>f Natural Historv bPJXSSF' Philadelphia. Pa. : ^mm I I : . ' U. (W. ON FOOD AND DIETETICS. TREATISE FOOD AND DIETETICS, PHYSIOLOGICALLY AND THERAPEUTICALLY CONSIDERED. BY F. W. PAVY, M.D., F.E.S., FELLOW OF THE ROYAL COLLEGE OF PHYSICIANS; PHYSICIAN TO, AND LECTURER ON PHYSIOLOGY AT, GUY'S HOSPITAL. PHILADELPHIA: HENRY C. LEA. 1874. PREFACE. IN the Preface to the second edition of my work on " Digestion, its Disorders and their Treatment," I mentioned that I had originally intended to add a section on Food to the contents of that volume, but that for the reasons given I afterwards determined to publish a separate treatise on the subject. Thus originated the present work, which, with the progress of time and a large consumption of mid- night oil, has grown to dimensions far exceeding those I had at first contemplated. From the fact that the subject of Food is one of deep concern, both to the healthy and the sick ; that the information which has been obtained during the last few years has completely revolutionized some of the cardinal scientific notions formerly entertained ; and that no modern systematic treatise of the kind here presented exists in the English language, I have been encouraged to think that the task I have undertaken may not be deemed superfluous. Whatever the results attained, I have steadily striven, sparing no pains for the purpose, to render the work produced instructive and useful. On account of the change recently introduced in chemical notation, I have given both old and new formulae, placing the latter within square brackets after the former. 35 GROSVENOR STREET, GROSVENOR SQUARE, March, 1874. CONTENTS. PAGE INTRODUCTORY REMARKS ox THE DYNAMIC RELATIONS OP FOOD, 17-24 Matter and Force, 17. Correlation of the Physical Forces, 17, 18. Equivalent of heat in mechanical motion, 18. Force and energy distinction explained, 18, 19. Analogy between living matter and a machine, 20. Forms of force derived from the sun, 20. Analogy between the animal system and a steam-engine, 22, 23. Life implies change, 23. Dormant vitality, 23. ON THE ORIGINATION OF FOOD, 25-35 Power possessed by animals of forming one kind of organic com- pound out of another, 25, 26. Influence of the solar force, 26-29. Action of vegetable life, 29-32. Formation of organic compounds, 33, 34. Kesults of animal and vegetable life, 34, 35. THE CONSTITUENT ELEMENTS OF FOOD, .... 36 ALIMENTARY PRINCIPLES : THEIR CLASSIFICATION, CHEMICAL RELATIONS, DIGESTION, ASSIMILATION, AND PHYSIOLOGI- CAL USES, a7-146 Distinction between alimentary principles and alimentary sub- stances, 37. Separation of food and drink not physiologically cor- rect, 37, 38. Classification of food, 38-40. THE NITROGENOUS ALIMENTARY PRINCIPLES, 41-96 : Albuminous or protein compounds animal and vegetable pro- tein compounds, 41-44. Gelatinous principles, 44, 45. Digestion of the nitrogenous principles, 45-48. Action of pancreatic juice, 48-50. Production of albuminose, 51, 52. Uses of nitrogenous matter, 53. Its relation to force-production, 54, 55. Experiments on the elimination of nitrogen, 55-72. Resume on nitrogenous food and muscular action, 72-74. Heat-production, 74, 75. Varied amounts of urea excreted on vegetable and animal diets, 75, 76 * CONTENTS. Metamorphosis of nitrogenous food, 77-86. Force value of nitro- genous food, 86-89. Nitrogenous matter as a source of fat, 90-93. Alimentary value of gelatinous principles, 93-96. THE NON-NITROGENOUS ALIMENTARY PRINCIPLES, 97-113: Hydrocarbons, or fats, 97-100. Uses of fat, 100-102. Fat as a heat-producing agent, 102-105. Oxidizable capacity of fat, 105, 106. Fat in relation to muscular force-production, 107-110. Ac- tual force-value of fat, 111-113. THE CARBOHYDRATES, 114-141 : Starch, 114-117. Sugars, 117-120. Gum, 120, 121. Dextrin, 122. Cellulose, 122. Lignin, 122. Lactic acid, 122. Assimila- tion and utilization of the carbohydrates, 123-126. Their destina? tion, 127. Power of animals to form fat, 128. Production of foie gras, 129-130. Conversion of the carbohydrates into fat, 130-134. Ultimate use of the carbohydrates, 134-136. Ternary principles not carbohydrates : Pectin, vegetable acids, alcohol, 136-141. THE INORGANIC ALIMENTARY PRINCIPLES, 142-146 : Water, 142. Saline matter, 142-146. ALIMENTARY SUBSTANCES, 147-396 ANIMAL ALIMENTARY SUBSTANCES, 147-223 : Their classification, 147, 148. Varieties of meat, 148-158. Un- wholesome meat, 158-167. Poultry, game, and wild fowl, 167-169. Fish, 169-177. Shell-fish, 177-181. Eggs, 181-185. Milk, 185- 199. Butter, 200-202. Cheese, 202-205. Animal foods sometimes but not ordinarily eaten, 206-223. Cannibalism, 206, 207. Mammals, 206-214. (Horseflesh, 211- 213.) Birds, 214-215. Reptiles, 216, 217. Fish, 217, 218. In- sects, 218, 219. Earth-eating, 219, 220. Table of references, 221- 223. VEGETABLE ALIMENTARY SUBSTANCES, 224-327 : Farinaceous seeds, 224-259. The cerealia, 225-254. Wheat and flour, 226-231. Bread, 231-237. Miscellaneous articles prepared from flour, 237-239. Unwholesome wheaten products, 240-242. Oats, 242-244. Barley, 