-Mt .nui InoTMMI (l>^ }UH>- dyotion frf ii- -i.n.,1 rim-.. di- uoounl oi mva is inoh^a^nl l\ -Ml Ml.1,.1 ,- ,.| Ill(l,.,' t -Il,.ll U.I (,-, .1-. hx |XtfOlM| Of ^ I" ret -I nUrogenoui matter ". d.,- bl^ad, as h t v a f\ll mt] oi animal UwL I -n the left is a tubule alone; in the middle is a tubule along with the bloodvessels ; on the right an- bloodvessels only. v. r., venae rectae; a. r., arterise rectse. Blood-supply of kidney : On entering the kidney the renal artery bivaks up into several brandies, which pass into the ti ne proper of the organ. Branches from these arteries URINE. 129 (arteria propria renales) have two determinations (1) into the cortex and (2) into the pyramids. (1) Those branches (interlobular) which pass into the cortex divide to become the afferent vessels to the Malpighian bodies, and, after there passing through the capillary tuft, the blood is re-collected and goes out by an efferent vessel. This effer- ent vessel in its turn is broken into a minute capillary plexus which surrounds the uriniferous tubules in the cortex of the kidney, and these capillaries unite to form the venous return- circulation (interlobular veins). Thus, this system has, it is to be noted, two capillary divisions, in the Malpighian tuft, and again about the tubules of the cortex. There is also a sub-capsular capillary division, derived from certain of the interlobular vessels that do not go to nor come from the Malpighian bodies. (2) Numerous minute branches (arterise recta?) are given off, which pass into the pyramids as far as their apices, and there form capillary divisions from which the venae recta3 return to join the branches from the cortex and form the venaB proprise of the kidney. The ureters : Leading from each kidney to the bladder is a duct the ureter about the size of a goose-quill and from twelve to sixteen inches in length. The ureters are lined with mucous membrane, continuous above with that of the pelvis of the kidney, and below with that of the bladder. The " pelvis of the kidney " is the upper dilated end of the ureter ; and, in its turn, is made up of the conjoined calyces (see Uriniferous Tubules). The bladder : This, which forms a temporary receptacle for the urine, is a hollow muscular organ of a pyriform shape, lined with mucous membrane, and situated in the pelvic por- tion of the abdominal cavity. The widest part is called the fuiHJH*, and the narrowed part, whence leads the urethra, is sometimes known as the cervix. Urine : It is a clear, amber-colored fluid of slightly acid reaction. It may develop a flocculent precipitate of a light cloud of mucus upon standing. It has a characteristic odor and a salty-bitter taste. Urine has a specific gravity of about 1020 ; but under con- 9 Phys. 130 SECRETION. ditions of health it may vary from 1010 to 1030, or even be- yond these limits. ( 'otwing-matter of urine: The coloring-matter of the urine appears to be identical with hydrobilirubin. It is derived from the bilirubin of the bile, which, being absorbed, passes out of the body in the urine rather than in the faeces. The varying tints are probably due to oxidation of this sub- stance. Composition of urine : The urine is an excrementitious fluid, and may be considered as a watery solution of the excrementi- tious products of the retrograde metamorphosis of nitrogenous bodies, resulting from the processes of life and action. Chem- ically, it is a solution of urea and urates with a small per- centage of organic salts. Table of the Chemical Composition of the Urine. Water . . . 967 Crystallizable nitrogenous bodies : Urea 14 Uric acid, free (trace), Uric acid in form of alkaline urates, Hippuric acid and hippurates, !> . 11 Pigments, extractives, and mucus, (All in small and constant amounts). Salts: Inorganic Chlorides of sodium and potassium, Sulphates and phosphates of sodium and potassium, Phosphates of magnesium and calcium, Silicates (trace), Organic Lactates, acetates, and formates, which only appear occasionally. Sugar (occasionally), a trace, Gases, nitrogen, and carbonic acid. 1000 Acidity of urine : The acidity of the urine is due to the AMOUNT OF URINE. 131 presence of acid sodium phosphate. There is no free acid present, as is shown by the fact that no precipitate is formed upon the addition of sodium hyposulphite. The degree of acidity varies, being less during active digestion and less after vegetable food. Herbivora have alkaline urine, while car- nivora have strongly acid urine ; but the herbivorous animal during fasting has acid urine, because it is then living from its own tissues and is for the time a carnivore. After excre- tion, however, the urine soon becomes more acid (probably because of the presence of some fermentation), and at this time uric acid and urates may precipitate. Upon further ex- posure it is attacked by micro-organisms, and the urea is changed to ammonium carbonate, the reaction becoming alka- line, and there is a precipitation of triple phosphates and alkaline urates. In the body these conditions do not occur in conditions of health. Secretion of urine : The kidneys secrete urine in two ways : (1) by filtration and (2) by real functional action of the epi- thelium. (1) In the circulation of the blood through the Malpighian tuft there seems to be no active separation of the urinary in- gredients by cell-power, but the water and saline elements are given off here by the blood by a process of simple filtration. The amount of fluid which passes here is governed by the blood-pressure in the arteries of the kidney and by the fluid- ity of the blood. (2) The epithelium of the uriniferous tubules has secreting function, and is able to separate from the blood foreign sub- stances (e. g. 9 urea or indigo-carmine) and eject them into the tubules, and to manufacture from material taken from the blood new substances not found there (e. g. y pigments). Amount of urine : The average amount of urine secreted daily is fifty-two fluidounces, though as little as thirty-five ounces, or as much as eighty-one, may be voided within the limits of health. The quantity varies greatly in health with the amount of fluid taken, of food consumed, of the activity of the skin evaporation, and somewhat with the character of the food. In a more general way it may be said to depend upon the condition of the blood, an excess of fluids demand- 132 SECRETION. ing increase of functional activity on the part of the kidneys. In conditions of disease or under the stimulus of drugs the limits mentioned are by no means final, for in certain patho- logical conditions the secretion may be almost wholly sus- pended or very greatly increased. Conditions affecting urinary secretion : The facto is that favor the increase of urine are those conditions which favor filtra- tion of water by the glomeruli of Malpighi that is, the presentation of a larger amount of blood to the action of these bodies. This is accomplished (1) By increasing the force of the heart. (2) Through the nervous system by its action upon the vascular, so as to produce local congestion. The effect of the nervous system in increasing the urine by reflex vaso-motor impulses is felt most in the glomeruli, and the urine is there- fore very watery. (3) By conditions which cause anemia of other parts, thus producing a greater determination of blood to the kidneys, and so increasing the urinary flow. So marked is this that the skin and kidneys may almost be said to be complementary in their action in eliminating water from the system ; and in this regard their relative activity may be said to be inversely proportional to one another. Kirke gives the following table (modified from Foster), which is useful for reference : Table of the Relation of the Secretion of Urine to Arterial Pressure (Kirke). A. Secretion of urine may be increased a. Ry increasing the general blood-pressure by 1. Increase of the force or frequency of the heart-beat. 2. Constriction of the small arteries of areas other than that of the kidney. b. By increasing the local blood-pressure by relaxation <>f flu' renal artery, without compensating relaxation < /*< - where by 1. Division of the renal nerves (causing polyuria). 2. Division of the renal nerves and stimulation of the cord below the medulla (causing greater polyuria). MICTURITION. 133 3. Division of the splanchnic nerves ; but the polyuria produced is less than in 1 or 2, as these nerves are distributed to a wider area, and the dilatation of the renal artery is accompanied by dilatation of other vessels, and therefore with a somewhat di- minished general blood-supply. 4. Puncture of the floor of the fourth ventricle or me- chanical irritation of the superior cervical ganglion of the sympathetic, possibly from the production of dilatation of the renal arteries. B. Secretion of urine may be diminished ft. By diminishing the general blood-pressure by 1. Diminution of the force or frequency of the heart- beats. 2. Dilatation of capillary areas other than that of the kidney. 3. Division of the spinal cord below the medulla, which causes dilatation of the general abdominal area, and urine generally ceases being secreted. b. By increasing the blood-pressure by stimulation of the spinal cord below the medulla, the constriction of the renal artery which follows not being compen- sated for by the increase of the general blood- pressure. c. By constriction of the renal artery by stimulating the renal or splanchnic nerves or the spinal cord. Course of the urine : The urine collected in the tubules of the kidney passes into the pelvis of the kidney, and is carried to the urinary bladder in irregular quantities by the ureter. The ureters simply act as ducts, and do not store up urine, nor do they usually actively eject it into the bladder. As a few drops of urine collect in the pelvis of the kidney, they run into the bladder, the action of the two kidneys not being in alternation nor absolutely regular in point of time. Re- gurgitation from the bladder is prevented by the oblique course of the ureter through the muscular wall of the blad- der. Micturition : When the bladder is filled the act of emptying 134 SECRETION. it is called micturition. It is a voluntary act, aided by the involuntary reflex contraction of the muscular coat of the oriran itself. The voluntary muscles involved are those of respiration the diaphragm and the abdominal muscles. So far as micturition is involuntary, it is a reflex depending upon a centre in the lumbar spinal cord. Urea is a chemical body, the formula being CH 4 N 2 O (Fig. 69). The nitrogen atom is derived from the "combustion. FIG. 69. Urea, prepared from urine, and crystallized by slow evaporation (Lehmann). of proteid material. Hence we infer that the amount of urea excreted gives us an index of the amount of proteid material consumed by the body. The used-up nitrogenous matters of the body may be derived from : (1) the food (urea being greatly increased by a nitrogenous meal) ; (2) the metabolism of the tissues. The result of the " combustion " of nitrogenous material is not at once found in the blood, but exists as some antecedent of urea (probably carbonate of ammonium) until the blood reaches the liver. Under the action of the liver-cells the an- tecedent of urea is converted into urea. The urea thus formed remains in the circulation until the blood reaches the kidney ; here the urea is excreted. Thus it is seen that the kidncv- VASCULAR GLANDS. 135 cells play no part in the formation of urea, but merely exert a selective action upon it for excretory purposes. Amount of urea excreted : The amount varies, but it may be considered to be about one-half the solid constituents of the urine. Roughly speaking, the urinary solids may be regarded as 4 per cent, of the total, and the urea (including the uric acid and urates) about 1.5 to 5 per cent. This proportion is very variable, and there may be urea in healthy urine to exceed 2J per cent., or in a much less ratio than 5 per cent. Method of estimating solids : A useful rule for approxi- mately estimating the total solids in any given specimen of healthy urine is to multiply the last two figures representing the specific gravity by 2.33. Thus, in urine of specific grav- ity 1025, 25' X 2.33 = 58.25 gr. of solids in 1000 gr. of urine. In using this method it must be remembered that the limits of error are much wider in diseased than in healthy urine. Urea as a waste : Urea is capable of still further oxidation, and so would be a source of further chemical potential energy ; but urea is excreted before it is fully oxidized, and so is a real waste. Uric acid : Of the other constituents of urine, uric acid is the most important, it being one of the forms in which nitrogen is eliminated. It usually exists in the urine in the form of urate of ammonium or sodium. It probably comes, like urea, from food-disintegration. Some consider it to be an imperfectly oxidized form of urea. The amount found in the blood is especially increased in gout. Hippuric acid : This is one of the few important constitu- ents of urine which is produced in the kidneys themselves, being formed from benzoic and amido-acetic acids. It is also formed in the liver. It is another form of nitrogenous elim- ination, but may come from tissue-waste as well as food-dis- integration. It exists in slightly larger quantity than does uric acid. Vascular Glands. The vascular glands, or ductless glands, are a collection of glandular structures that possess no ducts and apparently do 136 NUTRITION. not seem to be associated in either secretion or excretion. Ductless glands include the spleen, thyroid, thymus, and tonsils. Purpose of ductless glands : No definite function has l><, frag- ment of a detached fibre not touched with acid. MICROSCOPIC APPEARANCES. 143 width. The cells are bound into bundles by an albuminous cement, and these again into larger bundles by areolar tissue. Striated voluntary muscle-tissue consists of bundles of long muscle-cells or fibres. Each fibre is completely enveloped in a sheath, the sarcolemma, and the whole bundle of fibres is bound together by a delicate connective-tissue framework. FIG. 71. A, portion of a medium-sized human muscular fibre (magnified nearly 800 diame- ters). B, separated bundles of fibrils, equally magnified: a, a, larger, and b, b, smaller collections ; c, still smaller ; d, d, the smallest which could be detached. A number of these fasciculi are joined together to make up the gross anatomical muscle. On examining the muscle-fibre itself, it will be seen to consist of alternate segments of light and dark matter, giving the fibre a striped appearance (Fig. These fibres are about an inch in length and ^poth inch in diameter. They join the connective-tissue cells of a tendon 144 MUSCLE. FIG. 72. or aponenrosis or another muscle-fibre by adhesion of the sarcolerama at the ends, and thus unite the muscle-bundles in a firm mass ; and this union is further strengthened by the cohesion of the fibres. The st nation of the heart-muscle fibres is not so marked as in ordinary muscle, and the form of the fibres is different, for they are branched and more slender. Each fibre is nucle- ated, a large oval nucleus occurring at the centre. The appear- ance of the heart-fibres indicates that they occupy an internu- diate position between typical plain and striped fibres (Fig.72). Muscle general properties : ( bear the snne relation to living muscle as fibrinogen does to blood. It' 1>\ cold \ve delay the coagulation of muscles removed from animals immediately after death, we can express a viscid fluid of slightlv alkaline reaction, known as muscle-plasma. This muM'le-plasma, when exposed to ordinary temperatures, coag- ulates much in the same way as docs blood-plasma, inyosin Muscular fibres of the heart (Quain) PHYSIOLOGY OF MUSCLE. 145 being formed. In a short time the clot will contract and squeeze out a fluid resembling blood-serum. Myosin differs from fibrin in many of its reactions. It belongs to the glob ulin class of proteids. Fat, glyeogen, organic and inorganic salts (chiefly potassium) make up the remaining 10 per cent. The fat exists as minute strips between the fasciculi. The glyeogen is stored up to serve as a supply of chemical poten- tial energy. The amount of glyeogen present in resting mus- cle is about -J- per cent. Physiology of muscle : Muscle may exist in three different conditions : those of rest, activity, and rigor. Muscle-rest: During rest a muscle has a slight but very perfect elasticity. It can be stretched to a considerable ex- tent, but always returns at once to its former condition. In the living body the muscles are always in a condition of sligkt tension, which gives mechanical advantages. Even during a state of rest the muscle takes oxygen from the blood and gives up carbon dioxide to it. The reaction of a resting muscle is neutral or faintly alkaline. Muscle during activity : The peculiar property of muscle- fibre is its contractility, which is excited by all kinds of stim- uli direct and indirect. This property is soon lost, unless the supply of arterial blood is kept up. Muscles, especially the striated, possess a certain kind of sensibility due to the sen- sory nerve-fibres which end in them. They are but slightly sensible to pain, the sensations produced being rather those of the condition of the muscle, as to fatigue, cramp, etc.; or else of muscular sense, as to position of the muscles, com- parison of weights, etc. After activity the reaction is acid, due to the development of sarcolactic acid or lactates by the breaking up of the gly- eogen in the muscle. There is also an increase in water and carbon dioxide. Only a little proteid material is oxidized during muscular activity. This is shown by the fact that after severe exercise the amount of urea excreted, although increased somewhat, is not increased commensurately with what would have been the case had the proteid furnished the chemical potential energy. 10 Phys. 146 MUSCLE Actions of muscles as levers : Most of the voluntary mus- cles in the body may be regarded as sources of power for moving the bones viewed as levers. All levers are divided into three classes, according to the relative position of the power, the weight to be moved, and the axis of motion or fulcrum. The different movements of the foot offer an illus- tration of all three kinds of levers : The first kind (Fig. 73), FIG. 73. i ii in Illustration of levers of all three orders (Huxley). W, weight of resistance ; F, fulcrum ; P, power. where the fulcrum, F, is between the source of power, P, and the weight or resistance, ,W, is illustrated when the foot is raised and the toe tapped upon the ground, the ankle-joint being the fulcrum. The second kind of lever, where W is between F and P, is illustrated when the body is raised upon the toes, the ground being the fulcrum. The third kind of lever, where P is between F and W, is illustrated when a weight is held up by the toes, the ankle being the fulcrum and the anterior group of muscles on the leg the source of power. The forearm also acts as a lever of this sort when a weight is lifted in the hand. Oxygen-supply : Muscle receives its oxidizing agent oxy- gen from the blood coursing through the vessels contained in the muscle. Even during repose the muscle takes up oxygen from the blood and gives back carbon dioxide. The muscle also stores up within itself a certain amount of oxy- gen, which can be called upon to do work even if the blood- supply be stopped. To prove this it is only necessary to cut a muscle out of the body and cause it to contract in a cham- ber of nitrogen. As a result of these contractions the muscle LATENT PERIOD. 147 throws off a quantity of CO 2 as a product of oxidation. The oxygen necessary for this oxidation comes from the supply stored up in the muscle as the only source, there being no blood-supply and the atmosphere of the chamber in which the experiment is conducted consisting only of nitrogen. During muscular activity a greater supply of oxygen is needed, and this is supplied by a dilatation of the bloodves- sels of the part. Systematic exercise of a muscle educates the arterioles supplying that muscle to remain in a condition of dilatation. This increase in the blood-supply not only provides an increase of oxygen during activity, but during repose carries an increased amount of nourishment to the muscle, with the result that the muscle increases in size and power. Hence the value of " training." Muscle-fatigue : After performing its function of contraction for a certain length of time the muscle is less active in its response to stimuli, and finally, in spite of the strongest im- pulses, fails altogether to act. This is " fatigue " of a muscle, a phenomenon with which we are all familiar. After a suit- able rest the muscle recuperates and is as active as ever. The reason for this fatigue is that during the muscle's activity a number of effete poisonous products, the result of the active katabolism, have been formed more rapidly than the power of removal by the outgoing blood. After a time the accumulation of poison is so great as to paralyze the mus- cle, so that no further activity takes place until the excess has been removed. The first effect of fatigue is seen in the increased latent period and decrease in the strength of the contraction. Muscle latent period : By latent period is meant the time that elapses after a stimulus has been applied to a muscle and before the muscle acts. We speak of the " apparent " latent period and the "true" latent period. By " apparent " latent period is meant not only the time consumed by the impulse in awakening the muscle into activity, but also the time consumed by the impulse in reach- ing the muscle (say in travelling along the nerve). " True " latent period is the actual time consumed between 148 MUSCLE. the arrival of the impulse at the muscle and the beginning contraction. The time lor FIG. 74. the true latent period is about one one-hundredth of a second. Contractility : The im- portant use of this power is to do " work " /. e., the conversion of the potential chemical energy of the muscle into heat and visi- ble motion. If one watches a muscle contract, he will see that the muscle becomes much thicker and shorter; but the volume does not change. The fact that there is no chaiiyc of volume during contraction may be proved by placing a muscle in a vessel filled with fluid to a given mark. The muscle is now made to contract, and it is noted that the level of the fluid remains unchanged. Owing to the increased metabolism during func- tional activity, there is heat liberated, hence an increase of temperature. Path of stimulus for con- traction : In the living body the impulse causing contraction of a muscle arises in the centres of brain or spinal cord, and travels thence by means of anterior nerve-roots and the somatic nerves going to the particular muscle (Fig. 74). The nerve that supplies a given mii-cle as it enters that muscle breaks up into a num- representation of cerebral and spinal motor cells with axons. 1, cerebral cell ; 2, axon ; 3, 4, collaterals ; 4', end-tufts; 5, spinal cells; 6, axon; 7, limit of spinal cord ; A7. motor-nerve; S, muscle ; 9, muscle end-plate (Kauber). CONTRACTILITY OF MUSCLE. 149 her of branches that are distributed to all parts of the mus- cle. The terminal ends of the nerve-fibres are small cellular elements called nerve end-plates or end-organs (Fig. 75). As a rule, these end-organs are scattered in great numbers throughout the muscle. By means of these end-plates the impulse from the nerve is transmitted to the muscle-fibres. Contractility an inherent property of muscle : Although under ordinary circumstances an impulse to contract is sent to a muscle through the intervention of its nerve and end-plates, nevertheless the impulse may be administered directly to the muscle itself and the result be a contraction. FIG. 75. Nerve-ending in muscular fibre of a lizard (lacerta viridis). a, end-plate seen edge- ways ; b, from the surface ; s, s, sarcolemma ; p, p, expansion of the axis-cylinder. In b the expansion of the axis-cylinder appears as a clear network branching from the divisions of the medullated fibre. Highly magnified (Kuhne). To prove that a stimulus may be administered directly to muscle-fibre with a resulting contraction the following exper- iment will be described : Liberate the sciatic nerve from the surrounding tissues in the thigh (say the right) of a frog ; but do not destroy the connection of this exposed nerve with the gastrocnemius mus- cle or the spinal cord. Pass a ligature about the right thigh so as to include all the tissues except the sciatic nerve. Tie the ligature tightly. The purpose of this ligature is to shut 150 MUSCLE. off all blood-supply to the lower part of the limb. Xow poison the frog with curare. Curare is the Indian arrow- poison, and kills by paralyzing the end-plates in all the mus- cles. In the frog experimented upon, the poison is carried to all parts of the body except to the right leg below the liga- ture. The frog lies as though completely paralyzed. Now, with an electrical battery stimulate the right sciatic nerve ; the right gastrocnemius contracts. Stimulate the left sciatic ; no result. During the stimulation of both the right and the left sciatics the nerves showed that they were carry- ing impulses (proven by a galvanometer) ; hence the lack of activity in the left leg is due to the curare having affected either the muscle itself or the end-organs. On the right side the blood carrying the curare has been prevented (by the lig- ature) from reaching the muscle or its end-plates. Now apply the electrical stimulus directly to each gastroc- nemius in turn ; both will be found to contract and with equal vigor. This experiment proves the following facts : (1) Curare paralyzes a muscle by inhibiting the end-plates of the nerves ; hence" (2) the nerve-endings in the left gastrocnemius being paralyzed, the stimulus that made the left gastrocnemius con- tract necessarily exerted its influence on the muscle direct, and not by the intervention of the end-plates. Another proof may be offered to the same end, as follows : the gracilis muscle in the frog is practically free from nerve end-organs in its lower part, and yet the lower part of the muscle contracts as readily to direct stimulation as does any other muscle. An additional proof is that the heart of the em- bryo begins to beat some time before the nerve end-organ* have developed within the heart-muscle. Artificial stimuli: A muscle may be stimulated to activity either by irritating the muscle itself, or by sending impulses to the muscle through the intervention of its motor nerve. The stimuli employed are : (1) chemical, (2) mechanical, (3) thermal, (4) electrical. The last (electrical) is the most practical and most frequently used. To study the effects of muscular contraction a myograph is necessary. A myograph consist of a drum or cylinder cov- EFFECT OF GALVANIC SHOCK. 151 ered with smoked paper. The drum is made to revolve by clockwork at a definite rate. The nerve-muscle preparation is so arranged that when the muscle contracts a lever is made to trace a curve on the smoked paper. This curve is called a myogram, and by studying the curve one can learn the amount and character of the contraction. Galvanic, faradic, and interrupted currents : A galvanic cur- rent is a continuous flow of electricity directly from the bat- tery. A faradic current consists of a direct current passing through a coil of wire ; about this coil is wrapped a second coil, carefully insulated from the first coil. The current pass- ing through the primary coil, induces a current in the second- ary coil. It is the secondary current that is applied to the nerve to be experimented upon. An interrupted or alternating current is a faradic current in which the primary current is rapidly made and broken by an automatic interrupter. Response to stimuli : It has been found that striped and unstriped muscles react to stimuli in a somewhat different manner. This difference is probably due, for the most part, to their differing structure, but may possibly be due in some degree to their differing modes of connection with the nervous system. When a stimulus is applied to a striped voluntary muscle there is an instantaneous contraction of the part irri- tated, and of that only. This contraction ceases the moment the stimulus is withdrawn. If, on the other hand, any part having involuntary muscle for instance, the bladder or in- testines is stimulated, the contraction comes on more slowly, extends beyond the part stimulated, and continues for some time, with alternating relaxation, after the stimulus is with- drawn. Effect of galvanic shock : The instantaneous application and removal of a galvanic shock to a nerve-muscle preparation result in a single twitch of the muscle. Although it is stated above that the application is " instantaneous," it is shown by experiment that the impulse is not appreciated by the nerve unless the flow of current lasts at least 0.0015 of a second. On examining the curve traced by the muscle on the myo- 152 MUSCLE. graph, there are noted : (a) the latent period i. e., the time elapsed after the application of the current before the con- Base-line. Diagram of muscle-curve, a, point of application of current ; b, point of beginning contraction ; c, maximum ; d, return to normal. traction commenced ; (b) a gradual rise in the curve until the maximum is reached, and then a gradual curve down to the base-line (Fig. 76). Fio. 77. Opening shock. The time a-b is the latent period, and in the case of a frog's rust rocnemius is about y^th of a second. The total curve in the same muscle lasts about -t^th of a second. If, instead of applying and removing the galvanic current in close succession, the current be allowed to flow for some INTERRUPTED CURRENT. 153 time through the nerve-muscle preparation, two contractions and relaxations are noted. One contraction and relaxation takes place when the current is applied, but during the rest of the flow of the current the muscle remains in a state of repose. The second contraction and relaxation occurs when the current is broken. The ' contraction at the "make" of current is greater than that at the " break " (Figs. 77 and 78). From the fact that the muscle remains quiescent during the flow of the current after the first shock, the theory is deduced that it is not the change in state, but the rapidity of FIG. 78. the change in state, which irritates the muscle into con- traction. This is proven by the fact that a large amount of electrical cur- rent can be poured into a nerve-muscle preparation without exciting a contrac- closing shock, tion, provided the current be gradually applied and very slowly increased. On the other hand, a small amount of electricity if rapidly administered will produce a sharp contraction. Induced current : A single shock from an induced current produces the same result as that of a galvanic current of in- stantaneous duration. A prolonged application of the induced current produces the contraction on the " make " and a contraction on the " break " similar to that of the galvanic current, except that the " break " contraction is greater than the " make " contraction. Interrupted current: The interrupted current, consisting as it does of rapidly alternating " makes " and " breaks," produces a series of rapidly succeeding shocks on the nerve- muscle preparation. The result is that the muscle, after the period of contraction is over, has no time to relax before a second shock is received, which tends to a fresh contraction. Hence the muscle remains in a state of continuous contraction until the stimulus is removed or the muscle wears out. This condition of continuous contraction is called tetanus (Fig. 79). 154 MUSCLE. 7 Muscle electrical state : By a rise in electrical state is meant an increase in the height of the curve in succeeding contractions, on the application of single shocks. It is as though the muscle became more FIG. 79. sensitive and more responsive to the second shock than to the first, and more so to the third than to the second, etc. This rise in elec- trical state is observed for the first six to ten shocks. It is as though the muscle "limbered up " in the first few shocks by a little preliminary exercise until it reached its power of doing its best. Currents of rest : When a living curve of tetanus. muscle is tested by means of the galvanometer after removal from the body, it is found to develop certain electrical currents known as muscle-currents or currents of rest. They are strongest from the centre of the muscle toward the cut end, though certain minor currents are developed with the elec- trodes in closer proximity. The cut ends of a muscle are always electro-negative to its equator. This phenomenon cannot be observed in uninjured muscle when in the bodv, but any injury will render the injured portion electro-nega- tive to the rest of the muscle. This condition ceases with the power of contraction, and cannot be demonstrated in dead muscle. When the muscle is made to contract, the galvanometer- needle, which has indicated the passage of an electri- cal current during rest, flies quickly back toward the /ero, indicating the cessation of the current of rest. This action is known as the negative variation of the galvanom- eter, and as soon as the contraction of the muscle has ceased the instrument again indicates the presence of the current of rest. The causes are not yet fully determined, but these cur- rents are probably due to chemical changes resulting from physiological degeneration. It has been held that such PFLUGER'S LAW OF CONTRACTION. 155 currents occur naturally in muscle as the result of certain of the cells exciting electro-motive forces, but the former theory seems the more plausible. Excitability and conductivity : These terms, although similar as to the results, really mean very different things. Excitability, or irritability, refers to the activity of response a nerve may show to a given electrical shock ; whereas conduc- tivity means the power of one part of a nerve to conduct an impulse generated at another part of the nerve. A moderate current while passing through a nerve pro- duces no shock, but the excitability of the nerve at the point where the kathode is placed is increased (this increase at the kathodic area is called katelectrotonus) ; at the same time the excitability of the anodic area is lessened (anelectrotonus). On removing the current the excitability at the kathodic area falls, and the excitability at the anodic area rises. Ou the other hand, during the flow of a moderate current there is a marked diminution in the conductivity of the nerve at the kathodic area, even to the extent of rendering the kathodic area impervious to conduction, without seriously affecting the anodic area. On removing the moderately strong current the nerve instantly returns to its normal conductivity. If very strong currents be used, conduction is completely lost at both the anodic area and the kathodic area during the flow of the current ; and when the current is removed the nerve does not at once return to a normal state of conductivity, but remains impaired for a little while. Pfluger's law of contraction : It has already been shown that the stimulation of a nerve-muscle preparation by a galvanic current will result in a contraction only at the "make" and at the " break " of the current ; this is true only if the current be of medium strength, regardless of whether the current be " ascending" or "descending." But the results are different if the current be very weak or very strong. The following table, known as Pfluger's table, shows the effects at a glance : 156 MUSCLE. Current-strength. Ascending current. Descending current. "Make." "Break." " Make." "Break." Weak current .... Medium current . . Strong current . . . Contraction. Contraction. Contraction. Contraction. Contraction. Contraction. Contraction. Contraction. Ascending and descending currents : By an ascending cur- rent we mean a position of the electrodes such that the elec- trical flow is from the periphery (muscle) to the centre (brain) i. e.j the anode or positive pole is nearer the muscle and the kathode or negative pole is nearer the nerve-cell. The de- scending current is the reverse and the poles are transposed. The explanation of the results is simple, if we hear in mind the' effects on the nerve of the various strengths of currents. A galvanic current is stronger on the " make " than on the " break." Hence if we reduce the strength of our current so that the " break " shock is so weak as to fail to give any result, our " make/ 7 being somewhat stronger, would still influence the muscle so as to cause a contraction. This, too, would be regardless of the direction of the current. This explains the first part of the " law," namely, that regarding weak currents. In explaining the second part of the law, it is necessary to bear in mind that the nerve-vibrations start from the region of the kathode on closing the current; but on opening tin- current the nerve-vibrations start at the anodic area; also to bear in mind that moderate currents diminish the conductivity at the kathodic area without affecting the anodic region, anil on removing the current the nerve at once returns to normal. So we find on closing a descending current of moderate strength, the excitation starting at the kathode can readily pass to the muscle and produce a contraction. On opening the current, the excitation starts at the anode and readily travels along the nerve past the kathodic area (which, on opening the current, returned to normal conductivity), and so on to the muscle. If we take an ascending current, on closing, the impulse NERVE-MUSCLE PREPARATION. 157 goes from the kathode down the nerve, past the unaffected anodic area, to the muscle. On opening the current, the im- pulse passes from, the anode directly to the muscle. For the third part of the law we must remember that very strong currents reduce conductivity at both anode and kathode, and on removing the current the loss of conductivity does not at once disappear. So, to study it in detail, let us assume a strong ascending current. On making the current we re- move the conductivity of the anodic area, and the impulse starting at the kathode cannot travel by the impervious anodic area, and hence no contraction results. On opening the current the impulse from the anode passes to the muscle and a contrac- tion follows. With a descending current, we find on closing that the impulse starting from the kathode travels to the mus- cle unhindered and a contraction results ; but on opening the current the impulse starting at the anode travels along the nerve until it reaches the area where the kathode was. This area the impulse cannot pass (for the conductivity does not return for some time) and no contraction results. Position in the body: During repose a muscle is not in a state of complete relaxation, but in an intermediate condition between relaxation and contraction. The object of this is to save time ; for the muscle can at once exert its power, as it has no " slack " to gather in before it exerts its pull. Also there is no jerk or jar in the motion. Results of contraction : (a) heat ; (6) motion ; (c) sound ; (d) change of shape ; (e) fatigue ; (/) chemical changes ; (f the individual. In robust, healthy people >ii(lleuly killed, the muscles may live for six or eight hours longer. All the intermediate stages have been noted. The death of the muscle is characterized by a contraction, fixing MEDULLATED FIBRES. 159 the limbs and body in a rigid condition. This post-mortem rigidity constitutes rigor mortis. The muscles of the jaw and neck are usually first affected, and then the arm, trunk, thighs, and legs in the order named. All muscles are affected, both the voluntary and involuntary. The condition of rigor mortis lasts from ten to twenty- four hours, and passes off in the inverse order from its ap- pearance. First the legs relax, then the thighs, etc., and lastly the jaw. Cause of rigor mortis : Rigor mortis is caused by a fermen- tative change which produces a coagulation of the blood- plasma in the muscles. During this coagulation-process there is a development of heat, which causes a rise in the temperature of the body. It may amount to 5 or 10 F., or even more. Cause of disappearance of rigor mortis : After the fermenta- tive change which causes the coagulation of the muscle- plasma has ceased, a putrefactive process is set up which destroys the coagulum and the muscles become soft and flabby. NERVOUS SYSTEM. Fibres and Cells. The nervous system is an aggregation of tissues so arranged as to adjust the workings of all the parts of the body to one another and to suit the body to its environment. The elementary tissues of the nervous system are of two forms : nerve-fibres and nerve-cells. The nerve-fibres are of two kinds : (1) medullated or white fibres ; (2) non-medullated or gray fibres. The fibres are united in bundles to form nerve-trunks or " nerves/' The cells are in groups to form nerve-ganglia, but nerve- fibres are also found in the ganglia. Medullated fibres consist of (1) an external nucleated sheath, or neurilemma ; (2) an inner protective medullary sheath (the white matter of Schwann) ; and (3) internally the axis-cylinder (Fig. 80). 160 NERVOUS SYSTEM. The neurilemma is a pellucid structureless mem- brane. Within are seen at inter- vals nuclei surrounded by more FIG. 80. or less protoplasm. These nuclei and their protoplasm are relics of embryonic cells. The medullary sheath is a thick fatty semifluid substance. It is this substance which produces the peculiar white appearance of some nerves. Some authorities have claimed that it is made up of a fine network, in the meshes of which is embedded the bright fatty material. The axis-cylinder consists of a large number of primitive fibrill* Medullatcd ncr " librc . which vary considerably in size, A, node of but on the average may be said to be about Y^j-ytn inch in diam- eter. There is little doubt that the axis-cylinder is the essential part of the fibre, the other parts being merely for support or pro- tection. Nodes of Panvicr: Here and there at short intervals along the course of medu Hated fibres are found notches or constriction in the medullary sheath. These con- strictions are called the nodes of Ranvier. At these points there is no loss of continuity of the axis- cylinder nor of the neurilemma, but the medullary sheath is deficient and allows the neurilemma to dip in and touch the axis-cylinder. The nodes apparently divide the nerve-fibre off into histological units, for there is a nucleus for each segment marked off by the nodes. As the medullated nerves near their terminations, B, nucleus beloiii:in to the neurileinma : ('. axis-cylinder ; J'. neu- rilemma rendered dis- tinct by the retraction of the myelin of the medullary sheath. In the right-hand figure the clefts of LimtLT- mann arc shown as while lines in the dark myelin. The fi^r- arefl are taken from spe- cimens treated with osmic acid, which colors the tatty con- stituents of the myelin a dark brown or black (Key and Retzius). FUNCTION OF NERVE-FIBRES. 161 FIG. 81. the medullary sheath disappears and the axis-cylinder con- tinues with the neurilemma ; but this latter also disappears before the final ending of the fibre (axis-cylinder) in the tissues. The fibre then splits into two or more terminal branches. Thus white fibre becomes non-medullated fibre. Non-medullated fibres: They consist of the axis-cylinder alone, without the medullary sheath. They do not differ in any other regard from the white fibres. When collected in bundles to form nerves they have a yellowish or grayish color. They are found in the olfactory and auditory nerves and in the nerves of the sympathetic system, and they occur in greater or less number in the nerves of the cerebro-spinal system. In size these fibres are about one-third to one-half the diameter of the medullated. They are sometimes spoken of as the fibres of Remak (Fig. 81). Formation of nerve-trunks : To build up a nerve-trunk, whether from medul- lated or non-medullated fibres, the fibres are joined in bundles which are enclosed in a thin fibrous sheath (perineurium), and these bundles of nerve -fibres are bound in a firm connective-tissue which serves to protect and to unite them strongly. So far as we can see, the individual nerve-fibres, as a rule, are continuous and independent from their origin in the nervous centres to within a short distance of their peripheral termi- nation. When a nerve divides into several branches, or when adjacent nerves communicate, it is because certain fibres leave those with which they are associ- ated and pursue a different course. There is no real union of nerve-fibres. Function of nerve-fibres : The function of a nerve-fibres is the transmission of a stimulus. The axis-cylinder connects the centre and pe- riphery cells, and conveys between them the stimuli. This 11 Phys. Fibres of Remak; mag- nified 300 diameters. With the gelatinous fibres are seen two of the ordinary, dark bordered nerve-fibres (Robin). 102 NERVOUS SYSTEM. transmission for any particular fibre is in one direction only. Depending on the direction the impulse is carried, the nerve-fibres are classified into afferent (or centripetal), effer- ent (or centrifugal), and intercentral fibres. The former are those by which impressions are taken from the periphery to the brain, and are commonly called sensory fibres. Conduction in these nerves may cause (1) a sensation as of pain, heat, etc. ; (2) special sensation ; (3) reflex action of some kind ; or (4) inhibition or restraint of action. The second class conduct stimuli to the periphery, and are known as motor fibres. Conduction in these nerves may cause : (1) contraction of muscles (motor nerves) ; (2) control of nutrition (trophic nerves) ; (3) control of secretion (secre- tory nerves ; or (4) inhibition, augmentation, or checking of other efferent impulses. The last class includes nerve-fibres which connect more or less distinct nerve-centres, and may therefore be said to have no peripheral distribution. Nerve-fibres are mere conductors of impressions. An impulse started in any fibre is trans- mitted unchanged to its termination without being imparted to any of the fibres lying near it. Nerve-cells : As has been shown, the nerve-fibres are the conveyors of impulses, either from the brain to the periphery, or from the periphery to the brain. It is now in order to study the termination or starting-points of these impulses. Every nerve-fibre starts or ends in a nerve-cell, which is the origin of the impulse in the case of an efferent nerve-fibre, or the recipient of the sensation in afferent nerves. Nerve-cells or ganglion-cells present a great variety of shapes, and yet have common characteristics. The cell-body is granular, and contains a large nucleus which contain- a prominent nucleolus. The cells have at least one pncos, and often more (Fig. 82), and the cells are classified as uni- polar, bipolar, or multipolar. These processes are of two kinds one kind dividing and subdividing (branching or proto- plasmic processes ; or dendrites) until they become very deli- cate and seem to interlace with, but without joining, the NERVE-ENDINGS AT PERIPHERY. 163 equally fine processes from other cells ; another kind (axis- cylinder processes or axons) pass on without division and be- come axis-cylinders of medullated nerve-fibres. The nerve-cells vary greatly in size, and are very diverse in form, but the presence of a nucleus, a nucleolus and the processes is characteristic of nerve-cells. They may be FIG. 82. Is from the anterior horn of the gray substance of the spinal cord. enclosed in a delicate capsule, which becomes continuous with the netiri lemma (Fig. 83). Nerve-endings at periphery : Efferent nerves, as has been stated previously, on nearing their terminations undergo a loss of the neurilemma and the sheath of Schwann, the axis- cylinder itself splitting into several subdivisions. Each of these subdivisions ends directly in the substance of a cell be- longing to the organ supplied, by means of end- plates. Sensory nerves " ending" in the skin find their way to or take origin from certain bodies (sense-organs) which are essen- tial to the conduction of the sensory impression to the central nerve-ending. These sense-organs are of several kinds. In NERVOUS SYSTEM. the fingers and toes are found two kinds of sense-organs which may be especially mentioned : 1, touch-corpuscles FIG. 83. Nerve-cells, from spinal and sympathetic ganglia of man, enclosed in their cap- sular sheaths from hardened preparations (Key and Rctzius). (Fig. 85); 2, Pacinian corpuscles (Fig. 84). The exact anatomy and physiological use of these bodies are still some- what obscure; and, indeed, the whole subject of sensorv nerve-terminations is but ill understood. We may regard the fibres of sensory nerves, as a majority, as forming a minute plexus in the corium and to terminate in sense- organs in a way not always known. Some of the speeial sense-organs are possessed of nerve-endings which are more clearly observed. Neuron : By the term neuron (Fig. 80) is meant a nervous NATURE OF NERVE-JM PULSE. 165 entity that is, a nerve-cell, its axis-cylinder process, dendrites, and the terminals. Nature of nerve-impulse : The nature of the impulse that travels along the nerve is hard to determine. Nerve-fibres FIG. 84. FIG. 85. A, tactile corpuscle ; b, nerve (Quain). may be stimulated by any- thing which increases their irritability with sufficient suddenness, but they cannot of themselves originate such a condition. The stimulus produces its effect upon the nerve-termination. The re- sult, therefore, of any stimu- lation of a nerve depends solely on the character of the end-organ, and not upon the character of the stimulus. Artificially in a nerve-mus- cle preparation the impulse can be aroused by four meth- ods : chemical, thermal, electrical, and mechanical means. The impulse passing along the nerve is not chemical, be- cause there is no heat liberated ; nor is there an expansion of nerve-fibre, even after prolonged activity, and naturally one would expect heat and expansion as the result of chemi- Vater's or Pacini's corpuscle, a, stalk ; b, nerve-fibre entering it ; a, d, con- nective-tissue envelope ; e, axis- cylinder, with its end divided at / S (Quain). 166 NERVOUS SYSTEM. Fro. 86. Neuron with long axon proceeding as an uxis-cyliiiderof a nerve-fibre, n, nerve- cell proper ; r/,dendrites ; x, axon ; d. g., dendrite showing gemmulee; c, collutt r als; /. nui tut'ts or terminals. Pyram- i, spinal nerves of tin- right and left sides; (I, origin of the anterior rout : e, origin of the posterior root ; c, ganglion of the posterior root.' these spinal nerves is made up of an an- terior and a posterior root (Fig. 89), of which the latter is the larger. The ante- rior root arises between the anterior and lateral white columns, the posterior be- SPINAL CORD. 175 tween the posterior and lateral columns. On each pos- terior nerve-root is found a ganglion immediately beyond its point of emergence. The function of this seems to be trophic. The white substance is arranged externally to the gray in each half of the cord, and is so disposed as to be conve- niently divided for purposes of description into three columns, known respectively as the anterior, lateral, and posterior columns of the cord. There is also a thin band of white substance at the base of the anterior median fissure (the white commissure). The gray matter fills in the central portion of the cord, and is variable in its amount, the calibre of the cord at its enlargements being increased by the in- crease in the amount of gray matter at these points (Fig. 88). The white substance will be noticed to diminish quite regu- larly in the sections of the cord from above downward, as seen in this series. The gray substance is not completely halved by the anterior and posterior fissures of the cord, but is continuous across the mid-line (gray commissure) ; and in it at the centre is a minute canal communicating with the ventricles of the brain. The gray matter is more abundant between the lateral and anterior and between the lateral and posterior columns of the white substance ; the names anterior and posterior horns (cornua) are given to these regions re- spectively. Spinal cord microscopic examination: Under minute ex- amination the white matter is found to be made up of medul- lated nerve-fibres, which collect to form the anterior and posterior nerve-roots, and communicate with other regions of the cord. The gray matter contains multipolar cells of varying size and shape, with axis-cylinders and " branching 77 processes lying in the neuroglia (" connective tissue "). The multi- polar cells are some of them quite large. In the anterior horn of the gray substance the axis-cylinder-processes of the nerve-cells connect directly with fibres forming the anterior nerve-roots (Fig. 90) ; but in the posterior cornu the com- munication is indirect i. e., through the branching processes and the divided axis-cylinders of the collaterals of the pos- 176 NERVOUS SYSTEM. terior nerve-roots, forming thus by contiguity what are known as end-arborization^. Origin and course of nerve-roots : (a) Anterior nerve-roots are derived from (1) the anterior columns of the cord, but some fibres come through the commissure from (2) the oppo- site side, and some come from (3) the lateral tract. Still other fibres arise from (4) the multipolar cells in the anterior cornu of the gray matter. The fibres of the anterior nerve- Fio. 90. Transverse section of the spinal cord in man (lumbar region). roots are efferent i. e., they carry impulses from the centre (brain) to the periphery. (/>) Posterior nerve-roots give off collaterals which enter the posterior horn of the gray matter, and the ultimate fibres break up and form "indirect" communication with the branching processes of the large multipolar cells, but some fibres cross through the gray commissure to the opposite side. The fibres of the posterior roots arc afferent i. e., they cnrry impulses from the periphery to the centre in the cord, medulla, or brain. The course of all the fibres through the cord itself has not been determined with absolute accuracy. Certain fibres, how- TROPHIC CENTRES OF NERVE-ROOTS. Ill ever, have been traced through their length, notably the fibres of the pyramidal tracts, in the anterior and lateral columns. It has been calculated that only about one-half as many fibres enter the spinal cord from the brain as leave it through the nerves ; therefore it must follow that some fibres originate from the corcl. The increase in gray matter in the cervical and lumbar enlargements, where the fibres for the large plex- uses of the nerves (branchial and lumbar) are given off, con- firms this view. Spinal cord recurrent sensibility fibres : Certain afferent fibres in the nerve-trunks of the body do not pass directly into the posterior nerve-roots and so to the spinal cord, but take an erratic course as follows : After leaving the nerve-trunk these fibres, known as the recurrent sensibility fibres, pass into the anterior nerve-root, being mingled with the efferent fibres fora short distance ; then these fibres double on their course, return to the point of junction between the anterior and posterior nerve-roots, to follow by way of the posterior root the normal course to the cord. These fibres form but a small proportion of the afferent fibres, and the purpose of their erratic course is not easily explainable. Trophic centres of nerve-roots degeneration : The posterior nerve-roots seem to be dependent upon the ganglia which are found upon them for trophic influence. The anterior root in a similar way seems to depend upon a trophic centre in the gray matter in the anterior horn. Division of a nerve is followed by a degeneration or break- ing down of the axis-cylinders of its fibres within a day or two, the loss of function being an earlier and immediate manifestation. This degeneration is centrifugal that is, does not proceed toward the spine, but to the periphery. If the posterior root be cut, however, between its ganglion and its emergence from the cord, the degeneration is toward the cord i. c., centripetal and the nerve beyond the ganglion does not degenerate. The anterior root cannot, however, be divided at any point beyond its emergence without centrifugal degeneration of the fibres (Fig. 91). The regeneration takes place slowly if the continuity of the nerve is at once restored, and may even follow after the nerve has degenerated for some 12 Phys. 178 NERVOUS SYSTEM. months and a complete loss of function has affected the part supplied by it. The fact that the axis-cylinders are restored only in this way is of interest, as showing the influence of the trophic centres on the nerve-growth. FIG. 91. Degeneration of spinal nerves and nerve-roots after section. A, section of nerve- trunk beyond the ganglion ; B, section of anterior root ; C, section of posterior root ; D, excision of ganglion ; a, anterior root ; p, posterior root ; g, ganglion. The functions of the spinal cord may be tabulated as fol- lows : (a) Conduction, or carrying impulses between centre and periphery ; (b) Transference i. e., an apparent transferring of im- pression from one set of fibres to another ; (c) Reflex action i. e., the origination of action in response to stimuli from the periphery ; (d) Augmentation, or a resulting effect in excess of the exciting cause; (e) Automatic acts, from corresponding centres ; (/) Coordination, or the adjustment of the workings of different parts of the body to one another; (g) In hi /tit ion of reflex acts. Conduction : Than by the spinal cord there is practically no other nervous communication between the brain and the musculo-cutaneous system; hence through it must conic nil the nerve-impulses which pass to or from the brain. In other words, every sensory impulse that is felt and every motion that is willed, perception and volition being attributes CONDUCTION. 179 of the brain, must be conducted through the nerve-fibres of the spinal cord to the brain, and vice versa. The student must not suppose that in all instances a con- tinuous nerve-fibre runs from the central cell in the brain to the peripheral end-organ, and vice versa. The nerve-fibre is often broken at one or more places along its course, the end of the fibre terminating in a brush-like ending which " articu- lates " with the dendrites of a secondary or relay cell. The impulse is thus communicated to the second cell, which in turn transmits the impulse along its own nerve-fibre and so on. The conducting fibres in the cord lie chiefly in the white matter; whereas the gray substance represents the cell-ele- ments as well. The conducting fibres have definite positions in the cord, and each portion has fibres which always conduct the same kind of impressions. Most of the motor fibres cross to the opposite side in the medulla oblongata (decussation of the pyramids), and the im- pulses pass down by the lateral columns in the crossed or lat- eral pyramidal tract on the side opposite to that in which they originate. There is also a set of motor fibres which does not cross, but passes directly to the same side in the anterior col- umns, and decussates in the anterior or white commissure near the point of distribution. The destination of these fibres is variable, for the reason that the amount of decussation in the medulla is not constant ; but, as a rule, the fibres in the direct tract go to the upper portion of the body. The cells in the anterior cornu of the gray matter of the cord originate many of the fibres which go to the nerves. This is demonstrated after division of the cord by stimulating these fibres : a series of coordinated motions follows, and this stimulus may be ap- plied direct to the fibres or through the sensory nerves. The course of the sensory fibres (Fig. 92) is somewhat problematical, but certainly these impulses enter the cord by the posterior nerve-roots. The fibres conducting them break up as described, and cross to the opposite side of the cord through the gray commissure. It is probable that after decus- sating the fibres communicate with multipolar cells, and thus 180 NERVOUS SYSTEM. pass on as white fibres in the lateral columns. These fibres after entering the lateral columns (of the opposite side) pass to the medulla as a distinct tract the antero-lateral ascending FIG. 92. Diagram showing pathway of the sensory impulses. On the left side SS' represent afferent spinal nerve-fihres ; C, an afferent cranial nerve-fibre. These fibres ter- minate near central cells, the neurons of which cross the middle line and end in the opposite hemisphere (Van Gehuchten). tract at the periphery of the lateral, extending into the anterior column. REFLEX ACTION. 181 It is by this tract that sensations of pain and of tempera- ture are supposed to pass. There are also afferent fibres in the posterior columns the posterior median by which the sensations of touch and weight (or muscular sensation) are be- lieved to pass ; the latter, however, does not decussate. To recapitulate : sensations of pain and temperature are trans- mitted through the lateral columns, and those of touch and weight through the posterior columns. Transference : Impressions conveyed by a centripetal nerve- fibre, it has been stated, pass uninterruptedly throughout its whole length without being communicated to adjacent fibres. When such an impression reaches a nerve-centre in the cord, it may seem to pass to another fibre or set of fibres, so that the pain or other sensation is felt in an entirely differ- ent part from that in which the stimulus started. At the same time the primary impression may also be conducted to the brain, so that the sensation seems to come from two different places. This phenomenon, known as transference, cannot be well explained, yet instances of it are very frequent. In dis- ease of the hip the pain is frequently felt in the knee alone, yet the latter may be perfectly sound. A biliary calculus will often cause pain in the glans penis. Reflex action : Certain sensory stimuli are received, pass up the nerve-trunks to the posterior nerve-roots, and thus to the spinal cord. In the cord the impulse may be sent to the brain, producing the consciousness of pain, etc.; or else the impulse in the cord may be transmitted directly to some motor cell in the anterior horn, stirring it to activity, with the result that there is some muscular action. For example, a person may be tickled with a feather : the subject brushes away the offend- ing object either with or without consciousness of what he is doing. If he does it unconsciously, the reflex act takes place in the cord ; if he brushes away the feather as a result of the impression received in his brain, the reflex act took place in the brain. Furthermore, a person may perform a reflex act through the reflex centres in the cord, and yet after the act is completed he may receive the sensory impulse in the brain. From this we may deduce the following rule : 182 NERVOUS SYSTEM. A reflex act taking place through the cord, may or may not be accompanied by conscious sensation ; but if it is accompa- nied by conscious sensation, the consciousness is always later in point of time than the act. Thus we may say a reflex ac- tion is an action which results from a centripetal nerve-im- pulse passing to a nerve-centre in a ganglion, and there trans- forming to a centrifugal impulse passing to a muscle. Such an action may be simple and involve a single muscle, or com- plex and involve many : thus, a ray of light falling upon the retina causes a simple reflex contraction of a single muscle, and the iris contracts. As an illustration of a complex reflex action, however, irritation of the larynx causes not only a closing of the glottis, but a contraction of all the muscles in- volved in forced expiration or coughing. Reflex acts are more noticeable in the cerebro-spinal sys- tem ; but they may belong to either, or may be mixed, the impulse going by the one system and returning by the other. Examples: sneezing, coughing, swallowing are cerebro-spinal reflexes ; the vaso-motor reflexes are largely sympathetic, but the centripetal nerve is often cerebro-spinal, as in the secre- tion of saliva or in blushing. The spinal cord in man is so much under the control of the higher centres that its capabilities for reflex action nro often overlooked. After injury to the spinal cord the reflex acts are apt to be purposeless and fruitless. In many lower animals reflex actions, after the cord has been divided, are often followed by extensive and coordinated movements. In the frog this is especially marked. Yet the difference is one of degree only. In man many minor acts are per- formed as reflex movements occurring through the aid of the spinal cord, although the cord is incapable of initiating them of itself. Special reflexes of the cord : Under this heading we may enumerate the micturition, defecation, and genital centres : (1) Micturition: Here the sphincter of the bladder is kept in a normal state of contraction, until such time as afferent impulses are sent to the cord from the neck of the bladder owing to pressure of urine; an efferent impulse is then aroused which releases the sphincter and at the same time SPECIAL REFLEXES OP THE CORD. 183 FIG. 93. causes the wall of the bladder to contract (Fig. 93). though this act is a reflex one, it is under the control of the will to a large extent. (2) Defecation : The nature of the act is identical with that of micturi- tion : the impulse arising from the presence of flatus or faeces, with the re- sultant relaxation of the sphincter ani and contraction of the lower bowel. (3) Genital: Controls the erection of the penis and the ejaculation of semen. Under this heading we may also in- clude the centre for parturition. Stimulation of the interior of the uterus by its contents may excite the centre enough to cause it to send out impulses producing uterine contrac- tions and expulsion of its contents. That delivery can take place in a person under the influence of chloro- form shows that the centre is inde- pendent of the will. As in the case of micturition or defecation, there is an accessory action of the abdominal muscles, for the most part reflex and involuntary. Al- Schema of micturition. AC, EC, C, automatic, reflex, and cerebral centres; B, bladder ; S, sensory centre acted on by afferent im- pulses (Landois). All these reflexes have their seat in the lumbar enlarge- ment of the cord. Varieties of reflex acts : Reflex actions may be simple or coordinated, primary or secondary. In the simplest reflex act only a single nerve-cell with an afferent and an efferent fibre is concerned. In most reflexes, however, it is probable that several cells are involved and act in concert. Primary reflex acts are those such as sucking, which the infant can perform as well as the adult. Secondary or acquired reflexes are those which require an effort of the will for their first per- formance, but which become habitual after frequent repeti- tion. They are described under Coordination. 184 NERVOUS SYSTEM. Augmentation : The resultant reflex act of a stimulation may be far in excess of the amount of stimulation received. For example : a tiny drop of acetic acid applied to the skin of a decapitated frog may result in most frantic efforts on the part of the animal to rub off the offending fluid, although the irritation can be but slight. The explanation lies in the fact that the sensory nerve- fibres, as they enter the cord, in- dividually break up into an ascending and a descending limb. These two limbs in turn give out, at several points along their course, collaterals, which break up into sets of brush- endings. These " brush-endings " connect with the demlrites of motor cells. Thus it is seen that one sensory fibre may stir into activity several motor fibres, so that the resultant act is augmented. Automatic acts : In normal subjects the cord is not tho scat of many automatic acts. But if the brain be destroyed or severed from the cord, certain centres which normally are secondary assume primary importance and are capable of au- tomatic activity. The best examples are the vaso-motor centres of the cord. Under the capitulation of normal automatic coifrcx we may include those governing the sweat and general mus- cular tone. The student must bear in mind that there are many auto- matic acts whose centres are outside of the cord, their origin being in local centres. Thus, there are certain actions which continue, and, while they are closely related to reflex action, do not seem to be true reflexes, but to originate in the part. Thus, the peristaltic action of the alimentary tract is not de- pendent upon the presence of food in the intestines, but may be excited in the absence of food or checked when it is pres- ent. This action has been referred to small ganglia and nerve-plexuses found there (Auerbaeh's and Meissner's), and is considered to originate in the local nerve-centres. This is what is known as automatism or automatic nerve-action. Coordination: The coordination of the cord is nothing but a repetition of ordinary reflex acts for our daily lives. F<>r example, we look upon walking as a coordinate movement. It is performed unconsciously and depends on reflex activity GROSS ANATOMY. 185 resulting from cutaneous and muscular sensations, or the sense of position in space. Inhibition of reflex acts : This function does not originate in the cord, but in the brain ; but the transference of the impulse takes place in the reflex centres of the cord. It is the ability to control or modify reflex action by an effort of the will or by mental action which is not consciously voluntary. As an ex- ample of this, if the palm of a sleeping child be touched by the finger, the baby's hand will grasp the finger ; but if the child is awake, no such reflex occurs : it is checked by mental action. Again : one may avoid crying out when in pain by an effort of the will, or may hold the feet still when the soles are tickled. An example of this inhibition of reflex action is offered in the knee-jerk, an important sympton in the diag- nosis of certain nervous diseases. When the patellar tendon is tapped sharply with the tips of the fingers the foot and leg are suddenly jerked forward. If the patient is ignorant of what is to be done, the amount of motion obtained is diag- nostic ; but if he knows what is expected, the result is apt to be deceptive, as this reflex can be entirely prevented by the will. For this reason, if the patient knows the symptom, it is always tried when he is off his guard. The object of reflex acts is to save time, suffering, and wear and tear on the body ; but if we had no check on reflex activities, we should spend all our time making efforts to escape outside stimuli. Medulla Oblongata. Gross anatomy : The medulla oblongata, or bulb, is a column of white and gray nerve-substance, the lowermost portion of the brain, and connecting with the spinal cord below. The white substance is composed of the medullary fibres continued up from the cord, and the gray matter is arranged variously between the bands of white fibres. It has an an- terior and a posterior fissure, corresponding to those of the cord, and the central canal of the cord here opens into the fourth ventricle. The medulla continues in a general way the arrangement 186 NERVOUS SYSTEM. of the fibres in tracts of the cord below ; but as the diameter is greater, the general shape is pyriform and the shape of the columns is changed (Fig. 94). The anterior columns of the cord correspond in position to the pyramids of the medulla ; the posterior columns, to the restiform bodies ; and the lateral columns correspond to the lateral tract of the medulla with the olivary bodies. The fibres, however, do not follow this FIG. 94. Middle peduncle of cerebellum. Medulla oblongata and pons Varolii, anterior surface. arrangement so closely, but are distributed variously in the medulla. The anterior columns of the cord send some of their fibres (the direct pyramidal tract) into the pyramids, forming con- tinuous tracts. Fibres from the lateral columns also enter the pyramids, but cross in bundles to the opposite side. These fibres may be seen crossing the anterior fissure between the pyramids by gently separating them. This is known as the decussation of the pyramids. The fibres which cross in this way belong in the cord to the portion of the lateral column known as the crossed or lateral pyramidal tract. MEDULLA. 187 Further course of the fibres of the pyramids : Almost all of the fibres pass on through the pons Varolii and crusta of the mid-brain, to be distributed to the cortex of the hemispheres by way of the internal capsule and corona radiata. A few of the fibres (external arciform) pass to the cerebellum by way of the restiform body. Lateral columns : The lateral column of the cord is broken into three tracts in the medulla : one, we have just seen, joins the anterior pyramid of the opposite side by the decussation ; a second joins the restiform body on its way to distribution in the cerebellum ; while the third set of fibres passes through the deeper part of the medulla (formatio reticularis) and tegmentum of the mid-brain, and reaches the ganglia at the base of the brain ; some of these go to the fillet. Posterior columns : The fibres of the posterior column con- tinue on as the funiculi cuneatus and gracilis of the medulla. These in turn terminate in their own nuclei, from which new fibres arise and pass to the fillet of the opposite side. The restiform body, just above the ends of these funiculi, for the most part reaches the cerebellum. 1 Olivary bodies : Each is a mass of white nerve-substance containing a central gray nucleus. There are communications between it and some of the tracts from the cord, especially from those tracts of the anterior and lateral columns which go to the ganglia at the base of the brain. Medulla arrangement of the gray matter: As the fibres which form the crossed pyramidal tract pass from the lateral column to decussate into the pyramid of the opposite side, they push the anterior cornu of the gray matter backward ; and this is still further accomplished by the olivary body, until the gray matter is spread out toward the posterior sur- face of the medulla at its upper part. Here the central canal of the cord has widened out to form the lower part of the 1 In speaking of these fibres it has been convenient to say that they "pass" in certain directions or " are distributed" in some situation. It must not be forgotten that they are afferent and efferent medullated nerve-fibres, and that such terms must be considered as somewhat figurative. In reality, it would not seem proper to speak of an efferent fibre as being " distributed " at its origin, but convenience and usage permit the use of these and similar expressions. 188 NERVOUS SYSTEM. fourth ventricle, and the gray substance is aggregated just under the floor of the ventricle. There are also sonic other collections of gray matter for example, in the olivary bodies. In this gray substance are the nuclei of origin of some of the cranial nerves : the spinal accessory, hypoglossal, pneu- mogastric, and glosso-pharyngeal nerves ; and roots of the auditory, of the facial, and of the trigeminus nerves, arise in this important collection of gray matter. Of the smaller col- lections of gray substance, probably none has the peculiar interest which the floor of the fourth ventricle possesses for this reason. Function of the medulla : The functions of the medulla are practically of the same kind as those possessed by the spinal cord, but of a higher degree. They are as follows : (a) con- duction ; (6) reflex action ; (c) automatic action ; (d) inhibition. Conduction: As the medulla is the sole connecting link between the upper parts of the brain and the cord, it neces- sarily contains all fibres passing between these limits. Thus it conducts all impulses and transmits all sensations. Reflex action: The medulla resembles the spinal cord in being the seat of reflex acts; the only difference between them being in the fact that many of the reflexes performed by the former are of much greater importance to life than any performed by the latter. Of the many reflex acts arising from the medulla, the following are examples : (1) That part of digestion which is performed in the mouth is dependent upon medullary reflexes mastication, degluti- tion, and the secretion of saliva; and, probably, the secretion of the pancreatic and other digestive juices. In this con- nection the so-called vomiting-centre may be noted. (2) The respiratory functions are so-called automatic func- tions of the medulla, and are capable of being sustained by the nerve-force derived from the medulla alone. The centres for coughing and sneezing are also here. The pneumogastrie and phrenic nerves convey the afferent and efferent stimuli, though there may be communications with other nerves whereby sensory stimuli are applied. (3) " Cardiac depressor" is the name given to a bundle of nerve-fibres running from the heart to the medulla, which i< SPECIAL CENTRES. 189 stimulated when the heart is overworking against a high ten- sion in the bloodvessels. The result of the stimulation of this " cardiac depressor " is a stimulus to the vaso-motor nerves which results in a dilatation of the arterioles, and conse- quently the tension is diminished and the overwork of the heart ceases. This reflex act takes place in the medulla. Automatic action : The impulses which are sent out to muscles without apparent afferent stimuli, and without an effort of the will, are called automatic. Such rhythmic im- pulses as those which maintain the respiratory function be- long to this class. It is not to be doubted that such actions are reflex, and in response to stimuli. In the case of the lungs, for example, the presence of deoxygenated blood may serve to excite an afferent impulse. Nevertheless, some authors distinguish between automatism and reflex action. This automatic action cannot be considered as at all the same as an action of the brain proper, like volition, but rather as a high grade of reflex action. On the other hand, some authorities claim that it is truly automatic in some of its acts, in that it is capable of direct stimulation by the condition of the blood circulating through the environment. It is bilateral, and probably consists of both an inspiratory and an expiratory part. It may be influenced to a certain extent by voluntary impulses. Inhibition: This is the control or inhibition of action through the nerves which are distributed from this region and through the communications with other centres in the cord. Besides, there are supposed to lie in the medulla centres which maintain the nutrition and tone of the muscles. These are known as control and tonic centres. Special centres : In addition to the centres already given there is a considerable number of centres in the medulla which control many important and complicated coordinated muscular actions. These are centres of reflex action for the most part- that is, are called upon to act in response to stimuli derived from an afferent impulse or to a voluntary effort. The following are examples : (1) Regulation of the heart's action is found here, both inhib- itory and accelerator centres communicating through the vagus. 190 NERVOUS SYSTh'M. (2) Vaso-motor regulation of the unstriped muscular fibre of the arteries is al>o accomplished by the medulla. A peculiar vaso-motor disturbance is brought about by injury of one centre of the medulla namely, the interference with the glycogen-function of the liver and the appearance of sugar in the urine the diabetic centre. (3) Various centres which have to do with the regulation of the body-temperature. The vaso-motor centres we have already mentioned. There are also found special wct- centres ; and, furthermore, a control of the special sweat- centres found in the cord is here maintained. Upon plausible theoretical grounds there is also assumed to be a licut-iiiliibi- tory centre, by which the heat-production is controlled without reference to vaso-motor conditions. (4) Also see example under " Reflex action." Other functions : The origin of the roots of certain of the cranial nerves here has caused the special senses of hearing and of taste to be referred to this region ; and the connection with the sympathetic system through the cord has caused the centre for the dilatation of the pupil to be located in the medulla. Phonation is also dependent upon the action of nerves arising in this focus of gray matter, and no voluntary or reflex sound can be produced by an animal in which the speech-centre in the medulla is destroyed. The origin here of the hypoglossal and pneumogastric nerves, involving as they do the movements of the tongue and glottis, controls both the acts of phonation and articulation. Demonstration of respiratory functions of the medulla : For this purpose we select an active, healthy frog : If the spinal cord be removed up to the medulla, the respirations continue, and in the same way they do not cease if the hemispheres be removed without disturbing this organ ; or, if both cord and hemispheres be removed without disturbing the medulla, the movements of breathing will continue. If the medulla is injured at the origin of the pneumogastric nerve, however, the movements of respiration cease and the animal dies. The same occurs when a similar injury happens in the higher animals and in man. Death occurs instantaneously in this way when the medulla is broken near the axis in executions CEUEA CEREBRL 191 by hanging " the neck is broken " or when an animal is killed by "pithing" in laboratory experiments. Glosso-labio-laryngeal paralysis : It may be of interest at this point to observe the pathological changes that occur in the medulla in producing the disease known as glosso-labio- laryngeal or bulbar paralysis : It is a progressive degenera- tion of the gray matter of the medulla, and it shows itself first in a paralysis of the tongue, which renders articulation of certain sounds indistinct : as the degeneration progresses in the medulla articulation becomes more and more im- possible and deglutition is affected. The disease continues to aifect more and more of the functions dependent upon the medulla, until death ensues as a result of involvement of the cardiac and respiratory centres or of inability to take food. Pons, Crura Cerebri, and Corpora Quadrigemina. The pons Varolii is a collection of nervous tissue lying immediately above the medulla. It consists of white fibres, with areas of gray matter filling in the intervals between the fasciculi of white fibres. The white fibres connect the mid- brain with the medulla, and also pass between the hemi- spheres of the cerebellum. What is the function of the gray matter is little known ; but some of it is directly continuous with that of the medulla, and, like it, active as a centre of nervous force. In the pons is the decussation of many nerve- fibres, among which are some of the fibres of the facial nerve arising in the floor of the fourth ventricle. Paralysis following lesions of the pons : The so-called crossed paralysis may follow lesions in the lower portion of the pons that is, paralysis of sensation and motion, more or less complete, of the opposite side of the body, with paralysis of the facial muscles of the same side as the lesion. Crura cerebri : The crura are formed of fibres passing from the medulla through the pons Varolii, to the hemispheres of the cerebrum. They divide so as to form two sets of fibres : the more superficial (crustse) are mostly motor or efferent fibres which are continuous with the pyramidal tracts in the cord; while the deeper (tegmentum) layer of fibres are afferent or 192 NERVOUS SYSTEM. sensory, and are derived largely from the lateral and posterior tracts of the cord. Lying between these bands of fibre.- i- a mass of gray substance (locus nigcr) whose function as a nerve-centre is not understood, though it has to do with coordination of the muscles, and especially with regulation of the muscles controlled by the motor oculi nerve. Paralysis following lesions in the crura : This is paralysis of the opposite side of the body, both of sensation and motion, and of a degree of intensity depending upon the size of the lesion ; and, besides this, paralysis of the motor oculi nerve of the same side as the lesion. There is a derangement of the coordination of motions which follows lesions of this region beyond that which belongs to the motor paralysis; this is often shown in rotary movements when the subject attempts to walk. It is inferred that there are coordinating influences derived from the crura. The corpora quadrigemina are four rounded eminences placed in pairs, two anterior, two posterior. They are situ- ated on the dorsal aspect of the mid-brain below the posterior lobes of the cerebral hemispheres and between the optic thalami and third ventricle in front and the cerebellum behind. Functions: The anterior corpora quadrigemina are the homologues of the optic lobes in some of the lower animals, and the anterior pair may be regarded as important centres for the visual and motor functions of the eyes. The posterior pair are more intimately associated with the sense of hearing. The special functions of the corpora quadrigemina will be more fully discussed under the special senses. Not only doe- blindness follow lesions of the anterior corpora quadrigemina, but there is often atrophy of them when the eyes are de- stroyed.. Cerebrum. The cerebrum is composed of two hemispheres, connected by a commissure of white fibres, the corpus callow m, and of the third ventricle and optic thalami (di- or thalamencephalon). The two hemisphere* are separated by a deep fissure ex- tending from before backward, and in the interior of each is REGIONS OF THE CEREBRUM. 193 found a cavity known as the lateral ventricle. The hemi- spheres are connected directly with the spinal system by the crura cerebri and medulla, and with each other by the corpus callosum. They are composed of white and gray nerve- substance, and the latter is arranged largely at the periphery of the hemispheres ; the former being made up of communi- cating nerve-fibres which connect the various portions of the hemispheres with each other, and the hemispheres with other parts of the cerebro-spinal system, thus allowing a free control of the impulses arising from one cell or set of cells by other cells in the gray matter. The surface of the cerebrum is divided into regions by fissures, which separate one part from another. These fis- sures are always present, and upon them depends the deter- mination of the division of the cerebrum into lobes. Fissures which are of use in locating the lobes of cerebral matter are the fissure of Rolando, the fissure of Sylvius, and the parieto- occipital fissure (Fig. 95). Convolutions : The surface of the brain is further cut up by a number of other clefts, known as sulci ; and these separate the surface into a number of distinct masses or convolutions. The depth of the sulci and their number determine the quality of the brain in respect to its degree of development ; thus, the convolutions in man are much deeper and more numerous than in the lower animals. The sulci are not in- variable in position or number in different brains. Regions of the cerebrum: (1) Frontal lobe: This lobe is bounded by the fissure of Rolando, and contains several con- volutions which include the forward portion of the brain. (2) The parietal lobe lies behind the fissure of Rolando, and extends posteriorly to the occipito-parietal fissure. The convolutions are well marked, and are separated by a well- marked sulcus (sometimes known as the intra-parietal fissure) ; and the posterior branch of the fissure of Sylvi us is enfolded by an inferior parietal convolution, known as the " supramar- ginal convolution." (3) The tempo ro-sphenoidal, or temporal lobe, is below the Sylvian fissure and in front of the occipital lobe. Its convo- lutions are well marked. 13 Phys. 1 ( J4 NERVOUS SYSTEM. (4) The occipital lobe is found at the posterior end of the cerebral hemisphere, and its convolutions are continuous with those of the parietal and temporo-sphenoidal lobes, except within the longitudinal fissure, where it is cut off from the parietal by the parieto-occipital fissure. Plan of the human brain in profile, showing its fissures and convolutions. S, fissure of Sylvius ; S', anterior branch ; S", posterior branch ; R, fissure of Rolando ; P, intraparietal fissure. (5) The central lobe, or island of Rctt, is within the fissure of Sylvius, and is covered by convolutions of the frontal and parietal and temporal lobes (see 5 in Fig. 96). Besides these well-defined lobes, the portion of the cerebral surface which is trithin fin- longitudinal fixture is marked l>v sulci and convolutions. Convolutions of the frontal, parietal, and occipital lobes are found here: the marginal convolu- tion, the gyms fornicatus lying above the corpus eallosum, the paracentral lobule, the quadrate lobule, and the cuneato lobule are among the principal landmarks. ARRANGEMENT OF THE GRAY MATTER. 195 Arrangement of the gray matter : The increase in the area Horizontal section of the hemispheres at the level of the cerebral ganglia. 1, great longitudinal fissure between frontal lobes ; 2, great longitudinal fissure between occipital lobes : 3, anterior part of corpus callosum ; 4, fissure of Sylvius ; 5, convolutions of the insula ; 6, caudate nucleus of corpus striatum ; 7, lenticular nucleus of corpus striatum ; 8, optic thalamus ; 9, internal capsule ; 10, external capsule ; 11, claustrum. of the surface of the hemispheres by the infolding of the sulci adds very greatly to the amount of gray substance in 196 NERVOUS SYSTEM. FIG. 97. Gray matter of the cerebral cortex (Muyiicrt). the brain ; for the entire sur- face i.s composed of gray sub- stance, and this follows the sulci and fissures (Fig. 96) in all their folds, and is not cut into by them. Besides the gray matter in the convolutions, there are certain other grav masses in the substance of the white matter; the optic thalami, the corpora striata, and the claustrum (Fig. 96) are the chief of these gray masses. Minute structure of the gray matter : The gray matter of the cortex is made up of gan- glion-cells of various shapes and sizes lying in a loose neuroglia-stroma, more abun- dant at the surface. The cells are the source of numer- ous nerve-fibres, which par-s out into the white matter (Fig. 97). There are counted five layers of these ganglion ie tissues; and while these zones merge into one another, they are tolerably distinct. In the middle (and widest) Inver large multi polar cells are very numerous, and the fibres may be seen to pass through the deeper layers in bundles into the white matter. Chemistry of brain-tissue : Brain-tissue consists of some peculiar bodies allied to the rats, but containing nitrogen ; COURSE OF THE FIBRES IN THE HEMISPHERES. 197 of these, cerebrin and lecithin are the more prominent. Aside from these, the constituents are proteid and fatty substances, with salts, chiefly potassium and magnesium phosphates, and water. Weight of brain : About three pounds. In size it exceeds the brains of all the lower animals except those of the ele- phant and whale. Its weight is about one-fortieth of the total body-weight, and this ratio is greater than in the lower animals, with a few exceptions among the smaller birds and monkeys. In women the weight is about one-tenth less than in men. Relation of size of brain to the intellect: In some degree the size of the brain bears a direct relation to the intellect of the individual, but this is not absolute. The depth of the sulci and the consequent size and complexity of the convolu- tions are a more efficient measure of the brain-power. In the largest of the apes the brain of an adult animal is about the same in weight as that of a human infant at birth. Idiots, as a rule, have brains much smaller than the normal, and in them the convolutions are apt to be ill-marked and uncom- plicated, as is the case in the lower animals. Course of the fibres in the hemispheres : The course of these fibres may be classified in three groups : 1, commissural fibres; 2, fibres of association ; and 3, medullary fibres. (1) The eommissural fibres are those which connect one hemisphere with the other, and it may be said that these fibres connect each set of convolutions with the corresponding set of the opposite side. The convolutions of the portion of the brain lying above the fissure of Sylvius communicate by the corpus callosum. (2) Fibres of association are those fibres which connect the convolutions of one hemisphere. These fibres pass in bundles just beneath the cortical gray matter of the convolutions, and it is thought that most of the important convolutions of each hemisphere intercommunicate in this way. (3) The medullary fibres are those which connect the cere- brum and medulla, and are regarded as indirect and direct, according as they do or do not pass to the gray ganglia at the base of the cerebrum. In considering the course of the fibres 198 NERVOUS SYSTEM. from the medulla through the crura cerebri it was noted that the motor and sensory fibres were to some extent separated. The fibres pass from the crusta to the internal capsule, and hence the "direct fibres" pass to the cerebral convolutions through the corona radiata ; while the " indirect fibres " from the legmen turn pass to the corpora striata and optic thalami, and communicate with ganglion-cells in them. Function of the corpora striata and optic thalami: These "basal ganglia," with the other collections of gray substance, seem to have a controlling influence upon the spinal system. It is through these ganglia that all voluntary impulses, ex- cept those by the direct medullary fibres, must pass. These basal ganglia communicate through the corona radiata with the convolutions of the cortex, and it is probable that we may regard this part as acting as a middleman to elaborate and coordinate the voluntary impulses of the cortex and to act in matters not requiring the intervention of the higher endowments of the mind. This status of these ganglia is quite theoretical, but the function may be considered as a sort having the properties of both the automatism of the gray matter of the medulla and cord and the voluntary function of convolutions. In this consideration, however, we must not undervalue the communication with the cortex which these basic ganglia possess. Lesions of the basic ganglia : So far as has been observed, the corpora striata may be involved by considerable lesions without causing persistent motor or sensory disturbances, and the same may be said of the optic thalami ; but if the lesion encroaches upon the white matter of the internal capsule, or crura cerebri, the effect is to cause more or less paralysis, depending upon the severity of the lesion and its position. Functions of the cerebrum: The motor and sensory func- tions which have been seen to belong to other nuclei or gray matter are centred here, but infinitely broadened, for the cells in the convolutions of the cerebrum can originate the eilcivnt and perceive the afferent nerve-impulses. In fact, it is iu this portion of the brain that the intelligence is centred : it is the organ of the mind. Memory, reason, emotions, and all SENSORY AREAS. 199 the other attributes of the mind are dependent upon its func- tional power. Unilateral action of the brain : There are instances in which injury or disease of one-half of the cerebrum has left the intellectual faculties not gravely impaired. From a consider- ation of such cases it has been held that the action of one of the hemispheres is sufficient for the purposes of the mind. There is, however, an absolute dependence for motor and sensory functions upon the integrity of both sides, for the one side cannot act for the other in these functions. As a rule, it is safe to assume that the hemispheres act in unison. Localization of brain-function : While the brain is regarded as an organ of the mind, it is probable that the various func- tions may be regarded as belonging to definite portions of the convolutions which are appropriated for that purpose. The functions of the convolutions have not been assigned, except for a small portion of the brain-surface and for some of the simpler actions. For the most part, our knowledge of the exact localization of brain-functions is confined to "motor areas," in which it has been determined that stimulation of a certain group of cells will cause a definite action. Besides this, certain other centres are located, as of sight and speech. Determination of motor areas: When the surface of the brain is exposed in animals or in man, the stimulation of certain areas of the cortex by a mild electrical current will give rise to motion in the peripheral muscles ; and it is found that the stimulation of the same region in the same or other animals will cause the same results. These centres of motor impulses are situated almost entirely upon the convolutions about the fissure of Rolando (Fig. 98). Sensory areas: These have not been, by any means, so definitely fixed as the motor centres ; but the centres for sen- sation may be said to exist, and probably in the convolutions of the posterior portion of the cerebrum. The centre for vision in the convolutions about the posterior branch of the fissure of Sylvius is generally accepted ; or in the cuneate lobule (14, 15, in Fig. 98). The centre for hearing is toler- ably defined in the temporo-sphenoidal lobe along the pos- terior branch of this fissure (16, Fig. 98). The speecA-centre 200 NERVOUS SYSTEM. is also located with seeming accuracy along the anterior branch of the fissure of Sylvius and in the third frontal convo- lution. This centre seems to be much more developed upon the left side of the brain. In Fig. 98 this centre may be indicated roughly by reference to the tongue-centres (8 and (h. FIG. 98. 10 Brain of monkey, showing the position of the motor and sensory centres as ascer- tained by Ferrier. The actions all occur on the side of the body opposite to the part of the brain irritated. 1, the eyes open widely, the pupils dilate, and head and eyes turn toward opposite side ; 2, extension "forward of the opposite arm and hand, as if to reach something in front; 3, movements of tail (and trunk) : 4, retraction with adduction of opposite arm; 5, supination and llexioii of the forearm, by which the arm is raised toward the mouth ; 0, action of zygomatics, by which the angle of mouth is retracted and elevated; 7, elevation of ala of nose and upper lip; 8, opening of mouth with protrusion of tongue: 9. retrac- tion of tongue ; 10, retraction of opposite angle of mouth ; a, f>, <\ a- tion, motor nerves, and mixed nerves (/. e., sensorv and motor). C (I) olfactory, (II) optic, (VIII) XT f . , auditorv and parts of the (Y) Nerves of special sense, < * i / T v\ ] trigeminus and (IX) glasso- ^ pharyngeal. Nerves of common sen- f The greater portion of the (V) sation, \ trigeminus. " (III) motor oculi,(IV) pathcticus, (V) lesser division of the tri- Motor nerves, geminus, ( V [)abducens, ( V 1 1 ) facial, and (XII) hypoglossal, (XI) spinal accessory (?). ( (IX) crlosso-pharyngeal and (X) Mixed nerves, | pneumogastrio. 777 NERVE. 207 FIG. 100. Of the nerves of special sense, (I) olfactory, (II) optic, and (VIII) auditory will be explained later, and may be omitted from further consider- ation for the present. Ill nerve : It arises from a nucleus of gray matter just in front of the pons beneath the iter e tertio ad quartum ven- triculum (aqueduct of Sylvius), passing out through the cms cerebri, and emerging from the skull in the orbit (Fig. 100). It gives off some fibres to the lenticular ganglion. It is dis- tributed to all the muscles of the eyeball, with the exception of the superior oblique and the external rectus muscles. It also supplies the levator palpebrse superioris muscle, and by its connection with the lenticular ganglion con- trols the ciliary and pupillary muscles. It is purely a motor nerve, function is best described by showing the paralyses which "low its division : by paraly- of the elevator of the up- lid we have ptosis; by paralysis of the muscles of the eyeball we have inability to move the organ up or down or inward ; by the unopposed action of the external rectus the eyeball becomes turned outward (external strabismus) ; by paralysis of the muscle of the iris the pupil remains dilated and does not respond to A partly diagrammatic view of the floor of the aqueduct, looking upward (dorsally), nuclei of the third and fourth nerves, and the decussating fibres of the latter all shown; the third nerve nuclei are subdivided into an anterior nucleus, the Edinger- Westphal nucleus (a and 6), and a posterior nucleus ; the posterior nu- cleus has a dorsal, a ventral, and a mesal portion; the decussation of the fibres from the dorsal portion of the posterior nucleus of the third nerve is shown (Edingerj. 208 NERVOUS SYSTEM. light ; and by paralysis of the ciliary muscle the accommo- dation of the lens for near vision is prevented. The control of the pupil is not a voluntary one; but the effect of a strong voluntary effort, exerted through the third nerve, shows itself in contraction of the pupil, as when the eyeball is turned strongly inward and upward. IV nerve : It arises close by the third nerve beneath the aqueduct of Sylvius, and emerges, after decussation, from the valve of Yieussens. Thence, passing around the crus cere- bri, it runs parallel with the motor oculi (third) nerve to the orbit, where it is supplied to the superior oblique muscle. Its paralysis prevents the muscle from maintaining the hori- zontal plane of the eyeball. If this paralysis occurs, there is double vision, and the image seen by the affected eye ap- pears oblique and inferior to the image of the other eye. This may be corrected by inclining the head to the opposite side. This nerve is also known as the trochlearis or trochlear nerve. V nerve : This nerve resembles the spinal nerves in having a motor and a sensory root, the latter possessing a ganglion (Gasserian). The origin of the nerve seems to be in centres, separate for motor and sensory, in the floor of the fourth ven- tricle (Fig. 101). There are fibres which join the trunk of the nerve which are derived from the spinal cord. It emerges from the pons Varolii as two distinct nerve-roots. The larger of the two, the sensory, soon enters the Gasserian ganglion, the motor root passing beneath without communica- tion. The nerve then breaks up into three branches: of these the first and second are formed entirely from the sen- sory root, while the third carries all the motor fibres, and with them some of the sensory, so that the third branch of the nerve is partly sensory and partly motor. There is a partial decussation- of the fibres in the medulla, but many pass direct to the same side. Muscles supplied by the motor root, V nerve : The muscles supplied are those of mastication. The temporal, ma>-< !< T, and both pterygoid muscles, as well as the anterior belly of the digastric muscle and the mylo-hyoid, receive their inuer- vatiou from the motor root of the fifth nerve. Besides this, the tensor palati and tensor tympnni muscles are supplied SENSORY FIBRES. 209 by this nerve through its communication with the otic gan- glion of the sympathetic system. A branch to the buccinator View of the posterior surface of the medulla, the roof of the fourth ventricle being removed to show the rhomboid sinus clearly. The left half of the figure repre- sents : On, funiculus cuneatus and, (j, funiculus gracilis ; 0, obex; sp, nucleus of the spinal accessory ; p, nucleus of the pneumogastric ; p + up, ala cinera ; R, restiform body; XII', nucleus of the hypoglossal; t, funiculus teres ; a, nucleus of the acusticus; m, strife medullares'; 1, 2, and 3, middle, superior, and infe- rior cerebellar peduncles respectively ; /, fovea anterior; 4, eminentiateres (genu nervi facialis) ; 5, locus eceruleus. The right half of the figure represents the nerve-nuclei diagrammatically : V, motor trigeminal nucleus; V, median and, V", inferior sensory trigeminal nuclei; VI. nucleus of abducens; VII, facial nucleus; VIII, posterior median acoustic nucleus; VIIF, anterior median; VIII", posterior lateral: VIII'", anterior lateral acoustic nuclei; IX, glosso- pharyngeal nucleus ; A', XI. and XII, nuclei of vagus, spinal accessory, and hypoglossal nerves respectively. The Roman numerals at the side of the figure, from 'T to XII, represent the corresponding nerve-roots (Erb). muscle is probably not motor, but sensory. Lesions of the nerves paralyze these muscles. The sensory fibres of the fifth nerve are distributed in all three branches, and supply sensation to the skin of the face 210 NERVOUS SYSTEM. and anterior portion of the head, emerging from the bony canals upon the face at the supraorbital, infraorbital, and mental foramina. Sensory fibres are also supplied to the mucous membrane of the mouth and tongue (by the lingual branch) and to the muscles of the part (Fig. 102). FIG. 102. Diagram of the fifth nerve and its distribution. 1, sensitive root; 2, motor root; 3, Gasserian ganglion: I, ophthalmic division; II, superior maxillary division; III, inferior maxillary division ; 4, supraorbilal nerve, distributed to the skin of the forehead, inner angle of the eye, and root of the nose; 5, infra-orbital nerve, to the skin of the lower eyelid, side of the nose, and skin and mucous membrane of the upper lip; 6, mental nerve, to the integument of the chin and edge of the lower jaw, and skin and mucous membrane of the lower lip ; , //, external terminations of the nasal branch of the ophthalmic division, to the mucous membrane of the inner part of the eye and the nasal passages, and to the base, tip, and wing of the nose ; t, temporal branch of the superior maxillary division, to the skin of the temporal region ; m, malar branch of the superior maxillary division, to the skin of the cheek and neighboring parts: !>, bucral branch of the inferior maxillary division, passing along the surface of the buc- cinator muscle, and distributee! to the mucous membrane of the cheek and to the mucous membrane and skin of the lips; /.lingual nerve, to the mucous membraneof the anterior two-thirds of the tongue; of, auriculo-temporal branch of the inferior maxillary division, to the skin of the anterior part of the exter- nal ear and adjacent temporal region; x, x, x, muscular branches, to the tcin poral, masseter, and internal and external ptervgoid muscles: //, muscular branch, to the mylo-hyoid and anterior belly of the diagastric ; /, sensitive branch of communication to the facial nerve. Destruction of the sensory root results in complete anaesthe- sia of the skin of the face and mucous membrane of the mouth. VI NERVE, 211 The trophic influence of the sensory root is of very great value. If it be divided, the complete anaesthesia of the conjunctiva, of the nostrils, and of the lips prevents the reflex self-protec- tion which belongs to the parts, and they become injured very easily. Aside from that, the direct influence upon all the parts is great, so that when it is cut off there is a rapid degen- eration resulting, which is specially apparent in the mucous membrane of the nose and in the cornea. Influence of the sensory branch on the special senses : (1) Its division causes total anesthesia to the skin and mucous mem- brane ; the loss of the sense of touch in the part is of great importance, for the tongue and lips are used much for this purpose. (2) Upon the sense of sight it has a very control- ling influence, for, as we have seen, the trophic influence is essential to the maintenance of the integrity of the eye. (3) Upon the sense of smell. Here the influence is the same as with the eyes, trophic. The smell is soon lost on account of degeneration of the mucous membrane after division of the fifth nerve. (4) Taste, probably, is not a direct function of the nerve ; but if the tactile sensibility is gone and the tro- phic changes are begun, the sense of taste soon disappears in the anterior portion of the tongue. (5) Upon the hearing the effect is more gradual and less distinct. The secretions of the cavity of the tympanum and of the external auditory canal are of great importance in maintaining normal condi- tions. They are under the trophic influence of the fifth nerve, both through its auriculo-temporal branch and through its communication with the otic ganglion. The tensor tympani muscle is also supplied by the motor root. Thus, the auditory apparatus is considerably under the control of the nerve. Symptoms due to pathological changes in the sensory root : Headaches of the scalp and deeper tissues, and more especially the frontal sinuses, are common. Toothache and facial neu- ilgia are due to irritation or disease of parts of the nerves. douloureux is a persistent neuralgia of some or all of the inches of the nerve. VI nerve : It arises from a nucleus of gray matter in the floor of the fourth ventricle, and its nucleus is more or less directly connected with those of the third, fourth, and seventh 212 NERVOUS SYSTEM. nerves. It emerges without decussation at the posterior border of the pons Varolii, and passes forward to the orbit with the third and fourth nerves. In its course it has many commu- nications with the sympathetic nerves, but their significance is unknown. It is supplied to the external rectus muscle of the eye, and its stimulation causes external squint, and paraly- sis causes internal. VII nerve : It arises in the floor of the fourth ventricle, and its fibres emerge from the upper part of the groove be- Fio. 103. Diagram of the facial nerve and its distribution. 1, facial nerve at its entrance into the internal auditory meatus : 2, its exit at the stylo-mastoid foramen; 3,4, temporal and posterior auricular branches, distributed to the muscles of the external ear and to the occipilalis; 5, branches to the frontalis muscle; r>, branches to thestylo-hyoid and digastric muscles ; 7, branches to the upper part of the platysma myoides: 8, branch of communication with the superficial cer- vical nerve of the cervical plexus. tween the olivary and restiform bodies in company with the (eighth) auditory nerve (sometimes known as the portin mol- lis ; the facial being then called the portio dura of the "sev- enth" pair, when the classification of the cranial nerves is made into nine pairs). It passes into the internal auditory CHORDA TYMPANI BRANCH. 213 canal, and escapes from the skull by way of the aqueduct of Fallopius and the stylo-mastoid foramen. It is almost wholly a motor nerve, and is distributed to all of the muscles of the face (Fig. 103) except those mentioned as controlled by the motor branch of the trigeminus nerve. The muscles of the eyelids and some of the muscles of the palate in part are innervated by it, as well as the parotid and submaxillary glands through the chorda tympani. In the neck it supplies the posterior belly of the digastric and the platysma myoides muscles. It also sends branches to the stapedius muscle of the internal ear and to all of the muscles of the external ear. The branches passing to the salivary glands are secretory in their function ; and this is the only exception to the motor influence of the nerve. Function of the VII nerve : It is the motor nerve which parallels in its distribution the sensory root of the fifth ; it supplies the superficial muscles, as the latter does the skin. It is the nerve of "expression," by which the features are made to reflect the emotions. Paralysis of the VII nerve : If the nerve be divided or dis- eased, the face of that side is devoid of motion (Fig. 104), and becomes smooth and expressionless, while the sound side is held in its customary pose. The eyelids cannot close them- selves, and the lips do not oppose properly, on account of the defective action of the orbicular muscle. There is difficulty in drinking and in articulation for the same reason. The eyelids remain open in facial paralysis, and the con- junctiva is subject to injury by drying and by foreign bodies ; but the injury is not so great as in paralysis of the fifth nerve, because the seventh has no trophic influence. Chorda tympani branch : The chorda tympani is a small filament given off from the facial in the aqueduct of Fallo- pius, some of whose fibres are distributed to the submaxillary gland. If this nerve be divided, the secretion of saliva from the gland is greatly diminished, while stimulation of the nerve will excite a copious flow. This is an active secretion, and is not a simple filtration due to vaso-motor changes. A similar influence is noted in the corresponding half of the tongue. 214 NERVOUS SYSTEM. There is a similar distribution of fibres from the facial to the parotid gland, which also receives secretory fibres from the glosso-pharyngeal through the lesser superficial petrosal Fio 104. Facial paralysis of the right side. nerve, but their action has not been so thoroughly analyzed as in the case of the submaxillary and chorda tympani. The chorda tympani has still further an effect upon the sense of taste in the anterior portion of the tongue. If it be divided, the taste is much blunted on the affected side. It is FUNCTION OF THE IX NERVE. 215 now known that this is due to the communication with the glosso-pharyngeal nerve, thus : the chorda tympani has fibres from the otic ganglion, which, in its turn, receives the lesser superficial petrosal nerve from the glosso-pharyngeal. IX nerve : It arises in the medulla from centres near those for the vagus and spinal accessory nerves. Its fibres pass through the substance of the medulla and emerge in com- pany with those of the vagus and spinal accessory, to pass with them from the skull through the jugular foramen. It gives off a small branch, which passes to the tympanum and Eustachian tube (Jacobson's nerve) while in the jugular fora- men, and presents a small ganglion, the petrosal ; and it has communicating branches to the seventh and tenth nerves and to the otic ganglion. The nerve divides as it passes down, one branch passing forward to the tongue, and one going to the pharynx (whence its name). The portion which passes to the tongue is distributed to the posterior portion of the organ, to the circumvallate papillae, and the mucous membrane behind them, some fibres going to the lining of the soft palate, pillars of the fauces, and tonsils. The other branch is distributed to the mucous membrane of the pharynx, and by direct branches and communication with other nerves to all the muscles involved in swallowing. Function of the IX nerve : (1) It is the nerve of taste; and (2) it is essentially a nerve of deglutition. (1) It is only in the latter part of the stay of food in the mouth that it reaches the region directly supplied by this nerve. When the food is to be swallowed, it is pressed by the base of the tongue against the palate arch and pushed into the pharnyx. It is then that the sense of taste is here exercised. The reflex stimuli then excited start up the motor chain, which pushes the bolus on to the stomach. At one time there was considerable question as to whether the trigeminal or glosso-pharyngeal was really the conductor of this sense, but it is quite likely that both are essential to its proper appreciation. (2) Whether by reason of its communications with other nerves or not, in its distribution the nerve is a motor nerve as NERVOUS SYSTEM. well as sensory. Its distribution is to all the muscles of deglutition, and stimulation causes contraction of the muscles, while division paralyzes them. The very numerous connec- tions of the nerve complicate its anatomical origin very greatly, and interfere with a clear comprehension of the un- aided function of* the nerve. The reflex for swallowing originates in the medulla ob- longata, where the origin of the nerve is situated. X nerve : The Xth nerve is also known by two other names : " pneumogastric," from its distribution and function ; " vagus " or " par vagum," from its scattered distribution (vagus, Latin, wanderer). It arises from the gray matter in the floor of the fourth ventricle, its nucleus being very close to those of the glnsso- pharyngeal and spinal accessory. Its fibres pass through the substance of the medulla oblongata, and emerge from its lat- eral surface with the roots of its associate nerves, the glosso- pharyngeal, and spinal accessory. It passes from the skull with them by the jugular foramen. It has at this point a ganglionic enlargement. From here it passes down the neck, and is distributed more diffusely than any other cranial nerve. It is supplied to the organs by which air and food enter the body, and besides this has several important connections with the sympathetic system. (1) To the larnyx it supplies sensation and motion through the superior and inferior lari/n- geal branches. (2) In the chest it forms the pulmonary plexuses, which innervate the bronchi and lungs. (3) Branches to the cardiac plexus supply important stimuli to the heart and great vessels. (4) There are branches to the pharyngeal and cesophageal plexuses which are both sensory and motor, supplying both the mucous membrane and the muscular structures of the parts. (5) Its terminal branches supply the sensory and motor nerves to the stomach, the left nerve being distributed on its anterior wall, and the right poste- riorly. (6) Branches also pass to the liver and spleen and communicate with the solar plexus. Communication between X nerve and other nerves : Soon after leaving its origin in the medulla the vagus enters into EFFECT OF SECTION OF X NERVE ON RESPIRATION. 217 so many communications with other nerves, both sensory and motor, that it is difficult to know the real fibres of the original root and to determine what are original and what derived functions. The sympathetic system sends fibres in all the branches of the pneumogastric, and the pneumogastric sends branches to many of the important sympathetic plexuses and ganglia ; the pharyngeal, laryngeal, resophageal, pulmonary, cardiac, and solar plexuses are so made up by branches from both. The spinal accessory nerve is an important contributor, in that it sends a large branch which is incorporated in the vagus. The fibres from the spinal accessory nerve are motor, and are, most of them, carried in the recurrent laryngeal nerve. There are also communications to the glosso-pharyn- geal and hypoglossal nerves, and it also receives motor fibres from the facial and upper two cervical nerves. The original nerve is probably entirely sensory, and its motor function is derived from these connections with motor nerves. Function of X nerve in respiration : The nerve supplies, as has been said, the motor and sensory functions of the larynx, and in this is of value to the respiratory function both in the prevention of foreign substances entering the rima glottidis, and in the opening of that orifice for the entrance of air. Besides this, it supplies sensory fibres to the pulmonary plexus which transmit the reflex stimulus to the medulla, by which the motor apparatuses excited to action. The muscles of the larynx involved in the production of sound are supplied by this nerve, and, as the approxi- mation of the chordae vocales is necessary for this, it fol- lows that the voice is dependent upon the fibres of the pneumogastric supplied by the inferior or recurrent laryn- geal nerve. Effect of section of X nerve on respiration : Respiration is slowed immediately to about half its usual rate, and soon drops to five or six to the minute, and even slower. The respiration is easy inspiration slow and full, expiration harsh and sudden. Death follows this operation in a short time (one to six days), and the animal during the time is sluggish and apparently suffers from slow carbonic-oxide 218 NERVOUS SYSTEM. narcosis. It is inferred from this that the vagus is the nerve which carries to the automatic centre the stimuli which are needed to keep up the automatism, and that the medulla is incapable of originating the motor impulses unless controlled by afferent stimuli. Function of X nerve in deglutition: Deglutition both in the pharynx and the oesophagus is under the influence of the vagus, which gives innervation directly to the thoracic part of the latter and through the inferior laryngeal branch to the cervical part. The sensory fibres act as conductors of the stimulus which results in the reflex peristalsis by which the food is carried on through the oesophagus. The sensory dis- tribution to the larynx must not be forgotten in this connec- tion, for by it food is kept from the respiratory organs. Sec- tion of the vagi causes paralysis of swallowing, and food is apt to pass the glottis on an attempt to swallow, not even a cough being excited by such an accident. The closure of the glottis in swallowing is caused by a reflex action known as the "action of arrest," and is derived from the sensory fibres of the vagus. Relation of X nerve to stomach : The stomach receives both motor and sensory function through the vagus. The stomach receives its warning of the presence of food through the sen- sory fibres, and the muscular fibres excite the organ to con- tract upon it and " churn " it about during digestion. There is also a vaso-motor influence derived from the vagus. AVhen the nerve is cut but little food can reach the stomach, because of the paralysis of the oesophagus, and what food does enter is digested very slowly, so that the function of the pncu mo- gastric may be considered essential to stomach digestion. The connection with the solar plexus also involves the intes- tines in the action of the vagus. Influence of X nerve on the heart: There are numerous branches to the cardiac plexus from the trunk of the vagus and from its inferior laryngenl branch. Stimulation of the pheumogastric nerve diminishes the frequency, or, if strong, entirclv stops the heart in diastole. The nerve is therefore regarded as having an inhibitory (trfion. 'Phis is an nnnsual effect, for in other cases the stimulation of nerves going to FUNCTION OF XI NERVE. 219 muscles causes contraction : the heart, however, becomes flaccid under the influence of the stimulated vagus. Other functions of X nerve : The vagus nerve is the channel for excito-motor reflexes in coughing and vomiting, as well as for many other less essential reflexes, such as sighing, hiccoughing, and the like. XI nerve : The origin of the Xlth nerve is twofold. One root arises in the gray matter of the medulla near the nucleus for the vagus, while the other arises from the lateral tract of the cord as low as the fifth or sixth cervical vertebra, and passes up between the anterior and posterior spinal nerve-roots to join the medullary (or accessory) portion at its emergence from the medulla. The united nerve soon divides and passes out through the jugular foramen with the glosso-pharyngeal and pneumogastric nerves, the medullary portion joining the trunk of the pneumogastric, w r hile the spinal root supplies the sterno-mastoid and trapezius muscles. Function of XI nerve : The nerve is a motor to all intents, though it has some sensory fibres, as is shown by the pain caused by pinching it. (1) The anastomatic branch, which joins the pneumogastric, is apparently largely given off in the recurrent laryngeal nerve ; but its section does not produce the same effect upon the larynx as section of the trunk of the vagus or of its in- ferior laryngeal branch. There is paralysis of the voice, but not of the movements of the glottis for respiration. There are probably some fibres of this nerve also given off to the cardiac plexus. (2) The muscular branch supplies the sterno-mastoid and trapezius muscles; but these muscles are also supplied by the cervical spinal nerves, and their action is not paralyzed by the section of this branch of the spinal accessory. It is found, however, that the relation of these muscles to respira- tion is impaired by isolation from this nerve that is, when the breath is held in any violent exertion, as straining or pushing, or when a loud cry is uttered, the sterno-mastoid and trapezius muscles contract to fix the head and hold the spine steady. This action seems to be prevented by the section of this muscular branch of the spinal accessory. 220 THE SENSES. XII nerve : It arises in the gray matter at the inferior extremity of the floor of the fourth ventricle, mesial to the origin of the spinal accessory and pneumogastric nerves. The fibres pass through the substance of the medulla ob- longata, between the pyramid and the olivary body, and, emerging in a number of small bundles, connect into a nerve- trunk which emerges from the skull by the anterior condyloid foramen. It passes down the neck to about the level of the hyoid bone, where it curves forward and into the tongue, giving off branches to the muscles which move that organ. Function of XII nerve : It is a motor nerve, but po>- some sensory fibres derived from the cervical spinal nerves and from the trigeminus, with whose lingual branch it inos- culates on the side of the tongue. Filaments from it arc distributed to all the muscles which move the tongue, and to the depressors of the hyoid bone through its descendiixj branch. The latter branch, although in the anatomical sheath with the hypoglossal nerve, is not an integral part of the nerve, but is derived from the upper cervical nerves. Influence of XII nerve on digestion : It is important in mas- tication, for its muscles move the food about for the better action of the teeth. In animals, after division, drinking is impossible, because they are unable to lap up fluids, and the food is swallowed with difficulty because it is not carried back into the pharynx by the tongue after mastication. Connection of XII nerve with speech : Articulation of most sounds involves movements of the tongue. Impaired articu- lation is an early symptom in bulbar or glosso-labio-laryn- geal paralysis. THE SENSES. A peripheral organ for the reception of an impression, a nerve for its conduction, and a centre in the brain for the perception, are the organs necessary for sensation. It i> by means of impressions so received and conducted to it that the mind is able to control the body and to take cognizance of the external world. COMMON AND SPECIAL SENSATIONS. 221 Classification of sensations : Common sensations and special sensations. These last are commonly called " the senses." Common sensations : Such perceptions as cannot be dis- tinctly located in any organ or set of organs, such as fatigue, hunger, thirst, satiety. Besides these, there are some sensa- tions which involve certain organs which must be classed under this head ; thus inclinations to cough or to sneeze or to vomit are common sensations, and, similarly, to urinate or defecate. Many of these sensations occupy a border-line be- tween common sensibility and the special sense of touch, such as itching and tickling. Pain is a common sensation, but is very closely allied to the sense of touch. The two may be differentiated, however. If one touch a sharp instrument, he may perceive its shape and condition ; but if the pressure be increased, the ability to perceive its form is lost, and instead the sensation of pain is established. The relation of the two is curiously shown in partial anaesthesia by drugs, as when one takes nitrous- oxide gas for the extraction of a tooth, and is able to feel the operation and to know what has been done, without in the least feeling pain. The seat of the senses is the brain, or sensorium. The organ of the mind, which perceives the thing which the organ of sense has taken an impression from, is the fundamental struct- ure in the necessary chain. Hallucination is the perception of an object as a real presence without the presence of the object to justify the perception ; that is, it is an act of the brain which refers its action to an organ of the senses. Thus, in delirium tremens a person may perceive many curious and uncanny things, which his mind hears and sees and feels, but which his senses could not take cognizance of, because they are only " creatures of the mind." Difference between common and special sensations : The most important distinction between common and special sen- sations, is that the former are strictly limited to the condition of our bodies, while by the latter we gain, in addition, infor- mation respecting affairs outside of our bodies. This differ- ence may be explained if we compare the sensations of pain 222 THE SENSES. and touch, the one a common, the other a special sensation. For example, if the point of a needle be gently pi against the tip of the finger, we only feel this point by our sense of touch and refer the sensation to the object causing it. But if the needle be pressed harder, so as to enter the skin, we feel at once a sensation of pain, which is no longer referred to the needle, but to the finger itself. The sensation of pain is not able to cause us to recognize the object which caused it, nor its nature. Sensations and perceptions : Our habit of referring sensa- tions to outside causes leads us to consider as properties of external bodies the sensations which they excite in us. "When we speak of anything as having a bad taste, we forget that it only tastes bad to us. This habit pertains especially to sensations of touch and sight. From constant exercise of it we finally come to believe implicitly in the " evidence of our senses." Judgments : There is a distinction between a sensation and a judgment (which is often unconsciously based on sensations). When we estimate the distance of an object from ourselves we form a judgment based on past experience of many sensa- tions, such as the number of steps we must take before we touch it, etc. Nerves of special sense: The special nerves have no other function than the special one for which they are set apart ; and when they are separated from their special orpins for receiving impressions, they no longer respond to the custom- ary stimuli. The special senses are touch, taste, smell, hearin, vein ; c, capillar ered by epithelium (from K6 a fungiform papilla with the bloodvessels injected: y loops of simple papillae in the neighborhood, cov- lliker, after Todd and Bowman). The tactile sense is very important, too, in the sense of taste, for with many substances the taste is largely due to their me- Vertical section of a circnmvallate papilla, from the calf (35 diameters). A, the papilla; B, the adjacent surface. The figure shows the nerves of the papilla spreading toward the surface and toward the taste-buds which are imbedded in the epithelium at the sides; in the sulcus on the left the duct of a gland is seen to open (Engelmann). chanical condition : this is the case with mucilaginous, oily, and chalky tastes. Association of smell and taste : It is important, for with many substances particularly aromatic substances of food SMELL. 229 and drink the association of smell and taste is very essential to a thorough appreciation of a flavor. Most cooked foods lose their savor if the nose is obstructed ; thus with a " cold " in the nose " everything tastes alike." The principal tastes are sweet, bitter, acid, alkaline, and saline. Besides these, the general sensibility of the tongue detects pungent or caustic and styptic tastes, as well as the oily and mucilaginous tastes. Sensibility of tongue : It is quite acute. A solution of acid or bitter substances is tasted when very dilute : strychnine is said to be tasted in a 1 : 600,000 solution ; sulphuric acid, 1 : 1000. After-taste : After an aromatic substance has been tasted there remains in the mouth an impression of that flavor ; and if such substances be taken in rapid succession, the apprecia- tion of their flavor is lost. This impression, which is left by a strong flavor, is called the after-taste, and is utilized some- times to cover the taste of a disagreeable medicine, a strongly flavored aromatic preceding it. Smell. Smell conditions : The first essentials are a special nerve and nerve-centre, the changes in whose condition are per- ceived as sensations of odor. No other nerve-structure is capable of such sensations, even when acted on by the same cause. The special organs for this sense for the reception, conduction, and perception of the stimulus, as in the case of any of the senses, must be in their normal condition, and a stimulus (an odor) must be present to excite them. Odors are caused either by minute particles of solid matter or by gases which are in the atmosphere, and they must be capable of solution in the mucus of the Schneiderian mem- brane. The substance must pass in a current of air through the nostrils or it is not perceived as an odor. This is accom- plished by " sniffing" the air, and thus creating an intermit- ting current which is tested by the olfactory sense. In this way a trace of a gas or impalpable powder may be de- tected which cannot be traced by chemical or other means. 230 THE SENSES. If the substance be applied as a solution, it is not detected ; thus, rose-water in a nasal douche is not noticed while the nostrils are full of fluid, and yet as soon as the nostrils are free the odor appears. The olfactory nerves are the functional nerves of the sense, and are spread out in a fine network (Fig. 109) over the sur- FIG. 108. Cells and terminal nerve-fibres of the olfactory region, highly magnified. 1, from the frog; 2, from man: a, epithelial cell,' extending deeply into a ramified peripheral rods ; e, their extremities, seen their central filaments ; 3, olfactory ni-rvr- fibrilUe (Frey, after Schultze). tne irog; z, irom man: a, epitneiiai ceil,' exi process ; b, olfactory cells : c, their peripheral in 1 to be prolonged into fine hairs ; d, their cent fibres from the do} ; n, the division into fine fi face of the superior turbinated bone and on the upper third of the septum. The nerves end in special end-organs, known as olfactory celts (Fig. 108), which lie under the ciliated epithe- lium of the part. Origin of olfactory nerves: The nerves arise from a mass >!' gray matter lying beneath the anterior lobe of the brain upon ORIGIN OF OLFACTORY NERVES. 231 the cribriform plate of the ethmoid bone. This is the olfac- tory bulb, and it is connected by the olfactory tract with the cerebrum. Each olfactory tract arises from the cerebrum by three roots, two of which are composed of white matter, the other largely of gray matter. By these it is connected with the olfactory centres. The lining membrane of the nasal cavity is very sensitive to irritation, the nasal branch of the fifth nerve and branches from the spheno-palatine ganglion furnishing the ordinary FIG. 109. Distribution of nerves in the nasal passages; 1, olfactory bulb, with its nerves; 2. nasal branch of the fifth pair ; 3, spheno-palatine ganglion. and tactile sense. Therefore we can perceive, by the nose, the sensations of cold, heat, itching, tickling, pain, and ten- sion or pressure. The perceptions of the olfactory and of the nerves of touch often resemble each other, and some stimuli affect both nerves. The common sensibility is evoked by such substances as are irritating and acrid : ammonia gas has no odor, but it stimu- lates the mucous membrane by its irritating properties. The tactile or common sensibilities remain when the olfactory are gone. The relation between the two kinds of perception is 232 THE SENSES. lost to us, and we speak of the smell of ammonia or of alco- hol when it is probably not an olfactory, but a sensory, per- ception. Acuteness of smell : The sense of smell is very acute, but not so sharp in man as in many of the lower animals. The distribution of the olfactory nerves is much wider in some of the animals, and the cerebral development is correspondingly increased. In man the range of susceptibility is, however, probably greater. The variety of odors and the very minute quantity of stimulant substance required to produce a sensa- tion of smell are quite wonderful. The most delicate analy- sis may fail to show traces of the substances which can be appreciated by the sense of smell. For instance 0.000000005 gramme of oil of peppermint in 1 litre of air can be appreci- ated. There are some odors pleasant to some which others find almost intolerable. Musk, for example, is a pleasant per- fume to some, while to others it is quite unendurable. In the same way, the acuteness of this sense in some is more marked than in others, and yet this may apply only to certain kinds of odors. Like the sense of touch and the other special senses, that of smell can be very much developed by practice. Large salaries are paid to experts in discrimination of the quality of wines, etc. Often in cases of mental disease there are hallucinations of smell, which also may occur in cases of disease of the olfactory centres, when there may be frequent complaint of a bad smell. With normal organs there may be a sensation of an odor which cannot be detected by others present. Sneezing : The act of sneezing is a violent and sudden ex- pulsion of air through the nasal passages. The act is a reflex one and the exciting cause is a stimulation of the nasal ii la- ments of the fifth or trifacial nerve. Hearing. The auditory apparatus consists, in brief, of (1) the external ear; (2) the middle car; (3) the internal ear; and (4) the auditory nerve. MIDDLE EAR, 233 External ear : The external ear consists of the auricle and external auditory canal. The former serves to receive the sound-waves and to indicate the direction from which they come in animals which possess the power of moving the organ. Through the external auditory canal the sound-waves are conducted to the middle ear. The middle ear, or tympanum, is a cavity in the temporal bone which is shut off from the external auditory canal by the membrana tympani. The Eustachian tube connects this cavity with the pharynx. The lining of the middle ear is FIG. 110. Right temporal bone of the new-born infant, seen from its inner side, showing the membrana tympani and chain of bones in their natural position (Riidinger). partly ciliated epithelium, continued from the mucous mem- brane of the pharynx through the Eustachian tube. There are two openings of importance the fenestra rotunda and the fenestra ovalis in the bony wall, but they are covered, the former by a membrane, the latter by the stapes. The middle ear also communicates posteriorly with the mastoid-cells, which are air-cavities in the mastoid process of the temporal bone. Its only communication with the external air, how- ever, is through the Eustachian tube. There is a chain of small bones (ossicles) which connect the membrana tympani and the fenestra ovalis. 234 THE SENSES. Membrana tympani : It is a tough, tense, fibrous membrane set in the bony opening of the external auditory canal. The degree of tension of the membrane is regulated by the tensor tympani muscle. Ossicles: They are three in number (Fig. Ill), and are so Fco. ill. Bones of the tympanum of the left side. A, malleus : 1, long or slender process ; 2, near neck ; 3, the handle; 4, short process ; 5, head. B, incus: 1, body ; 2, short or posterior process ; 3, the long process with the orbicular process. C, stapes : 1 and 2, head ; 3, neck ; 4, 5, crura; 6, base. D, the three bones in their natural connection ; TO, malleus ; sc, incus ; , stapes. articulated as to communicate the vibration of the membrana tympani to the internal ear (Fig. 110). The handle of the malleus is attached to the membrane, so that this bone moves with each vibration. This motion is communicated to the incus, which passes it on to the stapes. The stapes rocks in the fenestra ovalis, and is therefore capable of transmitting to SEMICIRCULAR CANALS. 235 FIG. 112. the fluid in the cavity of the labyrinth the impulses which it receives. Eustachian tube : The Eustachian tube or canal leads from the cavity of the pharynx to that of the middle ear. It is completely pervious and allows a free passage of air from the pharynx to the middle ear. Its chief purpose is to provide for the maintenance of an equal pressure on both sides of the membrana tympani, by keeping the air in the middle ear in communication with the outside air. It also serves to render sounds clearer (as the apertures in violins do), and as an outlet for mucus. Internal ear : The proper organ of hearing is formed by the distribution of the auditory nerve within the internal ear or bony and membranous labyrinths. The bony labyrinth is situ- ated in the dense petrous portion of the temporal bone, and consists of three essential parts : the vestibule (Fig. 112), and open- ing from it the semicircular canals and the cochlea. There is another opening, the aqueductus vestibuli, whose use is doubtful, and still others for the entrance of the audi- tory nerve-filaments. Within the bony labyrinth is the membranous labyrinth, a series of tubes and sacs composed of fibrous tissue, lined with epithelium, which contains a colorless fluid, the endolymph, while a fluid surrounds this membranous labyrinth, the perilymph. Semicircular canals : These canals are arched cylindrical spaces in the solid bone which open at each end of the arch into the vestibule. They are three in number, and two are nearly vertical and one is horizontal. These canals are arranged in such a man- ner that the planes of the two vertical canals are at right angles, one being antero-posterior, and the other transverse (Fig. 112). These canals have within them complete mem- External view of a cast of the left labyrinth (Henle). /, fenestra rotunda, or round window ; a, fenestra ovalis, or oval window ; b, ampulla of superior semi- circular canal; c, ampulla of horizontal semicircular canal; (1, common shaft of union of these two canals ; e, ampulla of posterior semicircular canal ; g, promontory (the line should extend more to the right). 236 THE SENSES. branous tubes, considerably smaller than themselves ; so that the perilymph occupies the space between the bony wall and the membranous wall, while the endolymph occupies the cavity of the membranous semicircular canals (Fig. 113). Within the cavity of the membranous semicircular canals are found the terminations of some filaments of the auditory nerve. These filaments end in collections of cells called " cristae acustica?." A similar collection of nerve-cells, called " macula acustica," is found in the membranous chamber FIG. 113. Membranous labyrinth. Cs, semicircular canals; U, utriculus ; S, sacculus; A, aqueduct of vestibule; Cr, ductus reunions ; Co, cochlea. (utricle), situated in the vestibule, into which the membranous semicircular canals open. The me of the semicircular canals does not seem to be directly connected with the auditory function of the part, but to be connected more with the sense of equilibrium. The movement of the fluids in the canals, arranged in the direc- tions of the three dimensions, may serve to produce sensations upon the cristse acusticaB which lead to the formation of ac- curate judgment of changes in the position of the body. Cochlea : It is a part of the bony labyrinth which derives its name from its resemblance to a snail-shell. It is divided into two parts by a bony and membranous septum which runs parallel from base to apex of the spiral (Fig. 114). The " upper " passage opens out of the vestibule, and is known as the scala vestibuli ; the "lower," the scala tympani, is shut off by a membrane, which covers the fenestra rotunda, from the cavity of the 'tympanum. The scala vestibuli is sub- divided by a membrane, which passes from the bom lamina spiralis to the wall of the scala vestibuli, shutting off a tri- CANALIS COCHLEARTS. 237 angular space (canalis cochlearis). The floor of this space is the membranous partition (membrana basilaris) which sepa- FIG. 114. Bony cochlea of the human ear, right side, opened from its anterior face (Cruveilhier). rates the scala tympani from it, and upon this membrane is the organ of Corti. The roof of the triangular space is the membrane of Reissner. Canalis cochlearis : The canalis cochlearis, or scala media, is the representative of the membranous labyrinth occupying the bony cochlea. It is a single continuous tube, spiral in shape so as to conform to the shape of the bony cochlea. On cross-section it is seen to be triangular, or, more correctly speaking, the segment of a circle. Its apex lies just over the edge of the lamina spiralis, and its base against the circum- ference of the bony cochlea. Its upper side is the membrane of Reissner, and its lower side the basilar membrane. At the apex of the spiral cochlea the canalis cochlearis terminates as a blind end. At the base of the cochlea the canalis cochlearis connects by means of a tiny duct (canalis reuniens) with a round membranous chamber (the saccule). The saccule and utricle are those portions of the membranous labyrinth occupying the bony vestibule. The saccule and utricle communicate with each other by means of the "ductus endolymphaticus," a minute tube lying in the aqueductus 238 THE SENSES. vestibuli. Thus it is seen that the canalis cochlearis is filled with endolymph, freely communicating with the interior of the rest of the membranous labyrinth ; but nowhere does the endolymph communicate with perilymph. Organ of Corti : Upon the basilar membrane is arranged a series of rafter-like bodies which roof in a small canal (Fig. 115) ; upon this are spread the functional nerve-endings of the auditory nerve. These rafter-like bodies are large nucleated cells, the rods of Corti, having resting upon them other epithelial cells with hair-like processes which project into the canalis cochlearis or scala media. When looked at from above the cells have an appearance similar to the key- FIG. 115. Diagrammatic section of the organ of Corti. 1, membrana basilaris; 2, 3, internal and external rods of Corti ; 4, epithelial cells near inner and outer borders ; 5, hair-cells lying in contact with the rods (magnified 500 diameters). board of a piano. Fibres of the auditory nerve spread to these cells from the bony lamina spiralis. It is probable that each of the functional cells in the organ of Corti responds to a particular shade of sound. Auditory nerve : The Ylllth cranial or auditory nerve is purely centripetal in its functions. Anatomically we say that its fibres arise from a nucleus of gray matter in the floor of the fourth ventricle, and from this source pass out through the substance of the medulla in a number of small bundles which unite with another root which has connections with the cerebellum to form a trunk. This passes with the facial nerve into the internal auditory canal, and terminates in special end-organs in the internal ear. Its fibres contain numerous ganglion-cells. In the cochlea there are many of these cells, and they form plexuses of nerve-fibres to supply the hair-cells. The absence of neuri- COURSE OF SOUND-WAVES. 239 lemma in the auditory nerve gives it a soft feel which has caused the name " portio mollis" to be given to it when it and the facial were considered as a single nerve. Course of sound-waves : Sound-waves of the air are gath- ered by the concha, carried into the external auditory canal and vibrate against the membrana tympani. The mem- brane taking up the vibrations transmits them through the chain of ossicles to the stapes in the fenestra ovalis. The stapes imparts its motion to the perilymph of the vestibule FIG. 116. IDTMil membranous cochlea; LS, lamina 'ossea; EM, eustachian tube ;~ AN, auditory nerve ; N, canalis reuniens. (Fig. 116). There is now set up in the perilymph a fluid- wave that travels in all directions. Some of this fluid-wave travels along the scala vcstibtili to the apex of the cochlea, then through the aperture of communication with the scala tympani, then down the latter until it expends itself against the membrane in the fenestra rotunda. In its passage along the scala vestibuli and scala tympani, the fluid-wave vibrates against the membrane of Reissner and basilar membrane ; and this sets up similar vibrations in the endolymph of 240 Till-: SENSES. the canalis cochlearis. The fluid- wave in the canalis coch- learis irritates the hair-cells of the organ of Corti. These cells seem to be able to respond to particular tones by their sensitiveness in selecting each one its particular fluid-wave with its particular rate of vibration. The remainder of the fluid-wave is expended in a similar manner through the vestibule and semicircular canals, with similar effects on the saccule, utricle, and membranous canals. Localization of hearing : The branch of the eighth nerve, " arising" in the organ of Corti, having received its impulses from the organ of Corti cells, transmits its impulse to the FIG. 117. Position of the auditory centre in the first temporal convolution (Gowers). centre under the acoustic tubercle in the floor of the fourth ventricle; thence fibres pass by means of the trapezium (in the pons) to the opposite side, and thence through the lower fillet of that side to the posterior quadrigeminal body, whence, by means of its brachium, internal geniculate body, optic thalamus, and internal capsule, they proceed to the cortex of the first and second temporal convolutions (Fig. 117). Distance: We can only judge of the distance of the source of a sound by its intensity. The sound itself is in the ear. Ventriloquists take advantage of this fact, and, by modify- ing the intensity of the voice in imitation of the effect of distance, cause us to think that it really originates at a distance. Subjective hearing : By this heading is meant those sounds PRODUCTION OF VOICE. 241 that are distinctly heard and yet are not produced by phys- ical sound-waves, nor are they hallucinations. They may be due to disturbances of the auditory apparatus or to abnormal conditions of surrounding organs. Thus, buzzing or ringing in the ears may result from the hypersemia of the parts and exaggerated rush of blood, or from a defect in the circulating apparatus (as by an aneurism), or from disease in the auditory nerve or some other portion of the apparatus. Hallucina- tions of hearing are very common among the insane, and are purely creations of the disordered brain. Musical range of hearing : The range of musical notes that can be appreciated by the human ear is about seven octaves. There are about three thousand hair-cells in the organ of Corti, and it will be easily seen that this would allow an enormous capability to differentiate sounds and musical tones. This corresponds to a range of from 40 to about 4000 vibra- tions per second. The range of audibility, on the other hand, is about eleven octaves, or from 16 to 38,000 vibrations per second. With less than 16 vibrations per second we are conscious only of separate shocks, while with more than the larger number we are unconscious of sound altogether. Voice or Speech. Although the voice or speech cannot be called part of the senses, nevertheless it seems appropriate to describe its mechan- ism under the general subject of " The Senses." The larynx : The larynx is the organ of voice. It is a cavity closed laterally, but communicating with the trachea below and the pharynx above. The walls are made up of the thyroid, cricoid, and arytenoid cartilages, together with various muscles and membranes (Fig. 118). For a detailed description see Quain's or Gray's Anatomy. Across the cavity of the chamber are stretched in an antero-posterior direction the two vocal cords or " mem- branes." The free passage between the vocal cords is the glottis, the cords forming part of the rima glottidis. Production of voice : Voice is a result of the vibrations of the vocal cords. The vibration of the vocal cords is 16 Phys. 242 THE SENSES. produced by the passage of the air in expiration, never naturally in inspiration. The I' I<; - 118 - quality of the voice as regards pitch depends upon the length of the vocal cords, the crico- thyroid muscles acting to in- crease the tension, while the thyro-arytenoids relax the cords and the crico-arytenoids dilate and contract the rim a glottidis. Falsetto and high- pitched notes in a naturally low-pitched voice are due to vibration at the edges of the cords. The hollow spaces about the oral and nasal cav- ities are of use as resonators or sounding-boards. Articulate speech : The voice comes from the larynx ; but articulate speech is the result of the modification of the voice by the tongue, etc. (Fig. 119). The organs used in articu- late speech are : The tongue and teeth in the formation of the linguals and dentals ; the nasal sounds by the cavity of the nose ; the other sounds are formed largely by modi- fications in the shape of the mouth in one or another part. Musical range of the voice : The musical range of a hu- man voice is from one to three octaves. In this, culti- vation and natural aptitude are factors which permit great variability. The total range Longitudinal section of the human larynx, showing the vocal mem- branes. 1, ventricle of the larynx ; 2, superior vocal membrane ; 3, inferior vocal membrane ; 4, arytenoid car- tilage ; 5, section of the arytenoid muscle; 6, 6, inferior portion of the cavity of the larynx ; 7, section of the posterior portion of the cricoid cartilage ; 8. section of the anterior portion of the cricoid cartilage : 9, superior border of the cricoid cartil- age ; 10, section of the thyroid car- tila-c : 11, 11, superior portion of the cavity of the larynx ; 12, 13, aryten- oid gland; 14, 16, epiglottis; 15, 17, adipose tissue; 18, section of the hyoid bone ; 19, 19, 20, trachea. EYELIDS. 243 of the human voice from the highest soprano to the lowest bass is about four octaves. Thus it will be seen that the Section of the parts concerned in the formation of vowels. Z, tongue ; p, soft pal- ate ; e, epiglottis ; g, glottis ; h, hyoid bone ; 1, thyroid : 2, 3, cricoid ; 4, arytenoid cartilage (Landois). range of sounds which can be appreciated by the ear is far beyond the capacity of the voice. Sight. The visual apparatus is the eye with its accessory organs and the optic nerve. The function of the eye is the reception of stimuli of light, whereby we are able to perceive the impressions of form, color, and conditions of our surroundings in infinite variety. It is far the most complex in structure of any of the organs of special sense, and the most rapid and delicate in its actions. The stimulus received by the eye is transmitted along the optic nerve to the brain. Accessory organs of the eye : Under this heading we class the (1) eyelids, (2) lachrymal gland, (3) extrinsic muscles of the eyeball. Eyelids : Each eye has two lids, an upper and a lower. Each consists of a thin plate of elastic tissue with a covering of loose skin and a smooth lining of mucous membrane the conjunctiva which is reflected upon the eyeball. Along the 244 THE SENSES. edges of the lids are a number of short curved hairs which screen the eye from foreign bodies. The extreme sensitive- ness of the conjunctiva helps in this by giving immediate warning when any foreign substance gets in the eye. The muscles of the eyelids are the levator palpebrse supe- riores and the orbicularis palpebrarum. Lachrymal gland : It is a small racemose gland lodged in the upper and outer part of the orbit. It has several ducts, which lead to the surface of the conjunctiva of the upper lid. The secretion of the gland is usually just sufficient to keep the eye moist, but under the stimulus of pain or intense emotion the secretion is increased, and appears in drops which flow from the eyes tears. Under ordinary circumstances a slight ex- cess of this moisture is drained into the nasal cavity by the lachrymal duct. This secretion is slightly alkaline, and con- tains about 1 per cent, of solids, chiefly sodium chloride. Meibomian glands : They consist of a number of small racemose glands, lying beneath the conjunctiva, which secrete an oily protective substance. The ducts of these glands open along the edge of the lid. Extrinsic muscles of the eye : There are six muscles to each eye superior rectus, external rectus, inferior rectus, internal rectus situated respectively above, below, and to the inner, and outer sides of the eyeball ; also the superior oblique at the upper and inner angle of the orbit, and the inferior oblique at the lower and inner angle. The recti serve to turn the eye up, down, in, or out, accord- ing to which muscles or combination of muscles are acting. The oblique muscles tend to rotate the eyeball on an antero- posterior axis. The superior and inferior recti of both eyes work synchro- nously, but the action of the external rectus of one eye is synchronous with the internal rectus of the other; and the same is true as between the superior oblique of one eye ;m ,1 j j tace the ends Ol the rods and cones give the appearance of a fine mosaic. These organs are connected with the subjacent layers, and, ultimately, more or Diagrammatic section from the posterior portion of the human retina. 1, layer of rods and layer f nuclei 252 THE SENSES. less directly, with the axis-cylinders of the fibres of the optic nerve. The optic nerve pierces the eyeball not exactly at its most posterior point, but a little to the inner side. At the exact centre of the retina that is, the most posterior point of the eye there is a small yellow area (macula luted) with a cen- tral depression (fovea central-is). Here are found none of the fibres of the optic nerve, but a great increase in the numbers of the cones , as well as an increase in their size. Area of most acute vision : If the object looked at is focussed directly upon the macula lutea, the image is then seen with the greatest clearness. This is to be expected, for in the macula lutea we find the end-organs of the optic nerve most highly developed. In every-day life we look directly at an object so as to receive the image on the macula lutea and thus render the perception more acute. Rays of light enter- ing the eye on an angle are focussed on some other part of the retina and are not so clearly defined. Blind spot : If the left eye is covered and the right directed steadily upon the cross in Fig. 125, the circular spot will be FIG. 125. visible at the same time, though less distinctly. As the book is moved slowly backward and forward, a point will be found at which the round spot disappears, reappearing as the book is held nearer or farther or as it is inclined in cither direction and the image is carried away from the "blind spot." The blind spot is the point where the optic nerve enters the eye. If the image be focussed on this point no perception follows, as that part of the retina is deficient in rods and cones (Fig. 126). FIBRES OF NERVES AND TRACTS. 253 FIG. 126. The posterior half of the retina of the left eye viewed from before. Twice its natural size, s, cut edge of the sclerotic ; ch, chproid ; r, retina : in the interior at the middle the macula lutea with the depression of the fovea centralis is rep- resented by a slight oval shade : toward the left side the light spot indicates the colliculus or eminence at the entrance of the optic nerve, from the centre of which the arteria centralis is seen sending its branches into the retina, leaving the part occupied by the macula entirely free (Henle). Optic nerve : If one examines the optic nerves in a super- ficial manner, they will be seen to leave the back of each eye, passing backward through the optic foramina until they reach the body of the sphenoids. Here the optic nerves cross one another in the form of an X (the optic chiasm), the fibres intermingling, and the right nerve apparently passing over to the left side and the left nerve to the right side. The posterior limbs of the X pass backward and are called the optic tracts. The optic tracts in their backward course curve around the crura cerebri to terminate in the ganglion- cells of the pulvinar, anterior quadrigemina, and external geniculate bodies. From the ganglion-cells of the pulvinar, anterior quadrigemina and external geniculate bodies, fibres, called the optic radiations, pass backward to terminate in the ganglion-cells of the cortex of the posterior part of the occip- ital lobes. Fibres of nerves and tracts : A more minute examination of the optic nerves will show that each optic nerve consists of two distinct bundles of fibres laterally placed. The inner 254 THE SENSES. set of fibres being the fibres from the inner half of the retina ; the outer bundle of fibres coming from the outer half of the retina. If we trace these bundles backward to the optic FIG. 127. Left Eye Right Eye Optic ic nerve Optic radiation Optic radiation rlc'x of occipital lobe with cortical cells 1, external geniculate body; 2, pulvinar; 3, anterior quadrigeminate body; 4, in- ternal genioulate body ; 5, commissure of (Judden. chiasm, it is noted that only the inner bundles decussate and pass to the opposite side of the brain. The external bundles do not decussate, but pass directly backward as part BINOCULAR VISION. 255 of the optic tracts to terminate in the ganglia on their respec- tive sides of the brain. Thus the left pulvinar, left anterior quadrigeminate and external geniculate bodies receive the fibres from the inner half of the right eye and the outer half of the left eye. The right ganglia receive fibres from the inner half of the left eye and outer half of the right eye. Commissure of Gudden : There is also in the optic tracts a bundle of fibres arising from the internal geniculate body of one side, passing forward, as a third and innermost bundle of the optic tracts, to the back part of the optic chiasm ; here the fibres bend on themselves and pass backward along the inner margin of the opposite optic tract to terminate in the other internal geniculate body. These commissure! fibres form the commissure of Gudden, and serve to connect the two internal geniculate bodies. They play no part in vision. Nervous mechanism of vision : The image, properly re- ceived on the retina, stirs the rods and cones into functional activity. The u vibrations " are then passed along the optic nerves and optic tracts to terminate in the pulvinares, exter- nal geniculate and anterior quadrigeminate bodies. These structures may be called the primary vision centres. Here the impressions of sight are received in a physical sense, but the mind does not as yet appreciate the sight impression. The impression received by the ganglia are now trans- mitted along the optic radiations to the cortical cells of the occipital lobes. When the cortical cells are stimulated the mind is capable of appreciating that " we see" So we may call the ganglia the seat of physical sight, but the cortical cells are the seat of physiological sight. Binocular vision : In normal vision both eyes are used, so that a separate image is received on each retina. Through the intimate association of the two halves of the brain we are conscious of but one image. A further advantage of binocular vision is that each eye looking at an object from a slightly different standpoint sees a little more on one side or other of the object than does the other eye. Thus the combined image formed contains a little more of the object than would the image from one eye alone. 256 Till': SENSES. Thus the perceptive faculties can judge more correctly of the form and diddnce of an object. The stereoscope illustrates this point. In this instrument two photographs are taken by cameras so placed as to repre- sent the position of the eyes in vision, and the two views of one object are then superimposed by the use of prisms (Fig. 128). Photographic views seen through a stereoscope possess a rounded-out, lifelike appearance not seen in ordinary photo- graphs. Inversion of the image : From the laws of optics we know that the image formed on the retina is an inverted image of FIG. 128. Illustrating the principle of the stereoscope and binocular vision. the object. Yet we perceive the object in its upright posi- tion. This is the result of lifelong habit. A baby sees an object; the next step is to touch it; by practice the child finds out which is the top of the object through the touch- perception. He then corrects his mental impression. Very speedily the brain learns to make the correction, and the les- son once learned lasts through life. The correction is made by the brain in its perception of the image. It is an act of mental and not of physical origin. Tims, objects which are projected upon the left of the retinal surface look to be, as they are, on the right of the body ; and so with all the directions : the inversion of the retinal image is corrected by the mind. Duration of visual sensations: The duration of a visual sen- sation is always greater than that of the stimulus which has caused it. However brief the luminous impression, the effect on the retina lasts about one-eighth of a second. The spokes RETINAL RED. 257 of a rapidly revolving wheel for this reason do no not appear as spokes, but as a solid mass, each following one another so rapidly that one impression cannot fade away before another lias replaced it. Mental processes acting on visual perceptions : We are able to estimate by the aid of the brain the size, direction, distance, form, and speed of motion of a thing which we have seen. All of these are judgments based largely upon previous ex- perience. All of these deductions are liable to error by rea- son of faulty judgment or faulty vision, but this is the usual method of forming such estimates. As an example of visual illusion, the two centre squares shown in Fig. 129 are exactly the same size, yet the white square on the black ground appears larger than the black square on the white ground. Also in Fig. 130, the heavy black lines appear to converge, though in reality they are parallel. Illustration of irradiation , , , (McKendnck). Clearness of vision depends on the space between the cones in the point of clearest vision, the macula hi tea. It has been calculated that an object must subtend an arc of at least 60 to 70 seconds in the field of vision to be clearly seen. Such an object makes an image of about i2ihroth of an inch in the retina ; and this is about the distance between the cones at the macula lutea. Similarly, two points to be clearly distinguished must be separated suf- ficiently to allow this amount of separation in the retinal image. Retinal red : When the retina of a recently-killed animal is examined it is colorless ; but during life or if extracted without exposure to light, it is of a purple-red hue, and the color is found in the rods of the retina. It is derived from the pigment of the outer layer of the retina. It is the " ret- inal red " or " visual purple/ 7 as it is variously named, which one sees in the reflex of the retina. Exposure to light destroys it, and for this reason it was long unknown. It disappears after a brief exposure to sunlight, about half a minute. 17 Phys. 258 THE By throwing a beam of light into the eye by a mirror, as by the ophthalmoscope, a red glow is observed in the pupil. FIG. 130. Nl s s N N S S S N S N S s N S N S L. M S / s / V 1 \ X s / s / s x s x V II I 11 s x V V k. JM II S w 1 % II s s s s s s s s s s s s K K s s s Zollner's figure showing an illusion of direction (McKendrick). This is called the retinal reflex. The red glow is produced by the retinal red. Optograms are pictures which appear upon the retina after exposure to light. They are due to the fact that an exposure to light bleaches the retinal red, leaving it dark in the shaded portions. An optogram may be obtained by the following experi- ment: The eye is removed from an animal in a dark room and kept in a covered box until exposed to a brightly illumi- nated skylight or window for some minutes. The eye is then replaced in the dark room and the retina examined. It will he found that the panes of the window are shown in light color, while the sash is in dark outline. This soon fades on exposure to daylight; but if the retina is dried in the dark, the optogram is much more durable. AFTER-IMAGES. 259 Pigment of choroid and retina : The student must not con- found the " visual purple " with the pigment granules of the choroid and retina. The latter are dark brown or black, and serve to keep the chamber of the eye as a dark box. The black cloth of the photographer serves the same purpose. All optical instruments are painted black on the inside to prevent un desired reflection of light. So with the eye. Color-perception : It is probable that particular rods and cones are capable of responding to rays of light of a certain wave-length, and to those rays alone. It is well known that the rays of red light are of a certain length of vibration. The same is true of yellow and of green rays. We can con- ceive that each primary color has its own set of cones and rods capable of responding to its stimulus, and that by com- binations of these stimuli the complementary colors and varia- tions of shade may be perceived by the resulting stimuli act- ing upon the brain-centres. Such teaching is, of course, speculative, but this is one theory which has acceptance. Achromatism : If in obtaining an image of an object through a double convex lens the lens be too large, there will be seen around the image formed a halo of prismatic colors. This is called a chromatic ring, and is produced by an un- equal refraction of light-rays by the peripheral portions of the lens. The unequal refraction results in a dispersion of the light, so that it is broken up into the seven primary colors. To remedy this defect, we put a "photographic shutter" in front of the lens, thus limiting the entrance of light to the central portions of the lens, where the index of refraction is " constant " or " true." In the eye the iris acts as the pho- tographic shutter, thus rendering the image achromatic. It may, however, be said that there may be a visible band of color seen by some defective eyes where there is considerable fault in the focus of the image on the retina. After-images : It has already been noted that vision lasts longer than the stimulus which excites it. Under some con- ditions it may last a perceptibly long time : it is then known as an after-image. If one looks at an intense light, the sun, the sense of light remains for some time in the eye. Similarly, if one looks intently at a white spot on a black background, 260 THE SENSES. and then turns to a white surface, one has the image of a gray spot. The first of these conditions cited is known as a positive after-image, and the latter as a negative. In the first case the phenomenon results as a continuation of the same sensation, and in the latter a new perception results. These images appear to have the complementary color of the original object ; thus, green excites a reddish after-image ; orange, blue ; and so on. They may be explained as a result of exhaustion. The part of the retina on which the image has fallen becomes tired, and when the eye is turned upon a white ground, the white light coming to the retina docs not produce as much sensation in the tired portion. The colored negative after-images may be similarly explained. Near-point : The " near-point " is the nearest point to the eye at which vision is distinct, the shortest focus of the crys- talline lens. It is usually about five or six inches. Defective eyes : Although hardly within the scope of physi- ology, it may be perfectly appropriate to discuss here some of the more common defects of the eyes depending on abnormal conditions in the optical apparatus. Emmetropic eye : It is the normal eye that is, an eye in which parallel rays or rays from objects at a distance are focussed upon the retina without an effort at accommodation. Such a distance, for practical purposes, is considered to be any point beyond twenty feet. Absolutely emmetropic eyes are not common. Myopia or " near sight " is the term applied to an eye in which the rays from a distance are focussed in front of the retina, and the image is blurred. Such an eye is permanently focussed for near objects (Fig. 132). Myopia is produced in two ways by the antoro-posterior diameter of the eye being too great, or by the convexity of the lens being exaggerated. In either case the focus of the lens will fall in front of the retina. The first condition is essentially a congenital defect, whereas too great convexity of the lens may be either con- genital or the result of disease. Myopia is corrected by the use of a concave Ions which diverges the rays, and in this way prevents their coming to a PRESBYOPIA. 261 focus too soon. Such glasses are seldom needed except for distant vision. Hypermetropia, or "far sight/ 7 is the reverse of myopia (Fig. 131). In hypermetropia the antero-posterior axis of FIG. 131. the eye is too short, or else there is an abnormal flattening of the lens, which does not allow accommodation for near vision. The result is that the image of an object near by is focussed behind the retina ; but objects at a distance are clearly seen. FIG. 132. Hypermetropic eye and myopic eye (far-sighted and near-sighted eye). 1, hyper- metropic eye. The luminous rays arriving from an infinite distance (parallels) produce an ocular cone, the summit of which falls beyond the retina (at A), either because the cone is too long (lack of converging power in the media of the eye), or because the retina is too far forward (the eye being too short) ; 2, myopic eye ; the luminous rays from an infinite distance (parallels) produce an ocular cone, the summit of which falls in front of the retina (at B), either be- cause this cone is too short (excess of converging power in the media), or be- cause the retina is placed too far back (the eye being too long). Bonder's re- searches seem to show that short-sightedness is owing to this latter cause, as is well shown in the figure (the ocular globe being greatly elongated from back to front) (Kuss). Hypermetropia is corrected by the use of convex lenses, which add to the refractive power of the eye. Presbyopia is defective vision due to the loss of power in advanced years. The elasticity of the lens becomes less, and the convexity cannot be increased for near vision. The 262 THE SENSES. ciliary muscle may also be weaker and aid in the production of the error. A weak convex glass commonly corrects the lack of refraction-power. Thus we see that presbyopia is identical with hypermetropia due to flattening of the lens. Astigmatism is a defect in the vision due to the irregularity in the globe of the eye, whereby the diameter in one plane is greater than in another. Thus, the retina may be an uneven surface, and the image focus accurately in one part and falsely in another. In this condition vertical and horizontal lines are not seen with equal distinctness. Astigmatism is corrected by the use of cylindrical or pris- matic glasses, which have to be accurately adapted to the needs of each case. This error, if serious, is usually com- bined with other defects of vision, frequently myopia. Diplopia is the condition which results from a want of har- mony in the eyes, so that the image of each eye is perceived separately ; that is, one sees double. Diplopia is commonly caused by paralysis or spasm in one of the lateral straight muscles, which does not allow the eye to be turned in harmony with the other. If the eyes are turned so that the axes of vision are separated, the condition is known as external strabismus or squint ; if the axes are crossed, it is called internal strabismus or cross-eye. Color-blindness is an inability to perceive some colors. The colors which are usually mistaken are green and red. Fre- quently it is found that a distinction cannot be made between these colors. This is sometimes known as Daltonism. Theories of normal color-perception : Ordinary white light if decomposed is resolved into the seven primary colors violet, indigo, blue, green, yellow, orange, and red. Each of these primary colors has a different wave-length. Other colors than the seven primary colors are the result of the mixture of two or more of the primary colors in various proportions. We are ignorant of the manner in which the rods and cones are made to vibrate by ordinary images, and we are equally ignorant as to the nature of the process that allows the diiVcrcnt color-effects to be conveyed to the optic nerve. THEORIES OF NORMAL COLOR-PERCEPTION. 263 As it is impossible to go fully into the matter in this book, it is hoped that the following will suffice : The different color theories assume that there are different substances in the retina capable of responding to different wave-lengths of light (comparable to a photo-chemical pro- cess). Red, green, and violet (Fig. 133) are the fundamental FIG. 133. Diagram of three primary color-sensations. 1 is the so-called " red," 2 "green," and 3 "violet" primary color sensation; R, 0, Y, etc., represents the red, orange, yellow, etc., color of the spectrum, and the diagram shows, by the height of the curve in each case, to what extent the several primary color-sensations are re- spectively excited by vibrations of different wave-lengths (Foster). colors, and all others may be made from combinations of these three. Working on this basis, Young and Helmholtz assumed three chemical substances in the retina capable of replying to the three fundamental colors. Hering assumed three substances responding respectively to white or " black " (absence of light), red or green, and yellow or blue light. In this theory the white, red, and yel- low rays are katabolic in their effect on their individual re- cipient substances; the "black/ 7 green, and blue being ana- bolic e. g.j the substance responding to white is "broken down " by white light, but is regenerated ("built up") by "black light" i. e., absence of light. Mrs. Franklin assumes in her theory that in early life the eye possesses no color-per- ception, but merely perception of luminosity i. e., white or 264 THE SENSES. black. The substance responding to luminosity she calls " gray-perceiving." As the development progresses, some of the " gray " is differentiated into a blue- and a yellow-per- ceiving substance. The yellow-perceiving substance is still further differentiated in the course of development into a red- and a green-perceiving substance ; thus : Gray. I Blue. Yellow. I Green. Red. Many objections have been raised against each of these three color-theories, and it is true that each leaves some points unexplained ; but Mrs. Franklin's is, so far, the best, and more readily explains the peculiarities of color-blindness. Causes of color-blindness : The followers of Helmholtz and Hering say that color-blindness is due to an absence of one or more of the fundamental color-perceiving substances. Mrs. Franklin's theory assumes a lack of full development or complete absence of development of the " gray "-perceiv- ing substance. Thus, in a case of absolute color-blindness the " gray " substance has undergone no development, and the individual sees everything as without color, but in different shadings of light or darkness. To such a person a highly colored paint- ing would look like a black-and-white etching. Again, working on the development theory, we may assume that the gray differentiated into "blue"- and " yellow "-per- ceiving substances and there stopped. Clinically we find in- dividuals capable of distinguishing blues and yellows, but reds and greens are unknown to them. Color-blindness in different sexes: Males are far more liable to be color-blind than female (16 to 1). Only about one woman in four hundred is color-blind. The reason for the preponderance of color-blind men may be accounted for on the theory that although the differentiation of the "gray"- perceiving substance into red, green, and blue is a natural TEST FOR COLOR-BLINDNESS. 265 process, nevertheless it can be perfected by practice and color- education. Such an education is given early in life to little girls in matching colors for doll's clothing, etc., whereas, it is neglected in boys. Importance of the defect : In marine and land locomotion, red and green signals are used to indicate opposite conditions, and the failure to distinguish them has frequently been the cause of serious accidents. Test for color-blindness : By laying a number of skeins of yarn of various colors in a heap, and requiring the person to be tested to select all resembling a certain skein from the heap. EMBRYOLOGY. Embryology deals with the reproduction and development of individuals, whether of animal or plant life. AVe shall first study how the parent gives origin to the offspring, and later on the development of that offspring until it becomes a fully developed member of its species. REPRODUCTION. By species we mean a class of organized beings in which the individuals composing it die off, but which nevertheless repeats itself and maintains its complement by the continued accession of similar forms. Heredity is the inherent property of the individual by virtue of which the individual is of the same species as the parent, and furthermore has certain individual characteristic's of the parent. We are human beings because our parents were human beings, and we inherit the form and character- istics of the species. Still further, we may have certain tricks of speech, a peculiar gait or method of thinking, in- herited from our individual parents. Reproduction is the process by which a species is perpetu- ated, notwithstanding the limited existence of the individual members. The law governing reproduction is that the young are of the same kind as their parents. By this law, which i- s<> commonly observed as to seem a truism, is maintained the anatomical identity of individuals of a species, as well as the physiological fact of an unbroken continuance of the species by reproduction. Methods of reproduction : Reproduction takes place by one 266 THEORY OF REPRODUCTION. 267 of two entirely distinct methods asexual reproduction and sexual reproduction. Asexual reproduction is the usual method among plants and animals whose organism consists of but one cell. It is also the method in some of the lower multicellular animals and many higher forms of plants. When considering the amoeba it was stated that it sub- divided, with the result that from the single cell we had two distinct units, each being members of the same species. No sexual relations have taken place, and the offspring is the child of but one parent. Sexual reproduction : Sexual reproduction consists of the union of elements produced separately by the female and the male. In some of the lower organisms there is no direct output from the male and female, but the two parents fuse together, and the resultant mass develops into the offspring ; such a method is called " conjugation." As we rise in the animal scale we find that the female produces the ovum, or egg, which is capable of being devel- oped into a living offspring only when it is fecundated or im- pregnated by the seminal or spermatic element from the male. In some animals (worms) both the male and female ele- ments exist in the same individual ; but still the offspring is the result of " sexual reproduction," although there is but one parent. Theory of reproduction: Just why a cell should subdivide into two secondary cells, or why there should be the necessity for sexual reproduction, is a difficult problem to answer. Asexual generation may be explained according to certain principles laid down by Herbert Spencer. Spencer pointed out conclusively that the mass of a cell grows as the cube, whereas the surface grows only as the square ; from this it follows that the mass to be nourished soon outgrows the ab- sorbing or nourishing surface, hence fission results to insure a relative increase of surface to the mass. Such division of cells goes on for a variable number of times, until finally the cell-protoplasm becomes enfeebled and worn out. In certain infusoria the number of divisions may be from 150 to 450; after that conjugation is necessary for a continuance of the 268 REPRODUCTION. method. From such observations one may reason that sexual reproduction insures a fresh conjugation of protoplasm for each individual, thus keeping the standard up to the maxi- mum. Also sexual reproduction tends to preserve the type of the species ; for if the individual were the offspring of but one parent, certain characteristics of that parent would be exag- gerated in the course of several generations; but as the individual springs from two parents, it becomes the menu between those parents, with the result that the individual tends to become the composite of the entire species. Fecundation : The junction of the male and female delimit* in sexual generation is called fecundation. At the time when the male and female discharge their respective elements it is not necessary that the parents should be in juxtaposition. In deep-sea fishes the female discharges her spawn ; the male, which is destined to fecundate these ova, may be miles away and discharge his spermatozoa into the ocean ; the male and female elements may float about until finally they meet, with the result that the ova become fecundated. In fishes like trout, salmon, etc., the female lays her eggs on the spawning-bed, and later on the male deposits his ele- ments directly on the spawn. Still further along in the scale we find juxtaposition essen- tial, the male grasping the female and both discharging their elements at the same time, and the ova are fecundated as soon as they leave the female. Of course, in " viviparce " not only is juxtaposition essential, but there is an actual discharge of the male elements into the cavity of the female generative organs (such an act is called " copulation "). Human Female Organs of Generation. The female organs consist of two or (trie*, in which the ova are formed, and their oviducts or Fim//m>/r// and lactation. Duration of menstruation: The menstrual flow lasts on an average five days, the amount of blood lost gradually in- creasing until the third day, and then gradually diminishing. Nature of menstruation: At the beginning of menstruation there is a general congestion of the generative organs, in- cluding the breasts. A few days before menstruation there LIFE-HISTORY OF THE UNIMPREGNATED OVUM 275 has developed a hypertrophy of the mucous membrane (su- perficial layers) of the uterus. At the beginning of the menstrual period this hypertrophied mucous membrane is gradually shed, leaving its underlying vessels exposed, and they bleed. At the end of menstruation a new mucous mem- brane is developed. The shed mucous membrane is called the "decidua menstrualis." During menstruation, especially t the beginning of the period, the woman is apt to be peevish, irritable, complains of headache, loss of appetite, and a sense of pelvic oppression that may even amount to severe pain. Character of menstrual discharge : It is a thin, bloody fluid of a dark color and having a peculiar odor. It consists of blood, epithelium, and mucus from the uterus and vagina, to- gether with the decidua menstrualis. The blood does not clot. Ovulation : The commonly accepted theory is that about or shortly before the age of puberty the Graafian follicles begin to discharge their ova, and that this process continues until the menopause. Doubtless many Graafian follicles only par- tially develop and atrophy without discharging mature ova. The frequency with which well-developed ova are discharged is the subject of much dispute. The most conservative view is that there is one mature ovum discharged for each men- strual epoch. On the other hand, other embryologists claim that there is an almost daily discharge of ova, but only a few enter the Fallopian tubes, while the rest are lost in the peri- toneal cavity. Whichever view the student cares to follow is immaterial, but nevertheless we must agree that only about as many mature ova reach the uterine cavity as there are menstrual epochs. Life-history of the unimpregnated ovum : On leaving the ovary and entering the Fallopian tube the ovum is surrounded by a few cells derived from the discus proligerus of the Graafian follicle. These cells may serve as a source of nourishment, but soon disappear. There are now certain changes to be observed in the ovum itself; first, the vitelline substance seems to contract slightly, so as to leave a narrow rim or space between the vitellus and zona pellucida the peri vitel- line space. 276 REPRODUCTION. At this time the nucleus or germinal vesicle undergoes a karyokinetic division into two parts, one part then being ex- pelled from the ovum proper into the peri vitel line space as a "polar body" the remaining portion of the original nucleus, again subdividing into two parts by karyokinesis, with the protrusion of one part as a second " polar body." What is now left of the original nucleus is the "female pronudeus " ; the polar bodies are apparently useless. Up to this point all ova develop alike, and now are fit for impregnation. If im- pregnation does not take place, the ovum dies and is caM nil'. If impregnation does occur, there is further development of the ovum, as we shall see in a later paragraph. Corpus luteum : After the escape of an ovum there is an effusion of blood into the cavity of the Graafian follicle. The clot which follows is disposed of by the same retrogressive processes which extravasated blood may undergo in any part of the body. The serum is absorbed, the cells disintegrate. and the coloring-matter is in part taken up by the tissues and in part crystallizes or takes up other constituents, and pre- sents variations of coloring. Hand in hand with these changes in the blood go important changes in the surround- ing tissues. The epithelial cells which are left behind pro- liferate and form a sort of yellowish, very vascular tissue, which presently undergoes fatty degeneration. This yellow mass surrounding and enclosing the remains of the extrava- sated blood constitutes the corpus ufeum,and as it disappears its place is occupied by a dense, firm connective-tissue cica- trix, which may be pigmented. The changes described above are those that take place if the ovum fails to become impregnated. If, on the other hand, fecundation ocvurs, the corpus luteum undergoes certain characteristic changes, widely differing from the degenera- tion of the corpus luteum of non-pregnancy. If the ovum he impregnated, the corpus luteum then does not degenerate and disappear rapidly as after menstruation, but continues fully as large as at the beginning for several months, and at the end of pregnancy still remains as a clearly marked body. This is shown in the following table from Dalton : MALE GENERATIVE ORGANS. 277 At the end of three weeks. One month. Two months. Four months. Six months. Nine months. Corpus Luteum of Menstrua- tion. f inch in diameter ; central pale. Smaller ; convoluted wall bright yellow ; clot still reddish. Reduced to the condition of an insignificant cicatrix. Absent or unnoticeable. , \ Corpus Luteum of Pregnancy (Dal ton). lot reddish ; convoluted wall Larger ; convoluted wall bright yellow ; clot still reddish. | inch in diameter ; convo- luted wall bright yellow ; clot perfectly decolorized. Size about as at two months; clot pale and fibrinous; convoluted wall dull yel- low. Absent. Still as large as at the end of the second month. Clot fibrinous. Convoluted wall paler. Absent. inch in diameter ; central clot converted into a radi- ating cicatrix ; external wall tolerably thick and convoluted, but without any bright yellow color. Connection between ovulation and menstruation : Whether ovulation depends upon menstruation or menstruation upon ovulation, or whether either has any connection with the other, is a matter of lengthy controversy. However, the general view is that both ovulation and menstruation are the result of a common cause (cause un- known), but either may exceptionally occur without the other. It is generally accepted that ovulation, or discharge of yum from a Graafian follicle, takes place a few days before e onset of the menstrual period. Human male generative organs. The generative organs of the male consist of the two tes- icks, which produce the seminal fluid ; and the vas deferens, r duct leading from each to join with the duct of the corre- ponding seminal vesicle, in which the secretion is stored (?) until it is discharged through the penis and the prostate gland. 278 REPRODUCTION. FIG. 138. Testicles : Each testicle is made up of a dense connective- tissue framework and a secreting portion. The connective- tissue stroma, tunica albuginea, surrounds the outside of the organ, and sends incomplete partitions into the central por- tion of the organ, dividing it into a number of communicating cavities. In these cavities are winding tubules which con- stitute the secreting portion of the organ. These tubules in- osculate in a sort of mesh (rete testis), and finally all unite in the epididymis. The secreting tubules are called the semi- niferous tub lit ex. Each tubule has, in the active organ, a limiting membrane, upon which are a number of layers of flattened cells. In- ternal to these are seminal cells in two or more layers. The seminal cells contain nuclei which are capable of division, so that each nucleus may develop several new nuclei. The nuclei are the spermatoblasts, or cells from which the sper- matozoa originate. The cells before the di- vision of the nuclei resemble the ordinary cuboid epithelium, and it is in the superficial layers (i. e., toward the lumen of the tubuli) that this function of the cells takes place. The seminiferous tubules all converge to- ward the epididymis, a tortuous tubule which is lined with mucous membrane, and lies beside the testis in a long, convoluted mass which may be unravelled, and is found to be about 20 feet long. This empties contents, or rather continues on, into the va# dejer- ens, which conveys the semen to the junction with the seminal vesicle. During this passage the mucous membrane adds a viscid mucous secretion in which the spermatozoa are liber- ated and, so to speak, diluted. Spermatozoa : In the seminiferous tubules the developing spermatozoa may be seen with niMciii- i.irre; d, the heads all united in the cells from which tails; e, end-piece ,1 ,1 , i , of the tail. they arise, the tails projecting the cavity of the tube. Bu; separated. They then consist of a head and a THE URETHRA. 279 In length they are about ^th to -A^th of an inch. The head is somewhat elliptical and the tail gradually tapers. In other animals than man the size and form vary from those of man, though in a general way they conform. There is a very active vibratory motion of the tail of the spermatozoon, which allows it quite free motion in a fluid medium. It is by this swimming motion, in which it may be compared to a tadpole, that the seminal cell is able to reach the ovum against the action of the cilia in the uterus and Fal- lopian tube. Seminal vesicles : They are tubules which join the vasa deferentia, and lie upon the base of the bladder, emptying into the urethra by the ejaculatory ducts through the prostate gland. In structure the vesiculse seminales are convoluted and dendritic. They are lined by a mucous membrane and are con- voluted and folded so as to present a sacculated appearance. Prostate gland : It is a gland lying at the base of the bladder and surrounding the urethra at its beginning. It has the general structure of the glandular organs, and in addi- tion a considerable amount of muscular tissue. Its acini empty into ducts which empty into the urethra. Its function is not exactly known. The penis consists of three more or less cylindrical bodies of erectile tissue enclosed in fibrous sheaths. The two cor- pora cavernosa lie above, and receive between them, below, the corpus spongiosum, in which the urethra is contained. The glows penis is continuous with the corpus spongiosum. The covering of the penis is of loose skin, but over the glans penis and lining the prepuce it resembles mucous membrane. In this region there is an abundant subcutaneous nerve- plexus, and the Pacinian bodies are quite numerous, so that it is possessed of acute sensibility. The urethra extends from the bladder through the corpus spongiosum to the end of the penis. It is lined with mucous membrane, and is furnished in its deeper layers with numer- ous muscular fibres. There are a number of ducts of glands it whose function is not fully understood, though m is supposed to be added to that of the seminal nake up the semen. 280 REPROD UCTION. Erectile tissue of the penis : The erectile tissue consists of a system of distensible vacuoles containing venous blood, lying in the interstices of a fibrous connective tissue. The erector penis muscle by its contraction compresses the veins of the organ, and the veins become turgid with blood. The arteries enter the structure of the erectile tissue along the pubic bone, and are not pressed upon by the contraction of the muscle. Impregnation in the Human Species. Impregnation, or fecundation, are the terms applied to the junction of the male and female elements (spermatozoon and ovum), by virtue of which the joined elements become one, and by developmental growth eventually become an indi- vidual of the species. If no junction of sperm atazoon and ovum takes place, each dies and is cast off. If, however, the ovum is fertilized by the spermatazoon, a new life is begun. Methods of fertilization : The methods by which the male spermatazoon and ovum are brought in contact vary for dif- ferent classes of animals. As has already been mentioned, in some classes of fishes the female ejects her spawn and the male his elements, at different places, and owing to the ocean currents the male elements are brought into contact with the female elements. In frogs the male fertilizes the ova just as the latter leave the female. In mammalia the male intro- duces the spermatozoa into the genital passages of the female, and this act is called coitus. Coitus : The act of coitus is preceded by a preliminary period of sexual excitement, more marked in the male, during which the penis becomes swollen, turgid, and erect. The penis is then introduced into the female vagina. As a result of muscular movements there is friction upon the delicate sen- sory nerve end-organs of the glans penis and clitoris, pro- ducing intense nervous sensations, leading finally to a climax of excitement on the part of both male and female that gradually fades away. During the climax (or orgasm) there is an ejaculation <>i' seminal fluid on the part of the male into the upper end of the vagina of the female. There DETAILS OF IMPREGNATION. 281 is also a flow of secretion from the glands of Bartolini of the female, and also, presumably, a rhythmical opening and clos- ing of the cervical canal. Site of impregnation : Although the seminal fluid of the male is lodged, at the close of coitus, in the vagina of the female, the spermatozoa are not yet in contact with the ovum. By virtue of their inherent mobility the spermatozoa travel along the cervical canal into the cavity of the uterus, there to meet the ovum. The spermatozoa may even travel along the Fallo- pian tubes, impregnating the ovum in the lumen of the tube ; or may even leave the tubes by their fimbriated ends and impregnate the ovum on the surface of the ovary. Time of impregnation : Probably immediately before the menstrual period ; but owing to the fact that both the female and the male elements may remain in the genital passages of the female for some days in a healthy condition, it is difficult to fix the time of actual impregnation of the ovum. Impregnation is generally supposed to occur a short time before menstruation, on the following grounds : (1) It is probable that in most instances the rupture of the Graafian follicle occurs just before the menstrual period. (2) The uterus is in the most favorable condition to sustain the fecund ovum at that time, because of the presence of the decidua menstrual is. (3) Among the Jews, a remarkably prolific race, coitus is prohibited by the religious law for a week after menstruation. Details of impregnation : The spermatozoon travels along the uterus or Fallopian tubes until it comes in contact with the ovum. The spermatozoon, by lashing its tail, wriggles through the zona radiata (pellucida) of the ovum and enters the perivitelline space. Several spermatozoa may succeed in effecting an entrance as far as the perivitelline space ; but be- yond this, for some unknown reason, but one spermatozoon goes further and enters the vitellus proper ; the others die. The spermatozoon destined to fertilize the ovum now loses its tail, and the head and centre-piece are called the male pro- nucleus. The male pronucleus and female pronucleus now fuse together in the centre of the egg and form the first seg- mentation-nucleus. DEVELOPMENT. On the formation of the first segmentation-nucleus the act of imi>r<- i iw. with the allantois luiiy (see below) has already become tutted Sm^'SjSi with capillary loops, and has estab- lished a connection with the decidua of maternal growth. As the vessels of the allantois communi- cate more and more with the chorion, the embryo derives more of its sustenance from the mother, and the remains of the yelk-sac (umbilical vesicle) dwindle as the need is less and the substance is consumed (see Figs. 143, 144, 145). Subsequent history of the allantois : After the development of the placenta the allantois dwindles away, all except the por- tion nearest the foetus ; this becomes the urinary bladder and the urachus (the latter, in the adult, is an impervious cord extending from the bladder to the umbilicus). Chorion : the chorion is the outer zone of the ovum, and from its surface there project centrifugally a large number of villi. The chorion consists of several layers which fuse into one vascular membrane: the allantois, the outer layer of the amnion, and the vitelline membrane are united in the chorion. As the embryo develops the vessels of the chorion become thinner on the side toward the uterine cavity, and more dis- tinct on the opposite side. This change continues as the ASSOCIATION OF FCETAL AND MATERNAL BLOOD. 289 embryo increases, until the placenta is formed by the branch- ing of the embryonic vessels and the increase of the decidua at the corresponding point. Association of decidua and chorion : In the deeper part of the mucous membrane of the uterus, at the implantation of the chorion, there are hollowed-out spaces or sinuses in the tissues, which communicate both with a maternal vein and an artery that is, special arrangements are made for the rapid circula- tion of a large amount of blood in the uterine mucous mem- brane at the placental site. At the same time the glandular structures of the uterine mucous membrane are increased, and the follicles run deeply into the thick and succulent tissues. The villi dip down and develop new tufts of capillaries in the deepened crypts of mucous membrane, so that the tufts of capillaries of the chorion may be said to resemble in a way a glove filled with foetal blood dipping into a vessel filled with maternal blood. The placenta : So far the villi throughout the entire surface of the ovum become associated with the decidua. Now, as development advances, that portion of the chorionic villi that is lined by the central portion of allantois grows still further in size, while the rest of the chorionic villi atrophy and disap- pear. The loops of bloodvessels of the allantois project into the villi of the chorion (like a finger into a glove), and thus are brought into closer contact with the blood of the decidua (maternal blood). The portion of villi of chorion containing the vessels of the allantois is the placenta. Thus we see that the placenta consists really of two parts a foetal portion, consisting of fcetal bloodvessels contained in the chorionic villi ; and a maternal portion, consisting of follicles or depressions in the uterine wall, surrounded by bloodvessels so enlarged as to form blood-chambers (uterine sinuses). The chorionic villi are lodged in the uterine follicles, or between the follicles, according to some observers. Association of foetal and maternal blood : The foetal blood never comes directly in contact with the maternal blood. The two blood-currents are nearest each other while circulat- ing through the placenta. While in the placenta there are four layers of cells between 19 Phys. 290 DEVELOPMENT. the maternal and the foetal blood : 1, wall of chorion capil- lary ; 2, cells of chorion; 3, cells of uterine follicle; and 4, wall of the uterine sinus. Interchange between foetal and maternal blood : Although there is no direct communication between the blood of the foetus and that of the mother, nevertheless while passing through the placenta there is an exchange by osmosis between the mother's blood and the foetal blood. The mother's blood furnishes to the foetal blood food and oxygen, and in turn removes the carbonic acid and excre- mentitious material which the foetus must lose that is, the placental circulation supplies the place taken in after-life by the alimentary and the respiratory tracts. Umbilical cord : The umbilical cord or funis is the stalk connecting the placental with the foetus, and its purpose is to l>e a framework or support for the Fro. 146. umbilical arteries and veins. The tissues of the umbilical cord are formed from the vascular allantois, which carries the arteries and veins. It also has an external coating of the amnion and the shrivelled umbilical vesicle and its duct (Fig. 146). How this occurs will readily be seen by reference to the accom- panying cut. Human embryo and its envel- \, n opes at the end of the third At Iirst two arteries COI1VCV SenfoV'Smnio 1 ;' 6nlarge " the blood from the foetus to the placenta, and two veins carry the purified blood from the placenta to the foetus. Later on in foetal life one of the veins is lost. At term the umbilical cord is a rope-like structure between 18 inches and 36 indies long, and consists of one vein and two arteries spirally twisted ; the vessels are enclosed in a layer of jelly-like substance (Wharton's jelly), and the whole is surrounded by a con- nective-tissue sheath. The cord is about one-half inch in diameter. Destiny of placenta : The foetal part is almost all, excepting some of the capillary tufts which are torn off, discharged in PLACENTAL CIRCULATION. 291 the after-birth ; but the decidua is not entirely disposed of in this way, the portions remaining being in part absorbed and in part found in the lochia which occur for a fe\v days after the birth. When the placenta is expelled a part of the maternal tissue is left behind, and there is, of course, a loss of the blood contained in the uterine sinuses, but the general bal- ance of the circulation is not disturbed at childbirth. The reason for this is the oblique entrance of the placental vessels. They enter the sinuses at an angle, and are therefore com- pressed by the muscular tissue of the uterus in its contracted state. Appearance of placenta : The placenta at term appears as a thick, cake-like disk of vascular tissue. Its maternal and foetal portions are so intermingled that they cannot be sepa- rated. In size it covers about one-third of the uterine wall. Period of placental formation : The placenta is formed at about the third month of pregnancy. Before that time the chorion is covered by the decidua reflexa and nourishes the embryo, but as the placenta becomes more developed other parts of the chorion atrophy. Foetal Circulation. Types of foetal circulation : One meets with two distinct types of circulation in foetal life : the vitelline circulation and the placental circulation. In both types the blood is the circu- lating medium driven on by the heart, the essential difference being the site where the foetal blood is enriched. The vitel- line circulation precedes that of the placenta, and as soon as the latter is formed the former disappears. Vitelline circulation : In the vitelline circulation the vessels from the foetus (omphalo-mesentric vessels) pass over the yelk-sac and carry nutrition to it from the growing organism. The vitelline circulation in the human ovum is not very long- lived, for the chorion is early formed and the stock of nutri- ent protoplasm in the yelk-sac is very small. Placental circulation : In the placental circulation the foetal 292 DEVELOPMENT. blood is purified and enriched by osmosis with the maternal blood in the placenta. Development of circulatory apparatus : Among the earliest changes in the blastoderm, occurring in the second week of impregnation, is the formation of bloodvessels and blood- corpuscles. This occurs by the proliferation of certain branched cells of the mesoblast, and FIG. 147. these cells form a closed system of branching capillaries, their nuclei acquiring a red color and becoming the blood-corpuscles. This area is external to but connected with the embryo, and is known as the " area vasculosa." The area vasculosa extends all about the blastoderm upon the sur- face of the vitellus, and as the folds of the embryo occur the vessels are brought to enter the body through the space at which the vitellus is shut in to form the primitive gut. There are then two arteries and two veins which are known as the omphalo- mesenteric vessels. This form of circulation soon gives way to the placental, and the vessels passing to the umbilical vesicle waste, those belonging to that portion of the original vitelline cavity which forms the intes- tine, becoming the mesenteric vessels. Formation of heart : About the time of the formation of the area vaseulosa certain cells of the visceral layer of the incsoblast (splanchnopleure) develop a tube upon each side of the body, and these two tubes soon coalesce to form a single tube (Fig. 148), which receives two veins at its lower end and gives off two arteries at its upper. This is the primitive heart, and pulsations begin in it very feebly almost as soon as there is a trace of the originating cells. This structure soon develops a muscular tissue and a circulating fluid which shortly presents the character of blood. The heart then bends on itself so as to assume a U-shape, Diagram of embryo and its vessels, showing the circu- lation of the umbilical ves- icle, and also that of the allantois, beginning to be formed. COURSE OF THE FCETAL CIRCULATION. 293 which shortly is twisted in such manner that the arterial end of the heart crosses in front of the venous (Fig. 149), and the loop suggests the outline of the ventricles. In the next stage of development the septum between the ventricles grows, and separates the heart into two divisions ; and at about the same time the auricles are developed and the valves become well marked. These changes occur in the fourth to the eighth week of embryonic life. FIG. 148. FIG. 149. FIG. 150. FIG. 148. Earliest form of the foetal heart. 1, venous extremity; 2, arterial ex- tremity. FIG. 149. Foetal heart bent upon itself. 1, venous extremity ; 2, arterial extremity. FIG. 150. Foetal heart still further developed. 1, aorta; 2, pulmonary artery; 3,3, pulmonary branches ; 4, ductus arteriosus. Characteristics of placental circulation : After the cessation of vitelline circulation the needs of the fetus are supplied by the placental circulation until birth. The placental circula- tion presents two prominent features in which it diifers from adult circulation : (1) In the arterial circulation some conditions of the heart and great vessels are necessary to modify the pulmonary cir- culation before the air enters the lungs at birth. (2) In the circulation of the liver the veins present modifications to allow for the return placental circulation. Course of foetal circulation : Ductus venosus : The foetal blood, purified and enriched in the placenta, passes, by the umbilical vein in the umbilical cord, to the umbilicus. The umbilical vein then courses from the umbilicus to the under surface of the liver ; here the vein divides into two parts. One portion of the blood enters into the liver-substance, and after traversing its capillaries is poured out by the hepatic 294 DEVELOPMENT. veins into the inferior vena cava. The other portion of blood passes directly from the umbilical vein into the inferior vena cava by means of a blood-channel, the ductus venom n*. Foramen ovale : The blood in the vena cava inferior empties into the right auricle of the foetal heart. Instead of passing from the right auricle into the right ventricle, as is the case in the adult circulation, the blood from the inferior vena cava passes from the right auricle into the left auricle by means of an opening in the interauricular septum. The opening is called the "foramen ovale" The flow of blood from the inferior vena cava through the foramen ovale and into the left auricle is facilitated by the fact that the inferior vena cava points almost directly into the foramen ovale. The Eustachian valve also favors this peculiar course of the blood. The Eustachian valve consists of a crescentic fold of fibrous tissue covered with endocardium. The fold extends from a point between the opening of the superior and inferior venae cavse over to the lower and anterior margin of the fora- men ovale. The base of the fold lies on the right auriculo- ventricular ring, and the concavity of the fold is directed upward. From its position the Eustachian valve acts as a guiding-groove or gutter for passing the blood from the in- ferior vena cava to the foramen ovale. Ductus arteriosus : On entering the left auricle, the blood is passed into the left ventricle and thence into the aorta, to be distributed all over the body ; but principally to the head and upper extremities. The blood from the head and upper extremities returns to the heart by the superior vena cava. On entering the right auricle the blood from the superior vena cava passes in front of the stream that flows from the inferior vena cava to the foramen ovale, and enters the right ventricle. The direction in which the superior vena cava points (to- ward the auriculo-ventricular ring), and also the Eustachian valve, arc the factors that determine the separation of these two streams. On entering the right ventricle the blood from the superior vena cava is forced into the pulmonary artery toward the lungs. Before reaching the lungs this blood meets with a channel of communication between the pulmonary EFFECTS OF FCETAL CIRCULATION. 295 artery and the aorta. Into this channel (ductus arteriosus) the larger portion of the blood in the pulmonary artery enters and mingles with the blood of the aorta ; the remainder passes along the pulmonary artery to the structure of the lungs, which it nourishes, and thence back to the left auricle by means of the pulmonary veins. Hypogastric, arteries : The blood in the aorta that comes from the left ventricle passes, as has been already stated, largely to the head and upper extremities. The blood in the aorta that enters from the ductus arteriosus largely passes into the descending aorta. On passing down the descending aorta, some of the blood enters the mesenteric arteries, and thence back to the venous circulation by means of the portal vein and the liver. Some of the blood enters the iliac arteries and nourishes the lower extremities ; but the major part of the blood leaves the foetal body by the hypogastric arteries. The hypogastric arteries are branches of the internal iliacs and course along the abdo- men to leave the foetal body at the umbilicus. On emerging from the umbilicus, the hypogastric arteries change their name, and are now known as the umbilical arteries, and form part of the structure of the umbilical cord on their way to the placenta. Effects of arrangement of ftetal circulation : The liver, re- ceiving the freshest blood (from the umbilical vein), is the best nourished of all the organs of the foetus. The result is that the foetal liver is vastly larger in proportion than the adult liver. Also the circulation of the blood is made more perfect, for the branches of the aorta given off to the head and upper extremities distribute blood from the inferior vena cava ; while the ductus arteriosus, carrying the blood from the superior cava and right ventricle, enters the aorta in such a way that most of its blood is sent to the lower extremities and abdominal organs and umbilical arteries. In this way the deoxidized blood is sent back to the placenta for the renewal of its oxygen. The lower extremities are less well developed than the upper. There are probably two reasons for this : (1) the blood is less well aerated and less nutritious ; (2) the in- ternal iliac arteries, giving off the umbilical arteries, probably 296 DEVELOPMENT. divert a considerable portion of the blood-supply of the ex- ternal iliacs which go to the lower extremities. In addition, we note that, owing to the ductus arteriosus, only a little blood goes to the lung-tissues. The amount is quite sufficient to keep up the nutrition of the lungs, and as they have no function before birth they do not need a la rue blood-supply. Change from foetal to adult circulation : The respiratory cen- tre in the medulla, which has been quiescent because it has been sufficiently well supplied with oxygen, is awakened as soon as the connection with the uterine sinuses is interrupted. As soon as the supply of oxygen sinks to a certain point, an impulse of inspiration is generated, and the infant breathes and the lungs assume a condition of partial expansion. With the diminished resistance in the expanded lungs the amount of blood in the pulmonary circulation increases, and, the amount passing through the ductus arteriosus diminishing, this is soon obliterated. At the same time, the blood return- ing to the left auricle increases in quantity, and the intra- auricular pressure is greater; then, too, the inferior venacavu sends less blood, for the ductus venosus no longer carries the blood from the placental circulation, and, therefore, the fora- men ovale is not used, and is soon closed by the adhesion of its valve-like curtain. Thus, we have the adult circulation established in the place of the foetal in consequence of the respiratory movements. Also, owing to the division and occlusion of the umbilical cord, blood no longer passes through the umbilical vessels, with the result that the umbilical vein degenerates into a fibrous cord (round ligament of the liver). The hypogastric arteries remain pervious for the first part of their 'course, as the superior vesicle arteries ; but the rest of their course is obliterated ; they degenerate into fibrous cords. Development of vertebral column: Early in the development of an embryo there is formed, beneath the medullary groove, in the mesoblast a thin thread of soft cartilage known as the chorda dorsalis, or iwtorhorrf. This soon becomes in- cluded in a sort of fibrous sheath, and is the primary axis DEVELOPMENT OF THE FACE. 297 around which the bodies of the vertebrae are developed. On either side of the notochord are developed small centres which subsequently split (mesoblastic somites). These are the protovertebrce. From these are developed the vertebrae and the heads of the ribs by the inner part; and by the outer (or posterior) part the muscles and skin of the back, the epidermis being derived from the epiblast. The vertebrae are not formed by direct ossification of the protovertebrse, but they separate in such a way that adjacent protovertebrae each contributes half to the vertebra formed. That is, two protovertebrae form parts of two vertebrae, one above and the other below, and also form a whole vertebrae by their adjacent portions. Development of cranium : The cranium is developed from the prolongation of the tissues over the protovertebrae to the cephalic end of the embryo. Here it develops three seg- ments, corresponding to the three primary vesicles which are the forerunners of the brain. These centres of ossification are at the base of the skull, the bones of the vertex being de- veloped from membrane. Development of the face : At the head fold of the embryo the mesoblast does not split into two parts, as below, but folds in from the side, covered without and within by the epi- blast and hypoblast. These folds develop certain clefts from which the face is derived, the mesoblast furnishing the bone and muscle structures, and the epiblast the epidermis, while the hypoblast gives the mucous membrane which lines its cavities. Immediately below the anterior cranial vesicle there occurs on either side a cleft in the lateral fold of the embryo extend- ing to the vesicle for the eye. In the space of this cleft there is developed a sort of secondary cleavage of the parts, which by the rapid growth of the parts included between the clefts resembles a budding (Fig. 151). It is by the growth of these buds or processes that the outline of the face is formed. From each side sprouts the superior maxillary process, and the proc- esses unite in the median line, and with the nasal or inter- maxillary process from the upper border of the cleft. The portion below is cut off by a branchial cleft, below which is the mandibular process which forms the lower jaw. 298 DEVELOPMENT. When the processes do not unite as they should, various defects occur ; most common are those about the mouth viz., cleft palate and hare-lip, by failure of the superior maxillary j n-ix v-.-es to unite or by failure of the intermaxillary process to unite with the maxillary. The other branchial clefts do not persist in later life. They become closed as they accomplish their use in developing cer- tain organs: as pathological factors, however, we are often convinced of their non-union or of flaws in their develop- FIG. 151. Development during first month. ment, cysts and tumors of various kinds and certain fistula being attributable to this cause. Development of the extremities : They develop as buds from the somatoblast early in foetal life, and the formation of the joints by cleavage, and lesser details of structure are gradu- ally worked out. At about the third month the separation of the fingers and division of the extremity into joints is about completed. The arm develops somewhat in advance of the leg, and grows rather more rapidly in the earlier period of intra-uterine life. Formation of the spinal cord: It will be remembered that the medullary canal encloses in its cavity cells from the epi- blast which line it. These cells by proliferation and differ- entiation develop nerve-cells and nerve-fibres, the latter at first not medullated. The cells also gradually close in upon the medullary canal, and form a central lined with epithe- lium, a layer of nerve-cells (gray matter) and a layer of nerve-fibres (white matter). DEVELOPMENT OF THE BRAIN. 299 When the spinal cord first appears it fills the entire spinal canal, but at the time of birth the cord has apparently not grown so rapidly as the vertebral column, for it then ends at the third lumbar vertebra, and in the adult it ends at the first. Thus we are able to explain the apparent origin of the spinal nerves above their point of exit from the canal, and the increasing obliquity of the nerves from above down, until finally, in the tuft of vertical nerves below the extremity of the cord, we see the extreme degree of this peculiarity. Development of the spinal nerves : The axis-cylinders arise from cells of the epiblast lining the medullary groove. Before the closure of this groove to form the medullary canal an offshoot from the epiblast may be observed, which is the source of the posterior nerve-roots ; and they become attached to the cord as it develops. The anterior roots spring from the cord after it has developed fibres. The two roots then join and the nerve grows out into the mesoblast. Development of the cranial nerves : In much the same way the cranial nerves arise primarily, except the nerves of spe- cial sense. In function the motor nerves seem to form a sort of anterior root for the sensory, so that they may be arranged in pairs corresponding to the anterior and pos- terior roots of the spinal nerves ; and it does not seem entirely fanciful to regard their development as somewhat similar, thus : Third, fourth, sixth and seventh, motor; fifth, sensory. Twelfth, motor ; ninth, sensory. Eleventh, motor ; tenth, sensory. Development of the Brain. It has already been shown how from the growth and fusion of the two sides of the primitive medullary groove a tube is formed ; this tube representing the cerebro-spinal axis. Starting with this hollow cylindrical tube, closed at both ends, the spinal cord and brain are formed as a result of changes in the size of the lumen of the tube, alternate thick- enings and thinnings of the walls of the tube, and various foldings and reduplications of the tube on itself. 300 DEVELOPMENT. Earliest rudimentary brain: The first demarcation between brain and spinal cord consists of a widening of the medullary canal at its anterior end. At the same time FIG. 152. this primitive brain is subdivided into three portions by two transverse constrictions. The three chambers are known as the an- terior primary vesicle, middle primary vesicle, and posterior primary vesicle (Fig. 152). Secondary vesicles : As the result of further development the posterior primary vesicle is subdived into two parts by a transverse constriction. The anterior primary vesicle is also further developed by a forward growth of two projections, one on either side of the median line (hemisphere vesicles, Fig. 153). Ventricles of the brain : The various ven- tricles of the brain are the successors of the original lumen of the cerebral axis. Thus we see that the lumen of the prosencephalon is known in the developed brain as the lateral ventricles. The lumen of the thala- mencephalon is the third ventricle, and the communication between the two lateral ventricles and the third ventricle is called the foramen of Monro. The lumen of the mesencephalon (mid-brain) becomes the aqueduct of Sylvius. The cavity of the epencephalon (pons and cere- bellum) and of the metencephalon (medulla) is the fourth ventricle. Development of the walls of the cerebral tube : Commensu- rate with the division of the primary brain into its vesicles, there are changes going on in the walls of the tube. These changes consist of thickenings at some parts and thinnings at others. Thus the prosencephalon shows a more or less uni- form increase in thickness, destined to become the substance of the cerebral hemispheres, corpora striata, etc. The pons Varolii is a great increase ventrally of the wall halon, while the cerebellum is a corresponding the same vesicle. Formation of the ce- rebro-spinal axis. 1, anterior primary vesicle ; 2, middle primary vesicle ; 3, posterior primary vesicle. 3 R A R V DEVELOPMENT OF WALLS OF CEREBRAL TUBE. 301 FIG. 153. Anterior primary vesicle. Middle primary vesicle. Posterior primary vesicle. Formation of secondary vesicle, (diencephalon) 1, 1, prosencephalon ; mesencephalon ; 4, epencephalon ; 5, metencephalon. 2, thalamencephalon I. Anterior primary vesicle. II. Middle primary vesicle. III. Posterior primary vesicle. (Cerebral hemispheres, corpora striata, cor- pus callosum, fornix, lateral ventricles, ol- (^ factory bulb. 2. Thalamenceph- f Thalami optici, third 1. Prosencepha- lon. alon. \ 3. Mesencephalon. < ina 4. Epencephalon. 5. Metencephalon. ventricle,optic nerve. Corpora quadrigem- crura cerebri, aqueduct of Sylvius. Cerebellum, pons Va- rolii, anterior part of fourth ventricle. Medulla oblongata, posterior part of fourth ventricle, au- ditory nerve. 302 DE VELOPMEXT. The preceding table enumerates the changes described above. The roof of the fourth ventricle, in its lower half, is de- scribed as consisting of but a single layer of epithelial cells. This is a result of a thinning of the dorsal wall of the meten- cephalon. Folding of the brain-axis: So far we have described the brain as consisting of a straight tube with various constric- tions and changes in the thickness of its walls. But the de- velopment is not so simple as that. The brain is enclosed in an unyielding box (the skull), which develops and grows at the same time that the brain grows, but not commensurately. The brain tends to grow in the direction of its long axis, but the skull grows as a sphere; hence to accommodate itself to the containing cavity, the brain must fold on itself i. e., coil up. So changes take place, with the result that the cercf>ricle to fold in upon itself. The follicle of epiblast is shut off at the surface, and a ball of its substance left in the cup of the in- DEVELOPMENT OF THE ALIMENTARY CANAL. 303 folded optic vesicle. This ball forms the rudimentary lens, and the anterior layer of the vesicle is the retina. The muscular and vascular structures, as well as the con- nective tissue and humors, are derived from the mesoblast, which in part enfolds the ocular vesicle and in part enters it between the lens and the edge of the cup-like depression. The cornea is of later formation, and is derived from the epiblast of the skin. Development of the auditory apparatus : Very early in the life of an embryo there is a depression on either side of the head which passes through the same process as that men- tioned for the crystalline lens and for the germinal epithelium in the formation of ova. The mass of epiblast thus separated forms the epithelium of the labyrinth and vestibule, the sur- rounding mesoblast furnishing the bony and muscular struct- ures. The auditory nerve is developed with other cranial nerves, and grows in to its end-organs from its central origin. Development of the olfactory apparatus : In a similar way to the internal ear and the lens. The nasal fossa is primarily a depression in the superficial epiblast, which widens and deepens and receives the nerve-filaments from the olfactory lobe. This lobe is originally a bud from the prosencephalon. The primary olfactory depression continues to widen until it opens into the mouth, and is again shut oif by the growth of the " branchial arch/' which forms the superior maxilla. The nose is similarly derived from the mesial and lateral nasal processes. Development of the alimentary canal : As has already been explained, the primitive alimentary canal is formed from the involution of the splanchnopleure, and is really a portion of the yelk-sac partially shut oif from the rest. It is at each end a blind pouch which follows the head and tail folds. The portions have received the names fore-gut and hind-gut as they occupy one or other of these folds. The fore-gut joins with the mouth-cavity by the folding back of the epiblast in the formation of the stomadoeum, and from it are formed the pharynx, oesophagus, and stomach. By a similar involution of the epiblast the anus and lower 304 DEVELOPMENT. part of the rectum are formed, into which the hind-gut opens to complete the alimentary tract. The oesophagus is sometimes impervious at birth, and the rectum or anus may also be imperibrate. This is caused by the non-union of the segments developed from the epiblast with those developed from the hypoblast. Glands of the alimentary tract : (1) The salivary glands are developed from the epiblast lining the oral cavity. They a it- pear primarily as a simple tube which develops branches, about which the alveoli are formed. (2) The pancreas is similarly developed from the hypoblast of the fore-gut. (3) The liver is primarily a protrusion, into a mass of meso- blastic tissue, of the hypoblast of the fore-gut, which appears as soon as the bloodvessels begin to show themselves. The omphalo-mesenteric vein, from the umbilical vesicle, breaks up into a capillary plexus in this same tissue, and the hepatic cells develop about it. Derivation of the lungs : The lungs first appear as a bud at the junction of pharynx and oesophagus which soon forms a separate tube (the trachea). The cells from the hypoblast ex- tend into the surrounding mesoblast, and it is from this structure that all of the tissues of the lungs, except its epithe- lium, are formed. The Wolffian body is first seen as early as the third week as a series of transverse tubes which develop in the cells of the mesoblast, just inside of its division into parietal and visceral layers, on each side of the vertebral column. It is not a permanent organ. Internal to it develop the internal organs of generation, while behind the Wolffian body the rudimen- tary kidney develops. The Wolffian body is a sort of temporary kidney. At first it is a large vascular body, resembling the kidney in structure, and opens by its tubes (outer ends, the inner ends being blind) into the Wolffian duct, which leads to and opens into the cloaca. At about the sixth week of foetal life the kidney begins to grow and the temporary organ to atrophy. As this occurs a duct for the kidney (the ureter) is developed from the Wolffian duct. The use of the organ seems to be that of PARTURITION. 305 a temporary kidney ; but by the end of the third month it has been replaced by the permanent organ, and has almost entirely disappeared. Formation of the internal genitals : The body (germinal epithelium) which appears on the inner side of the Wolman body is the nucleus of the future testicle or ovary, while along the outer side are formed two ducts (Miiller's and the Wolf- fian), which pass down to the cloaca or lower end of the hind-gut. At first it is impossible to determine the sex of the foetus. If the foetus is to become a female, the ducts of Muller join to form the uterus and vagina, while the ununited por- tions remain as the Fallopian tubes. The Wolffian ducts are rudimentary in the female and appear as the ducts of Gart- ner. On the other hand, if the male type is to be formed, the Wolffian ducts become convoluted tubules, and each is at- tached to the testis as the epididymis and vas deferens. Miiller's duct is rudimentary in the male, and is only found as the sinus pocularis and the hydatids of Morgagni. Formation of external genitals : In both sexes in early foetal life the external genitals are alike, consisting of a body re- sembling a penis with a fold of skin at either side. In the female this body becomes proportionally smaller, and appears as the clitoris, the two lateral masses becoming the labia ma- jora. In the male a groove on the under surface unites at its borders to form the urethra, while the scrotum is formed from the folds of skin at the side. This differentiated condition may persist in adult life, and has been mistaken for hermaph- rodism. PARTURITION. By parturition is meant the expulsion of the foetus at a viable age from the mother. The average time for the full development of the foetus is given at two hundred and sev- enty-eight days. During this time (period of gestation) the foetus depends absolutely for its existence upon the mother. After parturition the child aerates its blood, ingests food, and purifies its body of wastes for itself,^ 20-Ph.ys. 306 PARTURITION. Method of expulsion of foetus : The foetus is expelled in part by the contraction of the uterine muscles, and in part by the pressure exerted by the abdominal walls. The uteri IK? contractions are the first to appear, and it is not until the foetus enters the vagina that the abdominal muscles are brought into play. Causes of uterine contractions : As to this no satisfactory answer has been given. Why the uterus should contain the growing embryo for months, and then be suddenly thrown into action to expel it, cannot be explained. Nature of parturition : It is a reflex action depending upon a centre in the lumbar spinal cord. Whence the stimuli are derived which excite the reflex is unknown, but probably from the organ itself. Character of uterine contractions : They are rhythmical in character, and may be compared to the contractions of the heart-muscle. Each "pain" begins feebly, gradually in- tensifies until it reaches a maximum, and then gradually de- clines until it entirely dies away, to be succeeded by another similar contraction and pause. This rhythmical action con- tinues until the uterine contents are expelled, and then the organ enters into a condition of tonic contraction. After parturition, by a process of involution lasting for a few weeks, the uterus returns to its normal unimpregnated state. APPENDIX. TABLE OF THE DEVELOPMENT OF AN EMBRYO. (Modified from Gray's Anatomy.) 1st Week. Ovum in Fallopian tube. Segmentation of vitellus. 2d Week. Ovum in uterine decidua. Chorion. Formation of blastoderm and division of mesoderm. Heart and medullary groove. Amnion and umbilical vesicle formed. Allantois. 3d Week. Head and tail flexures. Closure of medullary canal, and formation of primary cerebral vesicles and ocular and auditory vesi- cles. Branchial arches. Wolffian bodies. Limbs. 4th Week. Limbs increased. Anal opening. Interventricular sep- tum begins. Ant. spinal nerve-roots. Olfactory fossae. Lungs. Pleurae. 5th Week. Allantois vascular. Trace of feet and hands. Miiller's duct and genital gland. 6th Week. Umbilical vesicle disused. Branchial clefts close. Post- spinal nerve-roots. Membranes of the nervous centres. Bladder. Kidneys. Tongue. Larynx. 7th Week. Muscles perceptible. Many centres of ossification appear. 8th Week. Joints appear in extremities; fingers and toes separate. Crystalline lens. Salivary glands. Spleen. Interventricular septum complete. Sympathetic nerves. 9th Week. Distinction between ovary and testicle. Genital furrow. Pericardium. 3d Month. Formation of placenta. External genitals separate from anus. Eyelids, hairs, and nails. Duct of Wolffian body joins testicle. Jf,th Month. Middle- ear bones. Tympanum and labyrinth. Scrotum and prepuce. 5th Month. Germs of teeth. Hair- and sweat-glands. Brunner's glands. Uterus and vagina distinctly separate. 6th Month. Papillae of skin. Sebaceous glands. Peyer's patches. Free border of nails. 7th Month. Cerebral convolutions. Pupillary membrane disappears. 8th Month. Descent of testis. 9th Month. Opening of eyelids. Ossification of cochlea. CHEMICAL TESTS USED COMMONLY IN PHYSIOLOGI- CAL ANALYSIS. FOR PROTEIDS : Nitric Acid coagulates all except peptones. Heat. All are coagulated by boiling, except peptones. 307 308 APPENDIX. XanthojH'Oteic Reaction. A solution boiled with strong nitric acid becomes yellow : the color is deepened by the addition of ammonia. Biuret RatHinn. With a trace of copper sulphate and an excess of potassium or sodium hydrate they give a purple reaction. Millon's Reaction. With a solution of metallic mercury in strong nitric acid (Millon's reagent) they give a white or pinkish rear lion, and the color becomes more pink on boiling. FOR STARCH: Iodine Reaction. Add to a solution of starch a small quantity of tincture of iodine, and a blue reaction results. The color disappears on heating and returns on cooling. Glycoyen. Same test gives reddish reaction, port-wine color, which disappears on heating and returns on cooling. FOR SUGAR (GLUCOSE) : Moore's Test. Boil solution of sugar with an excess of potassium hydrate, brown color-reaction. Trommer's Test. Add to solution a sufficient amount of potassium hydrate to render it quite strongly alkaline. Then add a solution of copper sulphate, drop by drop, until a distinct blue tinge is visible. Heat, and the presence of sugar is shown by the appearance of red, yellow, or orange color-reaction. Fehling's Test Solution. An alkaline copper solution by which a quantitative test may be made. The solution is somewhat unstable, and is for this reason to be tested by boiling before using. The strength of the solution is such that 1 cubic cm. (15 minims) will be exactly decolorized by ^yth of a gramme (.075 grain) of glucose. This test is very delicate, and is quite commonly used for urinary examinations to detect glycosuria. The Fermentation Text. If a small quantity of yeast be added to a sugar solution, the fungus of the yeast (saccharomyces) will cause the sugar to be decomposed into carbonic acid and alcohol. If the process be continued until the sugar is entirely broken up, the amount of car- bonic acid evolved indicates the proportion of sugar present. FOR BILE-SALTS : Pettenkofer's Test. Upon the addition of sulphuric acid to a solution of bile-salts in water there is a precipitation of the salts, which are redissolved by a further addition of the acid. If a drop of a solution of cane-sugar be added, a deep cherry color is developed. FOR BILE-PIGMENTS : Gmeliii's Test. Add a small quantity of nitroso-nitric acid to a solu- tion of the bile-piirments, and a play of colors result, l>eirmnin<: with green and changing to blue, violet, red, and yellow. This is seen best on a white background ; therefore a plate is often used for this test. APPENDIX. 309 METRIC SYSTEM. 1 Inch 12 ' 3 1 1 1 1 1 1 1 1 2 i 3 ,4 15 |6 |7 18 |910 Millimetres. Centimetres. Square i Centi- ; The area of the figure within the heavy lines is that of a square decimetre. A cube one of whose sides is this area is a cubic decimetre or litre. A litre of water at the temperature of 4 C. weighs a kilogramme. A litre is 1.76 pint; a pint is 0.568 of a litre. The smaller figures in dotted lines represent the areas of a square centimetre and of a squre inch. A cubic centimetre of water at 4 C. weighs a gramme. Square Inch. Metre = 39f inches. Centimetre = f inch. Millimetre = ^ inch. Micromillimetre = ^tjnu inch. Gramme = 15^ grains. Centigramme = ^ grain. Milligramme = ^ifo grain. Kilogramme = 2.2 pounds. INDEX. A. Absorption, 111-115 carbohydrates, 113 destination of food, 113 fats, 113 favoring factors, 111 from stomach, 101 intestinal, 102 peptones, 113 salts, 113 sites, 111 sugars, 113 water, 113 Accommodation, 250 Achromatism, 259 Acid, hippuric, 135 hydrochloric, 97, 98 uric, 135 After-images, 259 negative, 260 positive, 260 Air, 82-84 complementary, 83 expired, 83 composition, 33 weight, 83 inspired, 83 composition, 83 reserve, 82 residual, 83 tidal, 82 vitiated, 88 Albumin, serum, 114 Allantois, 289 history, 289 purpose of, 289 Am n ion, 288 fluid, 289 purpose of, 288 Amoeba, 22 functions, 22, 23 Amylopsin, 105 Anabolism, 18 Anaesthesia, 201 Animal heat, 140-142 loss of, 141 regulation of, 141 centres, 142 sources of, 140 chemical action, 140 friction, 141 oxidation of tissues, 140 warm media, 141 Aphasia, 201 Apncea, 87 Apparatus, 23 Aqtieductus vestibuli, 237 Aqueous humor, 247 Area germinativa, 284 vasculosa, 293 Arterial contraction and dilatation (see also Arteries), 66 tension (see also Arteries), 59 Arteries, 54 action, 67 contraction, 66 dilatation, 66 elasticity, 58 nerve-supply, 67 vaso-constricior, 67 vaso-dilator, 67 resiliency, 56 size, 55 structure, 54 tension, 59 conditions modifying, 59 tone, 67 vasa vasorum, 54 vaso-motor nerves, 54 Asphyxia, 87 Assimilation, 140 Astigmatism, 262 Atmosphere, vitiated 'see also Air), 88 Auditory apparatus, 232 canal," 233, 238 external, 233 internal, 238 311 312 INDEX. Auditory nerve, 238 Auricle, 233 Auricles of heart, 51 action, ">1 regurcitation hindered, 52 Axis-cylinder, 159, 160, 161 Axons^ 163 B. Bacterium lactis, 121 Bile, 106 amount, 107 character, 107 composition, 107 pigments, 108 (imelin's test, 108 tests for, 310 salts, 107 Pettenkofer's test, 108 test for, 310 use of, in digestion, 109 Bilirubin, 108 Biliverdin, 108 Bladder, 129 Blastoderm, 284 Blastoderm ic membrane, 283 Blind-spot, 252 Blood, 30-46 arterial, 31 circulation (see also Circulation of blood), 48-70 frog's foot, 59 coagulation (see also Coagulation of blood), 42 coagulum, 44 buffy-coat, 44 corpuscles (see also Corpuscles of blood), 32-40 plates, 32-40 red, 32-38 white, 32-38 foatal, 290, 291 function of, 46 uases, 42 -i IKS appearance, 30 lack of oxygen, 87 liquor sanguinis, 32 maternal. 290, 291 opacity, 31 physical characteristics, 31 -plasma, :;_', 40 clotting, 41, 42 properties, 41 -plates, 40 -pivsee also Circulation of \ 57 Blood, quantity, 32 reaction, 31 -serum, 41 specific gravity, 31 taste, 31 venous, 31 Body, analysis, 24 energy of, 138 expenditures, 138 results of, 138 temperature of, 140 limits, 142 Brain (see also Cerebrum), 192 .'<)! development, 300-303 fibres, 197 association, 197 commissural, 197 medullary, 197 relation of size to intellect, 197 ventricles of, 301 weight, 197 Branchial cleft, 298, 299 Bronchi, 77 c. Canal is cochlearis, 237 reunions, 237 Capillaries, 54 Carbohydrates, 29, 30, 113 Cells, 18-24 blood (see also Blood), 23 connective tissue, 24 decay, 21 development, 21 endothelium, 24 epithelium, 23 varieties, 23, 24 gemmation, 20 karyokinesis, 20 nerve, (see also Nervous system', 23 nucleolus, 19 nucleus, 19 protoplasm, 19-22 difference between plants and animals, 23 specialization, 23 structure, 18 Centimetre, 311 Centres (see ^/limil cord, Mlull9, 281 Fenestra oval is, 233 rotunda, 233 Ferments, pancreatic, 105 Fertilization, methods, 281 Fibrin, 44 ferment, 44 Fibrinogen, 40, 41, 44 Foatal circulation (see also Circulation, festal), 292 Foetus, expulsion, 307 Folds, 285, 298 Food, 137 Food-stuffs, 30 carbohydrates, 29, 30 fats, 29, 30 proteids, 28, 30 Foramen ovale, 295 Fovea centralis, 252 Funiculus cuneatus, 187 gracilis, 187 G. Gall (see also Bile), 106 -bladder, 106 -ducts, 106 Ganglia of sympathetic (see also Sym- pathetic system}, 171, 172 Ganglion, 171 cervical, 172 cceliac, 172 impar, 172 ophthalmic, 171 otic, 171 semi lunar, 172 spheno-palatine, 171 submaxillary, 171 thoracic, 172 (iasrs in blood, 42 carbon dioxide, 42 nitrogen, 42 oxygen, 42 Gastric digestion, 100 conditions favoring, 100 Gastric digestion, nervous mechanism, 100 time required, 100 juice, 97 composition, 97 functions, 98 secretions, 98 Gelatins, 28 Gemmation, 20 Generation, asexual, 268 organs of, 269, 278 female, 269-278 male, 278-280 Geniculate bodies, 255 Germ-hill. 271 Germinal epithelium, 271, 300 membrane, 283 spot, 273 vesicle, 273 Gestation, 306 Glands, ductless. 135 purpose of, 136 of intestine (see also Intestine), 102- 104 lachrymal, 244 mammary (see also Mammary glands), 119 Meibomian, 244 prostate, 280 sebaceous, 122 sweat, 122 vascular, 135 Glandular activity, 117 conditions affecting, 117 amount of blood, 117 of material, 117 nervous system, 117 Glosso-labio-laryngeal paralysis, 191 Glottis, 241 Glycogen. 110, 114 in muscle, 145 Graafian follicle, 270 maturation of, 271 Gramme, 311 Gustatory-buds, 227 Gut, fore', 304 hind, 304 H. Hair, 124 -follicle, 124 Hallucinations, 221 11. a ring, 232-241 c.-ntre, 199 localization, 199, 240 musical range of, ~ 1 1 INDEX. 317 Hearing, subjective, 240 Heart, 51-54, 61 action, 61 apparent elongation, 61 auricles, 51 beat, 61 diastole, 61 systole, 61 cardiac cycle, 62 development of, 293 rate, 61 situation, 51 size, 51 ventricles, 51 weight, 51 Heat, animal (see also Animal 140-142 Hemiansesthesia, 201 Hemiplegia, 201 Hemispheres of cerebellum, 201 of cerebrum, 192 Heredity, 267 Hippuric acid, 135 Hymen, 274 Hypermetropia, 261 Hyperpnoea, 87 Hypoblast, 285 structures derived from, 287 I. Impregnation, 281 details, 282 site of, 282, 283 time of, 282 Incus, 234 Inhibition, 185-189 Insalivation, 92 Inspiration (see also Respiration), 81 Intestine, juice, 104 large, 110 absorption, 110 digestion, 110 structure, 110 peristaltic action, 102-111 small, 101-104 glands, 102-104 agminate, 104 Bru nner's, 103 Lieberkiihn's, 103 Peyer's, 103 solitary, 103 Iris, 248, 259 78- Judgments, 222 J. K. Caryokinesis, 20 Catabolism, 18 Kidneys, 126-135 blood-supply, 128 arteries, 129 afferent, 129 arterise rectse, 129 iuterlobular, 129 veins, 129 efferent, 129 interlobular, 129 venae rectse, 129 cortical portion, 127 medullary portion, 127 size, 126 weight, 127 L. Labyrinth, bony, 235 membranous, 235 Lacteals, 70-113 ^actose, 121 Lamina spiral is, 236 Larynx, 76, 241 lesions in brain and cord, 203, 204 Leukocytes, 38-40 ; 46, 59 characteristics, 38 chemical composition, 39 death, 40 function, 40 emigration, 38, 60 phagocytosis, 40 number, 32 occurrence, 38 origin, 39 varieties, 39 lymphocytes, 39 mononuclear, 39 polynuclear, 39 Life, 17 living body compared to machine, 17 Litre, 311 Liver, 106-110 functions, 106 elaboration of urea, 106, 110 excretion, 106, 109 glycogenic, 106, 109 secretion, 106 Lochia, 292 Lungs, 77 capacity of, 82 minute anatomy, 77 Lymph, 46-48 318 INDEX. Lymph coagulation, 47 chyle, 47 circulation, 70 composition, 46 flow, 70 factors in, 73 ganglia, 73 pressure, 72 Lymph spaces, 46, 70, 113 sources, 47, 72 intercellular spaces, 47 lacteals, 47 uses, 47 Lymphatic nodes, 73 purpose, 74 Lymphatics, 70 structure, 72 Lymphocytes, 39 M. Macula acustica, 236 lutea, 252 Malleus, 234 Malpighian bodies, 127 Mammary glands, 119 secretion, 119 structure, 119 alveoli, 119 lobes, 119 lobules, 119 Mastication, 90 muscles, 92 Mastoid cells, 233 Medulla oblongata, 185-191 automatic action, 189 centres, 189 conduction, 188 function, 188-190 gray matter, 187 gross anatomy, 185 inhibition, 189 reflex action, 188 Medullary canal, 299 groove, 285 sheath, 159, 160 Membrana basilaris, 237 tympani, 234 Membrane, blastodermic, 283 germinal, 283 of Reissner, 237 vitelline, 272 Menopause, 275 Menstruation, 274 connection with ovulation, 278 corpus luteum of, 278 Menstruation, definition, 274 discharge, 276 duration, 275 frequency of, 275 nature, 275 Mesoblast, 285 changes in, 286 structures derived from, 286 Metabolism, 18 Metric system, 311 Microcytes, 40 Micturition, 133 Milk, 119 composition, 119 secretion, 120 Mouth, 90 Mucin, 118 Mucus, 118 Muller's duct, 306 Muscle, 142-159 chemistry of, 144 conductivity, 155 contractility, 148, 149 contraction, 148 artificial stimuli, 150 chemical, 150 electrical, 150, 151, 152, 153 mechanical, 150 response to, 151 thermal, 150 path of stimulus, 148 Pfliiger's law. !."> results of, 157 currents, ascending, 156 descending, 156 of rest, 1 "> 1 electrical state, 154 excitability, 155 fatigue, 147 general properties, 144 latent period, 147, 152 apparent, 147 true, 147 as levers, 146 nerve-muscle preparation. 157 conditions influencing results, 157 order of fatigue, 157 oxygen-supply, 1 1<> physiology of, 145 activity, 145 rest, 145 rigor, 145 plasma, 144 varieties, 142 non-striated, 142 INDEX. 319 Muscle, varieties, striated, 142 involuntary, 142, 144 voluntary, 142, 143 Mutilations, 204, 205 brain, 204 cerebellum, 204 cord, 204 Myopia, 260 Myosin, 144 Myosinogen, 144 N. Nails, 124 matrix, 124 root, 124 Nasal cavity, 231 Near-point, 260 Nerve-cells, 159, 162, 175 ganglia, 159, 162 salivary secretion, 93 -degeneration, 167, 177 histologies! changes, 168 -endings, 163 efferent nerves, 163 sensory nerves, 163 -fibres, 159 centrifugal, 162 centripetal, 162 function, 161 of hemispheres, 197 rnedullated, 159 motor, 179, 203 non-medullated, 159, 161 of Eemak, 161 sensory, 179, 204 of sympathetic, 169 -impulse, 165 direction of, 166 experiments, 166 nature of, 165 olfactory, 230 origin, 230 speed of, 167 -regeneration, 168 -roots, 174, 176 anterior, 174, 176 course, 176 degeneration, 167, 177 posterior, 174, 176 course, 176 trophic centres, 177 -trunks, 161 vaso-motor, 66, 173 Nerves, auditory, 238 centres, 199 Nerves, cranial (see also Cranial nerves), 206 nuclei of, 188 nucleus, 238 optic, 253 fibres of, 251, 253 pneumogastric, in respiration, 86,217 section of, 87 spinal, 174 roots, 174 Nervous system, 159-220 Neurilemma, 159, 160 Neuroglia, 175 Neuron, 164 Nitrogen in atmosphere, 84 Nitrogenous equilibrium, Io9 Nodes of Eanvier. 160 Nose, 231 Nutrition, 136-138 o. Odors, 229 Olfactory bulb, 231 cells, 230 nerves, 230 tract, 231 Olivary bodies, 187 Optagrams, 258 Optic chiasm, 253 nerve (see also Nerves, optic), 253 radiations, 253 thalamus, 196, 198 tracts, 253 Organ of Corti, 238 Organs, 23 Osmosis, 111 Ossicles, 234 Ovaries, 270 Graafian follicles, 270 strom a, 270 Ovulation, 276 connection with menstruation, 278 frequency of, 276 Ovum, 271, 282 constriction, 285 unimpregnated, 276 history of, 276 Oxidation, 18 Oxygen, 137 P. Pacinian corpuscle, 164 Pain, 181, 221 Pancreas, 104 320 INDEX. Pancreas, juice, 104 composition, 105 function, 106 Paraglobulin, 41, 44 Paraplegia, 201 Parturition. :soi> nature, 307 time, 306 Penis, 280 erectile tissue, 281 glans, 280 Pepsin, 97, 98, 105 Pepsinogen, 98 Peptones, 91). li:; Perceptions, 222 visual, 257 mental processes of, 257 Perilymph, 235, 239 Perspiration, 122, 124 amount. 1 -.'."> insensible, 125 necessity, 126 nervous mechanism, 125 odor, 125 purpose, 125 Placenta, 290 appearance of, 292 destiny of, 291 foetal, 290 layers of cells, 290 maternal, 290 period of formation, 292 Place ntal circulation (sec also Circula- tion, plnceutal), 292, 294 Plasma, 40 of muscle, 144 Pleura, 77 Plexus, pelvic, 172 solar, 172 Polar body, 277 Pons Varolii, 191 Presbyopia, 261 Primitive groove, 285 trace, 2S4 Processes, intermaxillary, 299 maxillary, _'!>!> Pronucleus, female, 277 malr, 282 Prostate gland, 280 Proteids, 28, 30 albumins, 28 fibrin, 28 globulins, 28 peptones, 28 t<-sts for, 309 Proteolytic action, 99 Protoplasm, 19, 22 Protovertebne, 298 Proximate principles, 25-29 inorganic, 25, 26 distribution, 26 uses, 26 organic, 25 list of, 28, 29 nitrogenous, 27, 28 non-nitrogenous, 27, 28 source, 27 uses, 27 Puberty, 274 Pulse, 68-70 extinction, 69 sphygmograph, 68 tracings, (>9 varieties, 68 venous, 54, 69 Pyramidal tracts, 177, 186, 203 crossed, 1S6, 203 direct, 186, 203 Pyramids of medulla, 186 B. Reflex action of cord, 181, 182 of medulla, 188 Refraction (eye), 248, 249 Rennin, 121 Reproduction, 267-283 methods of, 267 asexual, 2(>S sexual, 268, 269 theory of, 268 Respiration. 75-90 abdominal, 80 act of, 75 associated movements, 82 automatic impulses, 86 centres, 86 synchronism of, 87 effects of, on circulation, 88 expiration, 81 muscles, 81 external, 84 force of, 83 forced, 81 frequency, 82 inspiration, 78-81 muscles, 78 internal. >.", movements. 75 nerves, 86 nervous mechanism, 85 reflex influences, 85 INDEX. 321 Respiration sounds, 82 special acts, 89 sighing, 89 sneezing, 89 laughing, etc., 90 thoracic, 80 tract, 75 Kestiform body, 187 Retina, 246, 250, 259 inversion of image, 25(5 pigment of, 259 Retinal red, 257 reflex, 258 Rigor mortis, 158 causes of, 159 disappearance of, 159 Rima glottidis, 241 Rods, 251 8. Saccule, 237 Saliva, 92 mixed, 92 secretion, 93 chorda tympani, 93 nerve-control, 93 Saponification, 105 Scala media, 237 tympani, 236 vestibuli, 236 Sclera, 246 Sebaceous glands, 122 secretion, 125 Secreting glands, 116 racemose, 116 tubular, 116 Secretion, 115-136 correlation, 118 discharge of, 117 milk, 120 mucous, 118 mucus, 118 processes of, 115, 116 chemical, 117 physical, 116 serous, 118 peritoneum, 118 pleura. 118 table of, 136 Segmentation, 283 nucleus, 283 Semicircular canals, 235 Seminal cells, 279 vesicles, 280 Seminiferous tubules, 279 21 Phys. Sensations, centres, 199 classification, 221 common, 221 organs necessary for, 220 pressure, 224 special, 221 visual, 256 duration of, 256 Senses, the, 220-265 nerves of, 222 seat of, 221 special, 222 muscle, 224 centre, 225 temperature, 225 Sense-organs, 163 Pacinian corpuscles, 164 touch-corpuscles, 164 Sensibility, tactile, 224 variations, 224 Serum, 41, 118 -albumin, 41 Sight, 243-265 Skin, 121-124 absorption by, 126 corium, 121, 122 cuticle, 121 cutis vera, 121, 122 derma, 121, 122 epidermis, 121 papillae, 122 Smell, 229-232 acuteness, 232 conditions for, 229 Sneezing, 232 Sodium glycocholate, 107 taurocholate, 107 Somatopleure, 286, 288 Somites, mesoblastic, 298 Sound-waves, 239 course of, 239 Species, 267 Speech, 241 articulate, 242 organs used in, 242 centre, 199 Spermatozoa, 279 Spermatozoon, 281, 282 Sphygmograph, 68 Spinal cord, 174-185 augmentation, 184 automatic acts, 184 columns, 175, 186 anterior, 186 lateral, 187 posterior, 187 322 INDEX. Spinal cord, coordination, 184 enlargements, 177 functions, 178 conduction, 178 gray matter, 175 gross anatomy, 174 lesions, 203 microscopic examination, 175 mutilations, 204 recurrent-sensibility fibres, 177 reflex action, 181, 182 special reflexes, 182 transference, 181 white substance, 175 Splanchnopleure, 286, 287, 288 Spleen, 136 Stapes, 234 Starch, tests for, 310 Starvation, 138 Steapsin, 105 Stercobilin, 109 Stercorin, 109 Stereoscope, 256 Stomach, 94-99 absorption from, 101 capacity, 100 digestion, gastric (see also Gastric digestion), 100 after death, 101 glands, 96 peptic, 96 pyloric, 96, 97 muscular action, 99 structure, 94 Stomata, 71 Strabismus, 262 external, 262 internal, 262 Succus entericus, 104 Sugar, test for, 310 Sweat, 122, 124 -glands, 122 Sympathetic system, 169-173 function, 173 ganglia, 169, 171-172 abdominal, 172 cephalic, 171 cervical, 172 thoracic, 172 relation to secreting glands, 173 sensory and motor influence, 173 vaso-motor fibres, 173 Systole (see also Heart), 61 Taste, ir>-J T. Taste, after-, 229 a.-v-MH-iatioi! with smell, 228 -goblets, 227 necessary conditions for, 225 origin of, 226 Teeth, 91 permanent, 91 temporary, 91 Temperature of body, 140 srii>ations, 181 Testicles, 279 tubules, 279 tunica albuginea, 279 Thoracic duct, 70, 114 Thymus, 136 Thyroid, 136 Tic douloureux, 211 Tissue, 23 Tongue, 90, 226 papillae, 226 sensibility, 229 taste-goblets, 227 touch, 227 varieties, 226 Tonsils, 136 Tooth, 91 cavity, 92 cement, 92 crown, 91 crusta petrosa, 92 dentine, 92 enamel, 92 fang, 91 neck, 91 pulp, 92 Touch, 181, 222-225 acuteness of, 223 measure of, 223 corpuscles, 164 organ of, 222 varieties of, 223 Trachea, 76 Tracts, 203, 204 motor, 203 lesions, 203, 204 sensory, 204 Trypsin, 105 Tympanum, 233 membrane, 233, 234 u. Umbilical arteries, 291, 296, 297 cord, 291 veins, 291 vesicle, 287 INDEX. 323 Urea, 134 amount, 135 as a waste, 135 Ureters, 129 pelvic, 129 Urethra, 280 Uric acid, 135 Urine, 129 acidity, 131 acid sodium phosphate, 131 amount, 131 coloring-matter, 130 composition, 130 course in tubules, 133 estimation of solids, 135 properties, 129 secretion, 130 conditions affecting, 132 epithelial action, 130 nitration, 130 relation to arterial pressure, 132 Uriniferous tubules, 127 course, 127 Uterine sinuses, 290 Uterus, 273 contractions of, 307 causes, 307 characters of, 307 Utricle, 237 V. Valves of heart, 53 of veins, 55 Valvulse conniventes, 102, 111 Vas defereus, 279 Veins, 55 contraction, 57 pulsation of jngular, 54 size, 56 structure, 55 Veins, valves, 55 Ventricle, fourth, floor of, 188 of heart, 51, 52 action, 52 lateral, 193 Vesicles, primary, 301, 302 secondary, 301, 302 Vestibule, 235, 237 Villi, 111, 112 structure, 112 Vision, area of most acute, 252 binocular, 255 centre for, 199 clearness, 257 nervous mechanism of, 255 Vitelline duct, 287 membrane, 272 Vitellus, 273, 287 Vitreous humor, 248 Vocal cords, 241 Voice, 241 musical range of, 242 production, 241 Vomiting, 101 w. Waste-products, 18 Whartou's jelly, 291 Wolffian body, 305 duct, 305 Y. Yolk-sac, 287, 292 Z. Zona pellucida, 272 Zymogen, 104 CATALOGUE OF PUBLICATIONS OF LEA BROTHERS & COMPANY, 706, 708 & 710 Saiisom St., Philadelphia. Ill Fifth Ave. (Cor. 18th St.), New York. 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Price per volume, cloth, $5 ; leather $6 ; half Russia, $7. Sold by subscrip- tion only. Prospectus free on application to the Publishers. ATTFDELD (JOHN). CHEMISTRY ; GENERAL, MEDICAL AND PHARMACEUTICAL. New (16th) edition, specially revised by the Author for America. In one handsome 12mo. volume of 784 pages, with 88 illustrations. Cloth, $2.50, net. It is replete with the latest inform- been adopted, bringing the work into ation, and considers the chemistry of close touch with the latest United every substance recognized officially States Pharmacopoeia, of which it is or in general practice. The modern a worthy companion. ThePittsburg scientific chemical nomenclature has Medical Review. BALLENGER (W. L.) AND WIPPERN (A. G.). Shortly. A POCKET TEXT-BOOK OF DISEASES OF THE EYE, EAR, NOSE AND THROAT. In one handsome 12tno. volume of about 400 pages, with many illustrations. Cloth, $1.50, net. Lea's Series of Pocket Text-books, edited by BERN B. GALLAITDET, M. D. See p. 18. BARNES (ROBERT AND FANCOURT). A SYSTEM OF OB- STETRIC MEDICINE AND SURGERY. Octavo, 872 pages, with 231 illus. Cloth, $5 ; leather, $6. 4 LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. BACOX (GORHAM). ON THE EAR. One 12mo. volume, 400 pages, 109 engravings and a colored plate. Cloth, net, $2.00. Just ready. It is the best manual upon otology, dents of medicine Cleveland Junr- An intensely practical book for stu- nal of Medicine. BARTHOLOW (ROBERTS). CHOLERA; ITS CAUSATION, PRE- VENTION AND TREATMENT. In one 12mo. volume of 127 pages, with 9 illustrations. Cloth, $1.25. BARTHOLOW (ROBERTS). MEDICAL ELECTRICITY. A PRACTICAL TREATISE ON THE APPLICATIONS OF ELEC- TRICITY TO MEDICINE AND SURGERY. Third edition. In one octavo volume of 308 pages, with 110 illustrations. BELL (F. JEFFREY). COMPARATIVE ANATOMY AND PHYS- IOLOGY. In one 12mo. volume of 561 pages, with 229 engravings. Cloth, $2. See Students' Series of Manuals, page 27. BILLINGS (JOHN S.). THE NATIONAL MEDICAL DICTIONARY. Including in one alphabet English, French, German, Italian and Latin Technical Terms used in Medicine and the Collateral Sciences. In two very handsome imperial octavo volumes containing 1574 pages and two colored plates. Per volume, cloth, $6 ; leather, $7 ; half Morocco, $8.50. For sale by subscription only. Specimen pages on application to the publishers. BLACK (D. CAMPBELL). THE URINE IN HEALTH AND DISEASE, AND URINARY ANALYSIS, PHYSIOLOGICALLY AND PATHOLOGICALLY CONSIDERED. In one 12mo. volume of 256 pages, with 73 engravings. Cloth, $2.75. Concise, practical, clinical, well illustrated and well printed. Mary- land Medical Journal. A concise, yet complete manual, treating of the subject from a prac- tical and clinical standpoint. The Ohio Medical Journal. BLOXAM (C. L.). CHEMISTRY, INORGANIC AND ORGANIC. With Experiments. New American from the fifth London edition. In one handsome octavo volume of 727 pages, with 292 illustrations. Cloth, $2 ; leather, $3. BROCKWAY (F. J.). A POCKET TEXT-BOOK OF ANATOMY. In one handsome 12mo. volume of about 400 pages, with many illus- trations. Shortly. Cloth, $1.50, net. Lea's Series of Pocket Test-books, edited by BERN B. GALLAUDET, M. D. See page 18. BRUCE (J. MITCHELL). MATERIA MEDICA AND THERA- PEUTICS. New (6th) edition. In one 12mo. volume of 600 pages. Just ready. Cloth, $1.50, net. See Student's Series of Manuals, page, 27. PRINCIPLES OF TREATMENT. In one octavo volume. Pre- paring. BRYANT (THOMAS). THE PRACTICE OF SURGERY. Fourth American from the fourth English edition. In one imperial octavo vol. of 1040 pages, with 727 illustrations. Cloth, $6.50 ; leather, $7.50. BURCHARD (HENRY H.). DENTAL PATHOLOGY AND THER- APEUTICS. Handsome octavo, 575 pages, with 400 illustrations. Just ready. Cloth, net, $5.00 ; leather, net, $6.00. In the treatment of the subject I is a valuable text-book on a subject the method pursued by the author which has heretofore not been ade- is logical ana sequential. The work I quately represented. -Dental Cosmos. LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. 5 BURNETT (CHARLES H.)< THE EAR : ITS ANATOMY, PHYSI- OLOGY AND DISEASES. A Practical Treatise for the Use of Students and Practitioners. Second edition. In one 8vo. volume of 580 pages, with 107 illustrations. Cloth, $4 ; leather, $5. CARTER (R. BRUDENELL) AND FROST (W. ADAMS). OPH- THALMIC SURGERY. In one pocket-size 12mo. volume of 559 pages, with 91 engravings and one plate. Cloth, $2.25. See Series of Clinical Manuals, page 25. CASPARI (CHARLES JR.). A TREATISE ON PHARMACY. For Students and Pharmacists. In one handsome octavo volume of 680 pages, with 288 illustrations. Cloth, $4.50. The author's duties as Professor student who cannot understand must of Theory and Practice of Pharmacy be dull indeed. The book is full of in the Maryland College of Phar- new, clean, sharp illustrations, which macy, and his contact with students tell the story frequently at a glance, made him aware of their exact ! The index is full and accurate. wants in the matter of a manual, j National Druggist. His work is admirable, and the i CHAPMAN (HENRY C.). A TREATISE ON HUMAN PHYSI- OLOGY. New (2d) edition. In one octavo volume of 921 pages, with 595 illustrations. Just ready. Cloth, $4.25 ; leather, $5.25, net. In every respect the work fulfils its promise, whether as a complete treatise for the student or as an ad- mirable work of reference for the physician. North Carolina Medical Journal. CHARLES (T. CRANSTOUN). THE ELEMENTS OF PHYSIO- LOGICAL AND PATHOLOGICAL CHEMISTRY. Octavo, 451 pages, with 38 engravings and 1 colored plate. Cloth, $3.50. CHEYNE (W. WATSON). THE TREATMENT OF WOUNDS, ULCERS AND ABSCESSES. In one 12ino. volume of 207 pages. Cloth, $1.25. One will be surprised at the amount of practical and useful in- formation it contains; information that the practitioner is likely to need at any moment. The sections devoted to ulcers and abscesses are indispensable to any physician. The Charlotte Medical Journal. CHEYNE (W. W.) AND BURGH ARD (F. F.) SURGICAL TREATMENT. In six octavo volumes, illustrated. Volume 1, 299 pages and 66 engravings, just ready. Cloth, $3.00 net. CLARKE (W. B.) AND LOCKWOOD (C. B.). THE DISSECTOR'S MANUAL. In one 12mo. volume of 396 pages, with 49 engravings. Cloth, $1.50. See Students' Series of Manuals, page 27. CLELAND (JOHN). A DIRECTORY FOR THE DISSECTION OF THE HUMAN BODY. In one 12mo. vol. of 178 pages. Cloth, $1.25. CLINICAL MANUALS. See Series of Clinical Manuals, page 25. CLOUSTON (THOMAS S.). CLINICAL LECTURES ON MENTAL DISEASES. New (5th) edition. In one octavo volume of 750 pages, with 19 colored plates. Cloth, $4.25, net. Just ready. ^^^FOLSOM's Abstract of Laws of U. S. on Custody of Insane, octavo, $1.50, is sold in conjunction with Clouston on Mental Diseases for $5.00, net, for the two works. 6 LRA BEOTHEBS & Co., PHILADELPHIA AND NEW* YORK. CLOWES (FRANK). AN ELEMENTARY TREATISE ON PRACTI- CAL CHEMISTRY AND QUALITATIVE INORGANIC ANALY- SIS. From the fourth English edition. In one handsome 12mo. volume of 387 pages, with 55 engravings. Cloth, $2.50. COAKLEY (COBNEJLIUS G.). THE DIAGNOSIS AND TREAT- MENT OF DISEASES OF THE NOSE, THROAT, NASO- PHARYNX AND TRACHEA. In one 12mo. volume of about 400 pages, fully illustrated. Preparing, COATES (W. E., JR.). A POCKET TEXT-BOOK OF BACTE- RIOLOGY AND HYGIENE. In one handsome 12mo. volume of about 350 pages, with many illustrations. Shortly. Cloth, $1.50, net. Lea's Series of Pocket Test-books, edited by BEKN B. GALT.AVDET, M. D. See page 18. COATS (JOSEPH). A TREATISE ON PATHOLOGY. In one vol. of 829 pages, with 339 engravings. Cloth, $5.50; leather, $6.50. COLEMAN (ALFRED). A MANUAL OF DENTAL SURGERY AND PATHOLOGY. With Notes and Additions to adapt it to Amer- ican Practice. By THOS. C. STELLWAGEN, M.A., M.D., D.D.S. In one handsome octavo vol. of 412 pages, with 331 engravings. Cloth, $3.25. COLLINS (C. P.). A POCKET TEXT-BOOK OF MEDICAL DIAGNOSIS. In one handsome 12mo. volume of about 350 PJI-.S, with many illustrations. Shortly. Cloth,$1.50, net. Lea? s Series of P< >f P,,,-l-,t Text-books, edited by BERN B. GALLATJDET, M. D. See page 18. HABERSHON (S. O.). ON THE DISEASES OF THE ABDOMEN Second American from the third English edition. In one octavo vol- ume of 554 pages, with 11 engravings. Cloth, $3.50. HALL (WINFIELD S.) TEXT-BOOK OF PHYSIOLOGY. Octavo about 500 pages, richly illustrated. In press. HAMILTON (ALLAN MCLANE). NERVOUS DISEASES, THEIR DESCRIPTION AND TREATMENT. Second and revised edition. In one octavo volume of 598 pages, with 72 engravings. Cloth, $4. HARDAWAY (W. A.). MANUAL OF SKIN DISEASES. New (2d) edition. In one 12mo. volume of 560 pages, with 40 illustrations and 2 plates. Cloth, $2.25, net. Just ready. The best of all the small books to recommend to students and practi- tioners. Probably no one of our dermatologists has had a wider every- day clinical experience. His great strength is in diagnosis, descriptions of lesions and especially in treat- ment. Indiana Medical Journal. HARE (HOB ART AMORY). PRACTICAL DIAGNOSIS. THE USE OF SYMPTOMS IN THE DIAGNOSIS OF DISEASE. New (4th) edition. In one octavo volume of 623 pages, with 205 engravings and 14 full-page colored plates. Cloth, $5.00, net. Jtmt r, <>,/>/. It is unique in many respects, and he will become a better diaguosti- the author has introduced radical cian. This is a companion to /Vac- changes which will be welcomed by | tical Therapeutics, by the same alJ. Anyone who reads this book i author, and it is difficult to conceive will become a more acute observer, of any two works of greater practical will pay more attention to the simple utility. Medical Review. yet indicative signs of disease, and LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. 13 HARE (HOBART AMORY). A TEXT-BOOK OF PRACTICAL THERAPEUTICS, with Special Reference to the Application of Reme- dial Measures to Disease and their Employment upon a Rational Basis. With articles on various subjects by well-known specialists. New (7th) and revised edition. In one octavo volume of 776 pages. Cloth, $3.75, net; leather, $4.50, net. Its classifications are inimitable, and the readiness with which any- thing can be found is the most won- derful achievement of the art of in- dexing. This edition takes in all the latest discovered remedies. The St. Louis Clinique. The great value of the work lies in the fact that precise indications for administration are given. A complete index of diseases and remedies makes it an easy reference work. It has been arranged so that it can be readily used in connection with Hare's Practical Diagnosis. For the needs of the student and general practitioner it has no equal. Medical Sentinel. The best planned therapeutic work of the century. American Prac- titioner and News. It is a book precisely adapted to the needs of the busy practitioner, who can rely upon finding exactly what he needs. The National Med- ical Review. HARE (HOBART AMORY) ON THE MEDICAL COMPLICA TIONS AND SEQUELAE OF TYPHOID FEVER. Octavo, 276 pages, 21 engravings and two full-page plates. Just ready. Cloth, $2.40, net. A very valuable production. One of the very best products of Dr. Hare and one that every man can read with great profit. Cleveland Journal of Medicine. HARE (HOBART AMORY, EDITOR). A SYSTEM OF PRAC- TICAL THERAPEUTICS. In a series of contributions by eminent practitioners. In four large octavo volumes comprising about 4500 pages, with about 550 engravings. Vol. IV., just ready. For sale by sub- scription only. Full prospectus free on application to the Publishers. Regular price, Vol. IV., cloth, $6 ; leather, $7 ; half Russia, $8. Price Vol. IV. to former or new subscribers to complete work, cloth, $5 ; leather, $6 ; half Russia, $7. Complete work, cloth, $20 ; leather, $24 ; half Russia, $28. The great value of Hare's System of Practical Therapeutics has led to a widespread demand for a new volume to represent advances in treatment made since the publication of the first three. More than fulfilling this request the Editor has secured contributions from practically a new corps of equally eminent authors, so that entirely fresh and original matter is ensured. The plan of the work, which proved so successful, has been fol- lowed in this new volume, which will be found to present the latest devel- opments and applications of this most practical branch of the medical art. The entire System is an unrivalled encyclopaedia on the practical parts of medicine, and merits the great success it has won for that reason. 14 LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. HARTSHORNE (HENRY). ESSENTIALS OF THE PRINCIPLES AND PRACTICE OF MEDICINE. Fifth edition. In one 12mo. volume, 669 pages, with 144 engravings. Cloth, $2.75 . A HANDBOOK OF ANATOMY AND PHYSIOLOGY. In one 12mo. volume of 310 pages, with 220 engravings. Cloth, $1.75. A CONSPECTUS OF THE MEDICAL SCIENCES. Comprising Manuals of Anatomy, Physiology, Chemistry, Materia Medica, Prac- tice of Medicine, Surgery and Obstetrics. Second edition. In one royal 12mo. vol. of 1028 pages, with 477 iUus. Cloth, $4.25 ; leather, $5. HAYDEN (JAMES B.). A MANUAL OF VENEREAL DISEASES. New (2d) edition. In one 12mo. volume of 304 pages, with 54 en- gravings. Cloth, $1.50, net. Just ready. It is practical, concise, definite and of sufficient fulness to be satis- factory. Chicago Clinical Review. This work gives all of the prac- tically essential information about the three venereal diseases, gon- orrhoea, the chancroid and syphilis. In diagnosis and treatment it is par- ticularly thorough, and may be relied upon as a guide in the man- agement of this class of diseases. Northwestern Lancet. It is well written, up to date, and will be found very useful. Inter- national Medical Magazine. HAYEM (GEORGES) AND HARE (H. A.). PHYSICAL AND NATURAL THERAPEUTICS. The Remedial Use of Heat, Elec- tricity, Modifications of Atmospheric Pressure, Climates and Mineral Waters. Edited by Prof. H. A. HARE, M. D. In one octavo volume of 414 pages,with 113 engravings. Cloth, $3. This well-timed up-to-date volume is particularly adapted to the re- quirements of the general practi- tioner. The section on mineral waters is most scientific and prac- tical. Some 200 pages are given up to electricity and evidently embody the latest scientific information on the subject. Altogether this work is the clearest and most practical aid to the study of nature's therapeutics that has yet come under our obser- vation. The Medical Fortnightly. For many diseases the most potent remedies lie outside of the materia medica, a fact yearly receiving wider recognition. Within this large range of applicability, physical agencies when compared with drugs are more direct and simple in their results. Medical literature has long been rich in treatises upon medical agents, but an authoritative work upon the other great branch of therapeutics has until now been a desideratum. The section on climate, rewritten by Prof. Hare, will, for the first time, place the abundant resources of our country at the in- telligent command of American practitioners. The Kansas City I Medical Index. HERMAN (G. ERNEST). FIRST LINES IN MIDWIFERY. In one 12mo. vol. of 198 pages, with 80 engravings. Cloth, $1.25. See Student's Series of Manuals, page 27. HERMANN (Li.). EXPERIMENTAL PHARMACOLOGY. A Hand- book of the Methods for Determining the Physiological Actions of Drugs. Translated by ROBERT MEADE SMITH, M. D. In one 12mo, volume of 199 pages, with 32 engravings. Cloth, $1.50. LEA BROTHEBS & Co., PHILADELPHIA AND NEW YORK. 15 HERRICK (JAMES B.). A HANDBOOK OF DIAGNOSIS. In one handsome 12mo. volume of 429 pages, with 80 engravings and 2 colored plates. Cloth, $2.50. Excellently arranged, practical, concise, up-to-date, and eminently well fitted for the use of the prac- titioner as well as of the student. Chicago Med. Recorder. This volume accomplishes its ob- jects more thoroughly and com- pletely than any similar work yet published. Each section devoted to diseases of special systems is pre- ceded with an exposition of the methods of physical, chemical and microscopical examination to be em- ployed in each class. The technique of blood examination,including color analysis, is very clearly stated. Uranalysis receives adequate space and care. New York Med. Journal. We commend the book not only to the undergraduate, but also to the physician who desires a ready means of refreshing his knowledge of diag- nosis in the exigencies of professional life. Memphis Medioal Mo HILL (BERKELEY). SYPHILIS AND LOCAL CONTAGIOUS DISORDERS. In one 8vo. volume of 479 pages. Cloth, $3.25. HILLDER (THOMAS). A HANDBOOK OF SKIN DISEASES. Second edition. In one royal 12mo. volume of 353 pages, Avith two plates. Cloth, $2.25. HIRST (BARTON C.) AND PIERSOL (GEORGE A.). HUMAN MONSTROSITIES. Magnificent folio, containing 220 pages of text and illustrated with 123 engravings and 39 large photographic plates from nature. In four parts, price each, $5. Limited edition. For sale by subscription only. HOBLYN (RICHARD D.). A DICTIONARY OF THE TERMS USED IN MEDICINE AND THE COLLATERAL SCIENCES. In one 12mo. volume of 520 double-columned pages. Cloth, $1.50 ; leather, $2. HODGE (HUGH L.). ON DISEASES PECULIAR TO WOMEN, INCLUDING DISPLACEMENTS OF THE UTERUS. Second and revised edition. In one 8vo. vol. of 519 pp., with illus. Cloth, $4.50. HOFFMANN (FREDERICK) AND POWER (FREDERICK B.). A MANUAL OF CHEMICAL ANALYSIS, as Applied to the Examination of Medicinal Chemicals and their Preparations. Third edition, entirely rewritten and much enlarged. In one handsome octavo volume of 621 pages, with 179 engravings. Cloth, $4.25. HOLMES (TIMOTHY). A TREATISE ON SURGERY. Its Prin- ciples and Practice. A new American from the fifth English edition. Edited by T. PICKERING PICK, F.R.C.S. In one handsome octavo vol- ume of lOOS.pages, with 428 engravings. Cloth, $6 ; leather, $7. A SYSTEM OF SURGERY. With notes and additions by various American authors. Edited by JOHN H. PACKARD, M. D. In three very handsome 8vo. volumes containing 3137 double-columned pages, with 979 engravings and 13 lithographic plates. Per volume, cloth, $6 ; leather, $7 ; half Russia, $7.50. For tale by tubscription only. 16 LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. HORNER (WILLIAM E.). SPECIAL ANATOMY AND HIS- TOLOGY. Eighth edition, revised and modified. In two large 8vo. volumes of 1007 pages, containing 320 engravings. Cloth, $6. HUDSON (A.). LECTURES ON THE STUDY OF FEVER. In one octavo volume of 308 pages. Cloth, $2.50. HUTCHISON (ROBERT) AND RAINY (HARRY). CLINICAL METHODS. A GUIDE TO THE PRACTICAL STUDY OF MEDICINE. In one 12mo. volume of 562 pages, with 137 engrav- ings and 8 colored plates. Cloth, $3.00. A comprehensive, clear and re- markably up-to-date guide to clinical diagnosis. The illustrations are plentiful and excellent. As exam- ples of the more recent additions to medical knowledge which receive recognition, we mention Widal's test for typhoid and the Neuron theory of the nervous system. Montreal Medical Journal. HUTCHINSON (JONATHAN). SYPHILIS. In one pocket-size 12mo. volume of 542 pages, with 8 chromo-lithographic plates. Cloth, $2.25. See Series of Clinical Manuals, p. 25. HYDE (JAMES NEVINS). A PRACTICAL TREATISE ON DIS- EASES OF THE SKIN. New (4th) edition, thoroughly revised. In one octavo volume of 815 pages, with 110 engravings and 12 full- page plates, 4 of which are colored. Cloth, $5.25 ; leather, $6.25. This edition has been carefully re- vised, and every real advance has been recognized. The work answers the needs of the general practitioner, the specialist, and the student. The Ohio Med. Jour. A treatise of exceptional merit characterized by conscientious care and scientific accuracy. Buffalo Med. Journal. A complete exposition of our knowledge of cutaneous medicine as it exists to-day. The teaching in- culcated throughout is sound as well as practical. The American Jour- nal of the Medical Sciences. It is the best one-volume work that we know. The student who gets this book will find it a useful investment, as it will well serve him when he goes into practice. Vir- ginia Medical Semi-Monthly. A full and thoroughly modern text-book on dermatology. The Pittsburg Medical Review. It is the most practical hand- book on dermatology with which we are acquainted. The Chicago Med- ical Recorder. JACKSON (GEORGE THOMAS). THE READY-REFERENCE HANDBOOK OF DISEASES OF THE SKIN. New (3d) edition. In one 12mo. volume of 637 pages, with 75 illustrations and a colored plate. Just ready. Cloth, $2.50, nff. Asa student's manual, it may be considered beyond criticism. The book is singularly full. St. Louis (td Surgical Journal. Without doubt forms one of the best guides for the beginner in der- matology that is to be found in the English language. Medic inc. JAMIESON (W. AJLL.AN). DISEASES OF THE SKIN. Third edition. In one octavo volume of 656 pages, with 1 engraving afld 9 double-page chromo-lithographic plates, Cloth, $6, LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. 17 JEWETT (CHARLES). ESSENTIALS OF OBSTETRICS. In one 12mo. volume of 356 pages, with 80 engravings and 3 colored plates. Cloth , $2 . 25. Just ready. An exceedingly useful manual for ing it in attractive and easily tangi- student and practitioner. The au- ble form. The book is well illus- thor has succeeded unusually well trated throughout. Nashville Jour. in condensing the text and in arrang- of Medicine and Surgery. American Authors. One large octavo volume of 763 pages, with 441 engravings in black and colors, and 22 full-page colored plates. Just ready. Cloth, THE PRACTICE OF OBSTETRICS. By ith 441 $5.00, net; leather, $6.00, net. A clear and practical treatise upon obstetrics by well-known teachers of the subject. A special feature of this work would seem to be the excellent illustrations with which the book abounds. The work is sure to be popular with medical students, as well as being of extreme value to the practitioner. The Medical Age. JONES (C. HANDF1EL.D). CLINICAL OBSERVATIONS ON FUNCTIONAL NERVOUS DISORDERS. Second American edi- tion. In one octavo volume of 340 pages. Cloth, $3.25. JULER (HEXRY). A HANDBOOK OF OPHTHALMIC SCIENCE AND PRACTICE. Second edition. In one octavo volume of 549 pages, with 201 engravings, 17 chromo-lithographic plates, test-types of Jaeger and Siiellen, and Holmgren's Color-Blindness Test. Cloth, $5.50 ; leather, $6.50. The volume is particularly rich in | color blindness, etc. The sections matter of practical value, such as | devoted to treatment are singularly directions for diagnosing, use of full and concise. Medical Age. instruments, testing for glasses, for | KING (A. F. A.). A MANUAL OF OBSTETRICS. Seventh edition. In one 12mo. volume of 573 pages, with 223 illustrations. Cloth, $2.50. From first to finish it is thoroughly j cyclopedias. The well-arranged ractical, concise in expression, well index renders the book useful to illustrated, and includes a statement of nearly every fact of importance the practitioner who is in haste to refresh his memory. Virginia discussed in obstetric treatises or ! Medical Semi-Monthly. KIRK (EDWARD C.). OPERATIVE DENTISTRY. Handsome octavo of 700 pages, with 751 illustrations. Just ready. See American Text-Books of Dentistry, page 2. We have only the highest praise for this valuable work. It is replete in every particular, and surpasses anything of the kind heretofore at- tempted. We can heartily recom- mend it to the profession. The Ohio Dental Journal. KLEIN (E.). ELEMENTS OF HISTOLOGY. New (5th) edition. In one 12mo. volume of 506 pages, with 296 engravings. Just ready. Cloth, $2.00, net. See Student's Series of Manuals, page 27. This work deservedly occupies a It is the most complete and con- cise work of the kind that has yet emanated from the press. The Med- ical Age. first place as a text-book on his- tology. Canadian Practitioner. 18 LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. L.ANDIS (HENRY G.). THE MANAGEMENT OF LABOR. In one handsome 12mo. volume of 329 pages, with 28 illus. Cloth, $1.75. LiA ROCHE (R.). YELLOW FEVER. In two 8vo. volumes of 1468 pages. Cloth, $7. LAURENCE (J. Z.) AND MOON (ROBERT C.). A HANDY- BOOK OF OPHTHALMIC SURGERY. Second edition. In one octavo volume of 227 pages, with 66 engravings. Cloth, $2.75. LEA'S SERIES OP POCKET TEXT-BOOKS, edited by BKRN B. GALLAUDET, M. D. Covering the entire field of Medicine in a series of 16 very handsome cloth-bound 12mo. volumes of 350-450 pages each, profusely illustrated. Compendious, clear, trustworthy and modern, and issued at the very moderate price of $1.50, net, per volume. The following volumes constitute the series. Co ATES' Bacteriology and Hygiene. BROCKWAY'S Anatomy. COM .INS and ROCKWELL'S Physiology. MARTIN and ROCKWELL'S < 'hemistry and Physics. NICHOLS and VALE'S Histology and Pathology. SCHLEIF'S Materia Medica, Therapeutics, Medical Latin, etc. MALS- BARY'S Practice of Medicine. COLLINS' Diagnosis. POTTS' Nervous and Mental Diseases. GALLAUDET'S Surgery. LIKES' Genito- Urinary and Venereal Diseases. GRINDON'S Dermatology. BALLKN- GER and WIPPERN'S Diseases of the Eve, Ear, Throat and Nose. EVANS' Obstetrics. CROCKETT'S Gynecology. TUTTLK'S Diseases of Children. For separate notices see under various authors' names. LEA (HENRY C.). A HISTORY OF AURICULAR CONFESSION AND INDULGENCES IN THE LATIN CHURCH. In three octavo volumes of about 500 pages each. Per volume, cloth, $3.00. CHAPTERS FROM THE RELIGIOUS HISTORY OF SPAIN; CENSORSHIP OF THE PRESS; MYSTICS AND ILLUMINATI- THE ENDEMONIADAS; EL SANTO NlftO DE LA GUARDIA; BRIANDA DE BARDAXI. 12mo., 522 pages. Cloth, $2.50. FORMULARY OF THE PAPAL PENITENTIARY. In one octavo volume of 221 pages, with frontispiece. Cloth, $2.50. SUPERSTITION AND FORCE; ESSAYS ON THE WAGER OF LAW, THE WAGER OF BATTLE, THE ORDEAL AND TORTURE. Fourth edition, thoroughly revised. In one hand- some royal 12mo. volume of 629 pages. Cloth, $2.75. STUDIES IN CHURCH HISTORY. The Rise of the Temporal Power Benefit of Clergy Excommunication. New edition. In one handsome 12mo. volume of 605 pages. Cloth, $2.50. AN HISTORICAL SKETCH OF SACERDOTAL CELIBACY IN THE CHRISTIAN CHURCH. Second edition. In one hand- some octavo volume of 685 pages. Cloth, $4.50. LEHMANN (C. G.). A MANUAL OF CHEMICAL PHYSIOLOGY. In one 8vo. volume of 327 pages, with 41 engravings. Cloth, $2.25, LEA BROTHERS & Co., PHILADELPHIA AND NEW "XORK. 19 LIKES (SYLVAN H.). A POCKET TEXT-BOOK OF GENITO- URINARY AND VENEREAL DISEASES. In one handsome 12rao. volume of about 350 pages, with many illustrations. Shortly. Cloth, $1.50, net. Lea's Series of Pocket Text-books, edited by BEEN B. GALLAUDET, M. D. See page 18. LOOMIS (ALFRED L.) AND THOMPSON (W. OILMAN, EDITORS). A SYSTEM OF PRACTICAL MEDICINE. In Contributions by Various American Authors. In four very hand- some octavo volumes of about 900 pages each, fully illustrated in in black and colors. Complete work now ready. Per volume, cloth, $5 ; leather, $6 ; half Morocco, $7. For sale by subscription only. Full prospectus free on application to the Publishers. See American System of Practical Medicine, page 2. LUFF (ARTHUR P.). Students of Medicine, engravings. Cloth, $2. MANUAL OF CHEMISTRY, for the use of In one 12mo. volume of 522 pages, with 36 See Student's Series of Manuals, page 27. LYMAN (HENRY M.). THE PRACTICE OF MEDICINE. In one very handsome octavo volume of 925 pages, with 170 engravings. Cloth, $4.75 ; leather, $5.75. Complete, concise, fully abreast of i Practical, systematic, complete and the times and needed by all students well balanced. Chicago Med. Re- and practitioners. Univ. Med. Mag. corder. An exceedingly valuable text-book. ' LYONS (ROBERT D.). A TREATISE ON FEVER. In one octavo volume of 362 pages. Cloth, $2.25. MACKENZIE (JOHN NOLAND). ON THE NOSE AND THROAT. Handsome octavo, about 600 pages, richly illustrated. Preparing. MAISCH (JOHN M.). A MANUAL OF ORGANIC MATERIA MEDICA. New (7th) edition, thoroughly revised by H. C. C. MAISCH, Ph. G., Ph. D. In one very handsome 12mo. volume of 512 pages, with 285 engravings. Just ready. Cloth, $2.50, net. Used as text-book in every college of pharmacy in the United States and recommended in medical col- leges. American Therapist. Noted on both sides of the Atlantic and esteemed as much in Germany as in America. The work has no equal. Dominion Med. Monthly. The best handbook upon phar- macognosy of any published in this country. Boston Med. & Sur. Jonr. LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. MALSBARY (GEORGE E.). A POCKET TEXT-BOOK OF THEORY AND PRACTICE OF MEDICINE. In one handsome 12mo. volume of about 350 pages. Cloth, $1.50, net. Shortly. Lea's r/Vx of Pocket Text-books, edited by BERN B. GALLAUDET, M. D. See page 18. MANUALS. See Students Quiz Series, page 27, Student's Series of Manuals, page 27, and Series of Clinical Manuals, page 25. MARSH (HOWARD). DISEASES OF THE JOINTS. In one 12mo. volume of 468 pages, with 64 engravings and a colored plate. Cloth, $2. See Series of Clinical Manuals, page 25. MARTIN (EDWARD). A MANUAL OF SURGICAL DIAGNOSIS. In one 12mo. volume of about 400 pp., fully illustrated. Preparing. MARTIN (WALTON) AND ROCKWELL, (WM. H.). A POCKET TEXT-BOOK OF CHEMISTRY AND PHYSICS. In one hand- some 12mo. volume of about 350 pages, with many illustrations. Cloth, $1.50, net. Shortly. Lea's Series of Pnr/.-rf Text-books, edited by I'.KKN B. GALLAUDET, M. D. See page 18. MAY (C. H.). MANUAL OF THE DISEASES OF WOMEN. For the use of Students and Practitioners. Second edition, revised by L. S. RAU, M. D. In one 12mo. volume of 360 pages, with 31 engrav- ings. Cloth, $1.75. MEDICAL NEWS POCKET FORMULARY, see page 32. MITCHELL (S. WEIR). CLINICAL LESSONS ON NERVOUS DISEASES. In one 12mo. volume of 299 pages, with 19 engravings and 2 colored plates. Cloth, $2.50. Of the hundred numbered copies with the Author's signed title page a few remain ; these are offered in green cloth, gilt top, at $3.50, net. The book treats of hysteria, recur- rent melancholia, disorders of sleep, choreic movements, false sensations of cold, ataxia, hemiplegic pain, treatment of sciatica, erytnromelal- gia, reflex ocularneurosis, hysteric contractions, rotary movements in the feeble minded, etc. Few can speak with more authority than the author. The Journal of the Ameri- can Medical Association. MITCHELL (JOHN K.). REMOTE CONSEQUENCES OF IN- JURIES OF NERVES AND THEIR TREATMENT. In one handsome 12mo. volume of 239 pages, with 12 illustrations. Cloth, $1.75. Injuries of the nerves are of fre- quent occurrence in private practice, and often the cause of intractable and painful conditions, conse- quently this volume is of especial interest. Doctor Mitchell has had access to hospital records for the last thirty years, as well as to the government documents, and has skilfully utilized his opportunities. The Med. Age. MORRIS (MALCOLM). DISEASES OF THE SKIN. New (2d) edition. In one 12mo. volume of 601 pages, with 10 chromo-litho- graphic plates and 26 engravings. Cloth, $3.25, net. Just ready. MULLER (J.). PRINCIPLES OF PHYSICS AND METEOROL- OGY. In one large 8yo. vol. of 623 pages, with 638 cuts. Cloth, $4.50. LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. 21 MUSSER (JOHN H.). A PEACTICAL TEEATISE ON MEDICAL DIAGNOSIS, for Students and Physicians. New (3d) edition, thor- oughly revised. In one octavo volume of about 1000 pages, with about 220 engravings and 48 full-page colored plates. In press. Notices of previous edition are appended. We have no work of equal value in English. University Medical Magazine. His descriptions of the diagnostic manifestations of diseases are accu- rate. This work will meet all the requirements of student and physi- cian. The Medical News. From its pages may be made the diagnosis of every malady that afflicts the human body, including those which in general are dealt with only by the specialist. North- western Lancet. It so thoroughly meets the precise demands incident to modern research that it has been adopted as a leading text-book by the medical colleges of this country. North American Practitioner. Occupies the foremost place as a thorough, systematic treatise. Ohio Medical Journal. The best of its kind, invaluable to the student, general practitioner and teacher. Montreal Medical Journal. NATIONAL DISPENSATORY. See Stille, Maisch & Caspari, p. 27. NATIONAL FORMULARY. See Stille, Maisch & Caspari' 8 National Dispensatory, page 27. NATIONAL MEDICAL DICTIONARY. See Billings, page 4. NETTLESHIP (E.). DISEASES OF THE EYE. New (5th) American from sixth English edition, thoroughly revised. In one 12mo. volume of 521 pages, with 161 engravings, and 2 colored plates, test-types, formulae and color-blindness test. Cloth, $2.25. Just ready. By far the best student's text-book on the subject of ophthalmology and is conveniently and concisely ar- ranged. The Clinical Review. It has been conceded by ophthal- mologists generally that this work for compactness, practicality and clearness has no superior in the English language. Journal of Medicine and Science. The present edition is the result of revision both in England and America, and therefore contains the latest and best ophthalmological ideas of both continents. The Phy- sician and Surgeon. NICHOLS (JOHN B.) AND VALE (F. P.). A POCKET TEXT- BOOK OF HISTOLOGY AND PATHOLOGY. In one handsome 12mo. volume of about 350 pages, with many illustrations. In press. Cloth, $1.50, net. Lea's Series of Pocket Text-books, edited by BERN B. GALLATJDET, M. D. See page 18. NORRIS (WM. F.) AND OLIVER (CHAS. A.). TEXT-BOOK OF OPHTHALMOLOGY. In one octavo volume of 641 pages, with 357 engravings and 5 colored plates. Cloth, $5 ; leather, $6. A safe and admirable guide, well best, the safest and the most conif re- qualified to furnish a working he nsive volume upon the subject that knowledge of ophthalmology. has ever been offered to the Ainer- Johns Hopkins Hospital Bulletin. ican medical public. Annals of It is practical in its teachings. Ophthalmology and Otology. We unreservedly endorse it as the 22 LEA BROTHKBS & Co., PHILADELPHIA AND NEW YOBK. OWEN (EDMUND). SURGICAL DISEASES OF CHILDREN. In one 12mo. volume of 525 pages, with 85 engravings and 4 colored plates. Cloth, $2. See Series of Clinical Manuals, page 25. PARK (ROSWEL.L.). A TREATISE ON SURGERY BY AMERI- CAN AUTHORS. New and condensed edition. JH /y/vxx. In one royal octavo volume of about 1250 pages, witli about 1000 engravings and many full-page plates. JS7' This work is also published in a larger edition, comprising two volumes. Volume I., General Suryery, 799 pages, with 356 engravings and 21 full-page plates, in colors and monochrome. Volume II., Special Surycry, 800 pages, with 430 engra- vings and 17 full-page plates, in colors and monochrome. Per volume, cloth, $4.50 ; leather, $5.50. Net. The work is fresh, clear and practi- way that they add great force to the cal, covering the ground thoroughly j text. The Chicago Medical Re- yet briefly, and well arranged for ! corder. rapid reference, so that it will be of The various writers have em- special value to the student and busy bodied the teachings accepted at practitioner. The pathology is the present hour. The North Amer- oroad, clear and scientific, while the ican Practitioner. suggestions upon treatment are Both for the student and practi- clear-cut, thoroughly modern and tioner it is most valuable. It is admirably resourceful. Johns Hop- I thoroughly practical and yet thor- kins Hospital Bulletin. oughly scientific. Medical News. The latest and best work written A truly modern surgery, not only upon the science and art of surgery. Columbus Medical Journal. The illustrations are almost en- tirely new and executed in such a in pathology, but also in sound surgical therapeutics. New Or- leans Med. and Surgical Journal. PARK (WILLIAM H.). BACTERIOLOGY IN MEDICINE AND SURGERY. 12mo., about 550 pages, fully illustrated. In press. PARRY (JOHN S.). EXTRA-UTERINE PREGNANCY, ITS CLINICAL HISTORY, DIAGNOSIS, PROGNOSIS AND TREAT- MENT. In one octavo volume of 272 pages. Cloth, $2.50. PARVIN (THEOPHIL.US). THE SCIENCE AND ART OF OB- STETRICS. Third edition. In one handsome octavo volume of 677 pages, with 267 engravings and 2 colored plates. Cloth, $4.25 ; leather, $5.25. In the foremost rank among the most practical and scientific medical works of the day. Medical News. It ranks second to none in the English language. Annals of Gyne- cology and Pediatry. The book is complete in every de- partment, and contains all the neces- sary detail required by the modern practising obstetrician. Interna- tional Medical Magazine. Parvin's work is practical, con- cise and comprehensive. We com- mend it as first of its class in the English language. Medical Fort- nightly. It is an admirable text-book in every sense of the \vord.-Nashville Journal of Medicine and Surgery. LEA BKOTHEKS & Co., PHILADELPHIA AND NEW YOKK. 23 PEPPER'S SYSTEM OF MEDICINE. See page 3. PEPPER (A. J.). FORENSIC MEDICINE. In press. See Student's Series of Manuals, page 27. SURGICAL PATHOLOGY. In one 12mo. volume of 511 pages, with 81 engravings. Cloth, $2. See Student's Series of Manuals, p. 27. PICK (T. PICKERING). FRACTURES AND DISLOCATIONS. In one 12mo. volume of 530 pages, with 93 engravings. Cloth, $2. See Series of Clinical Manuals, page 25. PLAYFAIR (W. S.). A TREATISE ON THE SCIENCE AND PRACTICE OF MIDWIFERY. Seventh American from the ninth English edition. In one octavo volume of 700 pages, with 207 engravings and 7 plates. Cloth, $3.75 net; leather, $4.75, net. Just ready. In the numerous editions which obstetrician. It holds a place among the ablest English-speaking authori- have appeared it has been kept con- stantly in the foremost rank. It is a work which can be conscientiously recommended to the profession. The Albany Medical Annals. This work must occupy a fore- most place in obstetric medicine as a safe guide to both student and ties on the obstetric &rt. Buffalo Medical and Surgical Journal. An epitome of the science and practice of midwifery, which em- bodies all recent advances. The Medical Fortnightly. THE SYSTEMATIC TREATMENT OF NERVE PROSTRA- TION AND HYSTERIA. In one 12mo. volume of 97 pages. Cloth, $1. POCKET FORMULARY, see page 32. POCKET TEXT-BOOKS, see page 18. POL.ITZER (ADAM). A TEXT-BOOK OF THE DISEASES OF THE EAR AND ADJACENT ORGANS. Second American from the third German edition. Translated by OSCAR DODD, M. D., and edited by SIR WILLIAM DALEY, F. R. C. S. In one octavo volume of 748 pages, with 330 original engravings. Cloth, $5.50. The anatomy and physiology of ment are clear and reliable. We each part of the organ of hearing are carefully considered, and then follows an enumeration of the dis- eases to which that special part of the auditory apparatus is especially liable. The indications for treat- can confidently recommend it, for it contains all that is known upon the subject. London Lancet. A safe and elaborate guide into every part of otology. American Journal of the Medical Sciences. POTTS (CHARLES S.). A POCKET TEXT-BOOK. OF NERVOUS AND MENTAL DISEASES. In one handsome 12ino. volume of about 450 pages. Cloth, $1.50, net. Shortly. Lea's Series of Pocket Text-books, edited by BERN B. GALLAUDET, M. D. See page 18. PROGRESSIVE MEDICINE, see page 32. PURDY (CHARLES WA BRIGHT'S DISEASE AND ALLIED AFFECTIONS OF THE KIDNEY. In one octavo volume of 288 pages, with 18 engravings. Cloth, $2. 24 LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. PYE-SMITH (PHILIP H.). DISEASES OF THE SKIN. In one 12mo. vol. of 407 pp., with 28 illus., 18 of which are colored. Cloth, $2. QUIZ SERIES. See Student's Quiz Series, page 27. KALFE (CHARLES H.). CLINICAL CHEMISTRY. In one 12mo. volume of 314 pages, with 16 engravings. Cloth, $1.50. See Student's Series of Manuals, page 27. RAMSBOTHAM (FRANCIS H.). THE PRINCIPLES AND PRAC- TICE OF OBSTETRIC MEDICINE AND SURGERY. In one imperial octavo volume of 640 pages, with 64 plates and numerous engravings in the text. Strongly bound in leather, $7. REICHERT (EDWARD T.). A TEXT-BOOK ON PHYSIOLOGY. In one handsome octavo volume of about 800 pages, richly illustrated. Preparing. REMSEN (IRA). THE PRINCIPLES OF THEORETICAL CHEM- ISTRY. New (5th) edition, thoroughly revised. In one 12mo. vol- ume of 326 pages. Cloth, $2. A clear and concise explanation that the work has met with general of a difficult subject. We cordially favor. This is further established recommend it. The London Lancet. \ by the fact that it has been trans- The book is equally adapted to the lated into German and Italian. The student of chemistry or the practi- tioner who desires to broaden his theoretical knowledge of chemistry. New Orleans Med. and Surg. Jour. The appearance of a fifth edition of this treatise is in itself a guarantee treatise is especially adapted to the laboratory student. It ranks unusu- ally high among the works of this class. This edition has been brought fully up to the times. American Medico-Surgical Bulletin. RICHARDSON (BENJAMIN WARD). PREVENTIVE MEDI- CINE. In one octavo volume of 729 pages. Cloth, $4 ; leather, $5. ROBERTS (JOHN B.). THE PRINCIPLES AND PRACTICE OF MODERN SURGERY. New (2d) edition. In one octavo volume of about 800 pages, with about 500 engravings. Shortly. THE COMPEND OF ANATOMY. For use in the Dissecting Room and in preparing for Examinations. In one 16mo. volume of 196 pages. Limp cloth, 75 cents. ROBERTS (SIR WILLIAM). A PRACTICAL TREATISE ON URINARY AND RENAL DISEASES, INCLUDING URINARY DEPOSITS. Fourth American from the fourth London edition. In one very handsome 8vo. vol. of 609 pp., with 81 illus. Cloth, $3.50. ROBERTSON (J. MCGREGOR). PHYSIOLOGICAL PHYSICS. In one 12mo. volume of 537 pages, with 219 engravings. Cloth, $2. See Student's Series of Manuals, page 27. ROSS (JAMES). A HANDBOOK OF THE DISEASES OF THE NERVOUS SYSTEM. In one handsome octavo volume of 726 pagee, with 184 engravings. Cloth, $4.50 ; leather, $5.50. SAVAGE (GEORGE H.). INSANITY AND ALLIED NEUROSES, PRACTICAL AND CLINICAL. In one 12mo. volume of 551 pages, with 18. typical engravings. Cloth, $2. See Series of Clinical Man- ualt, page 25. LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. 25 SCHAFER (EDWARD A.). THE ESSENTIALS OF HISTOL- OGY, DESCRIPTIVE AND PRACTICAL. For the use of Students. New (5th) edition. In one handsome octavo volume of 359 pages, with 392 illustrations. Cloth, $3.00, net. Just ready. Nowhere else will the same very moderate outlay secure as thoroughly useful and interesting an atlas of structural anatomy. The American Journal of the Medical Sciences. The most satisfactory elementary text-book of histology in the Eng- lish language. The Boston Med. and Sur. Jour. A COURSE OF PRACTICAL HISTOLOGY. New(2d) edition. In one 12mo. volume of 307 pages, with 59 engravings. Cloth, $2.25. The book very nearly approaches perfection. Methods are given with an accuracy of detail and prevision of difficulties which can hardly be overpraised. It bears eloquent tes- timony to the wide knowledge and untiring industry of its author. The Scottish Med. and Surg. Jour. SCHLEIF (WILLIAM). MATERIA MEDICA, THERAPEUTICS, PRESCRIPTION WRITING, MEDICAL LATIN, ETC. 12mo., 352 pages. Cloth, $1.50, net. Just ready. Lea's Series of Pocket Text-books. Edited by BERN B. GALLATJDET, M. D. See page 18. SOHMITZ AND ZUMPT'S CLASSICAL SERIES. Advanced Latin Exercises. Cloth, 60 cts. Schmidt's Elementary Latin Exer- cises. Cloth, 50 cents. Sallust. Cloth, 60 cents. Nepos. Cloth, 60 cents. Virgil. Cloth, 85 cents. Curtius. Cloth, 80 cents. SCHOFIELD (ALFRED T.). ELEMENTARY PHYSIOLOGY FOR STUDENTS. In one 12mo. volume of 380 pages, with 227 engravings and 2 colored plates. Cloth, $2. SCHREIBER (JOSEPH). A MANUAL OF TREATMENT BY MASSAGE AND METHODICAL MUSCLE EXERCISE. Octavo volume of 274 pages, with 117 engravings. SENN (NICHOLAS). SURGICAL BACTERIOLOGY. Second edi- tion. In one octavo volume of 268 pages, with 13 plates, 10 of which are colored, and 9 engravings. Cloth, $2. SERIES OF CLINICAL MANUALS. A Series of Authoritative Monographs on Important Clinical Subjects, in 12mo. volumes of about 550 pages, well illustrated. The following volumes are now ready : YEO on Food in Health and Disease, new (2d) edition, $2.50; CARTER and FROST'S Ophthalmic Surgery, $2.25 ; HUTCHINSON on Syphilis, $2.25; MARSH on Diseases of the Joints, $2; OWEN on Surgical Dis- eases of Children, $2; PICK on Fractures and Dislocations, $2; SAVAGE on Insanity and Allied Neuroses, $2. For separate" notices, see under various authors' names. SERIES OF STUDENT'S MANUALS. See page 27. SIMON (CHARLES B.). CLINICAL DIAGNOSIS, BY MICRO- SCOPICAL AND CHEMICAL METHODS. New (2d) edition. In one very handsome octavo volume of 530 pages, with 135 engravings and 14 full-page colored plates. Cloth, $3.50. Just ready. This book thoroughly deserves its success. It is a very complete, authen- tic and useful manual of the micro- scopical and chemical methods which are employed in diagnosis. Very excellent colored plates illus- trate this work. New York Medical Journal. In all respects entirely up to date. spects entirely up Record. Medical The chapter on examination of the urine is the most complete and advanced that we know of in the English language. Canadian Prac- titioner, 26 LEA BROTHERS & Co., PHILADELPHIA AND NEW YORK. SIMON (W.). MANUAL OF CHEMISTRY. A Guide to Lectures and Laboratory Work for Beginners in Chemistry. A Text-book specially adapted for Students of Pharmacy and Medicine. New (6th) edition. In one 8vo. volume of 536 pages, with 46 engravings and 8 plates showing colors of 64 tests. Cloth, $3.00, ><> t. Jnut ready. It is difficult to see how a better the covers of this book. The North- book could be constructed. No man western Lancet. who devotes himself to the practice Its statements are all clear and its of medicine need know more about teachings are practical. Virginia chemistry than is contained between Med. Monthly. SLADE (D. D.). DIPHTHERIA; ITS NATURE AND TREAT- MENT. Second edition. In one royal 12mo. vol., 158 pp. Cloth, $1.25. SMITH (EDWARD). CONSUMPTION; ITS EARLY AND REME- DIABLE STAGES. In one 8vo. volume of 253 pp. Cloth, $2.25. SMITH (J. LEWIS). A TREATISE ON THE DISEASES OF IN- FANCY AND CHILDHOOD. Eighth edition, thoroughly revised and rewritten and much enlarged. In one large 8vo. volume of 983 pages, with 273 engravings and 4 full-page plates. Cloth, $4.50; leather, $5.50. The most complete and satisfac- j can more than hold its own against tory text-book with which we are [ any other work treating of the same acquainted . American Gynecologi- cal and Obstetrical Journal. 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THE NATIONAL DISPENSATORY: Containing the Natural History, Chemistry, Pharmacy, Actions and Uses of Medicines, including those recognized in the latest Pharmacopeias of the United States, Great Britain and Germany, with numerous refer- ences to the French Codex. Fifth edition, revised and enlarged, including the new U. S. Pharmacopoeia, Seventh Decennial Revision. With Supplement containing the new edition of the National Formu- lary. In one magnificent imperial octavo volume of about 2025 pages, with 320 engravings. Cloth, $7.25; leather, $8. With ready reference Thumb-letter Index. Cloth, $7.75 ; leather, $8.50. Recommended most highly for the amount of information contained in physician druererist.- and invaluable to the ruggist. Therapeutic Gazette. It is the oflicial guide for the Med- ical and Pharmaceutical professions. Buffalo Med. and Sur. Jour. The readiness with which the vast this work is made available is indi- cated by the twenty-five thousand references in the two indexes. Bos- ton Medical and Surgical Journal. Should be recognized as a national standard. North Am. Practitioner. STIMSON (LJEWIS A.). A MANUAL OF OPERATIVE SURGERY. New (3d) edition. In one royal 12mo. volume of 614 pages, with 306 engravings. Cloth, $3.75. A useful and practical guide for all students and practitioners. Am. Journal of the Medical Sciences. The book is worth the price for the illustrations alone. Ohio Medical Journal. STIMSON (LEWIS A.). A TREATISE ON FRACTURES AND DISLOCATIONS. In one handsome octavo volume of 831 pages, with 326 engravings and 20 plates. Just ready. Cloth, $5.00, net ; leather, $6.00, net. Preeminently the authoritative Taken as a whole, the work is the text-book upon the subject. The best one in English to-day. *. vast experience of the author gives j Louis Medical and Surgical Journal. to his conclusions an unimpeachable i Pointed, practical, comprehensive, value. The work is profusely il- ! exhaustive, authoritative, well writ- lustrated. It will be found indis- ten and well arranged. Denver pensable to the student and the prac- Medical Times. titioner alike. The Medical Age. \ STUDENT'S QUIZ SERIES. Thirteen volumes, convenient, author- itative, well illustrated, handsomely bound in cloth. 1. Anatomy (double number); 2. Physiology; 3. Chemistry and Physics; 4. Histol- ogy, Pathology, and Bacteriology; 5. Materia Medica and Thera- peutics ; 6. Practice of Medicine ; 7. Surgery (double number); 8. Genito- Anatomy and Surgery, which being double numbers are priced at $1.75 each. Full specimen circular on application to publishers. STUDENT'S SERIES OF MANUALS. 12mos. of from 300-540 pages, profusely illustrated, and bound in red limp cloth. 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Ill Fifth Avenue, New York. 16 28 8-/ Y