245, 246. Eye, 246-248. Indian corn, 248-250. Rice, 250-252. Millet, 252, 253. Buckwheat, 253. Quinoa, 254. Leguminous seeds, or pulses, 254-259. Oleaginous seeds, 259-264. Tubers and roots, 264-276. Potatoes, 264-269. Herbaceous articles, 276-285. Products of the cabbage tribe, 277, 278. Various vegetables, 279-285. Fruity products consumed as vegetables, 285, 286. Esculent fungi, 286-289. Varieties of fruit, CONTENTS. XI 290-314. Bark, 315. Sawdust and wood}' fibre, 315. Saccharine preparations, 316-320. Saccharine products, 320-322. Farinaceous preparations, 322-327. BEVERAGES, 328-395: Water, 328-335. Non-alcoholic, exhilarating, and restorative beverages, 335-357. Tea, 336-342. Kepresentatives of tea, 343, 344. Coffee, 344-350. Fictitious coffee, 350. Chiccory, 350, 351, Gua- rana, 351, 352. Cocoa, 352-357. Fictitious cocoas, 357. Alcoholic beverages, 357-395. Effect of alcohol on the system, 358-361. Beer, 361-365. Cider, perry, 365, 366. Wine, 366-390. French wines, 381-384. German wines, 384, 385. Hungarian wines, 385. Greek and Italian wines, 386. Australian wines, 386. Port and other wines of Portugal, 386, 387. Sherry and other Spanish wines, 388, 389. Marsala, 389. Madeira, 389. Cape or South African wines, 389, 390. Fruit wines, 390. Mead, 390. Spirits, 390-394. Liqueurs, 394, 395. CONDIMENTS, 396. THE PRESERVATION OF FOOD, 397-402 Modern processes of preservation, 897. Four means of preserving food, viz., by cold, drying, exclusion of air, and use of antiseptics, 398-402. PRINCIPLES OF DIETETICS, 403-442 Composition of milk and the egg, 403. Researches of the Paris Gelatin Commission, 404-406. Position held by nitrogenous matter, 407-409. Question as to the necessity of fats and carbohydrates, 410. Adaption of food to demand, 411, 412. Liebig's estimate of the nutritive value of food, 413. Frankland's estimate of the force- producing value of food, 414-418. The appetite as a measure of capacity for work, 419. Nitrogenous matter required for physical development, 419, 420. Human labor more expensive than steam work, 421. Moleschott's table of a standard or model diet, 422. Adjustment of food to climate and work, 423-426. Table showing percentage composition of various articles of food, 427. Playfair's dietaries, 428-430. Workhouse dietaries, 430. Prison dietaries, 431, 432. Tables of hard and light labor diets, 433. Industrial employment, penal, and punishment diets, 434, 435. Instances of limited diet, 436, 437. Table from Payen of percentage value of food in nitrogen and carbon, 438, 439. Outgoing of nitrogen and carbon as a diet basis, 440-442. PRACTICAL DIETETICS, 443-502 Kind of food best adapted for the support of man, 443. Varieties of diet consumed by different nations : Arctic regions, 444-446 ; Xll CONTENTS. North American Indians, 446; Mexico, 447 ; Pampas Indians, 448 ; Guachos, 448, 449. Natives of Australia, New Zealand, 449, 450 ; of the Friendly Islands, 450; Otaheite, 450; Feejee Islands, 451 ; Tanna, New Caledonia, Savu, 451 ; Sandwich Islands, 452 ; China, 452,453; Japan, 453, 454; India, Ceylon, 455; Africa, 455-459. Mixed food the natural diet of man, 459. Vegetarianism, 460, 461. Dietetic value of meat often overestimated, 461. A certain amount of fresh food necessary to health, 462. Effects of animal and vege- table food compared, 463-467. Proper amount of food, 467-472. Effects of excess and deficiency of food, 472-476. Times of eating, 476-484. Culinary preparation of food, 484-491. DIET or INFANTS, 492-496. Woman's milk, 492-494. Milk of lower animals, 494, 495. Fari- naceous food, 496. Liebig's food, 496 DIET FOR TRAINING, 497-502. Object of training, 497. Old and new systems, 498-500. Oxford and Cambridge systems, 501, 502. THERAPEUTIC DIETETICS, . . . . . . . 503-559 General considerations, 503-506. Diet for gout, 506, 507. Influ- ences of food, 508. Principles of dieting for thinness and stoutness, 509, 510. Keduction of corpulency, 511-513. Dietary for the dia- betic, 513-515. Ill effects of restriction to salted and dried provi- sions, 515. Eegulation of amount of fluid, 516. Effect of varieties of food on the urine, 517-519. Food for weak digestion, 520-522. Food for dyspepsia, 522-524. Food for disordered states of the in- testinal canal, 525, 526. DIETETIC PREPARATIONS FOR THE INVALID, 527-536. HOSPITAL DIETARIES, 537-558. Guy's Hospital, 537. St. Bartholomew's Hospital, 538, 539. St. Thomas's Hospital, 539. London Hospital, 540. St, George's Hos- pital, 541. Middlesex Hospital, 542, 543. University College Hospital, 543, 544. King's College Hospital, 544. St. Mary's Hospital, 545. Westminster Hospital, 546. Seamen's Hospital, 547. Leeds General Infirmary, 547, 548. Manchester Royal Infir- mary and Dispensary, 548, 549. Birmingham General Hospital, 549, 550. Newcastle-upon-Tyne Infirmary, 551. Edinburgh Royal Infirmary, 551, 552. Glasgow Royal Infirmary, 553. Richmond, Whitworth, and Hardwicke Hospitals (Dublin), 553, 554. Bethlem Lunatic Hospital, 555. St. Luke's Hospital for Lunatics, 555, 556. Han well Lunatic Asylum, 556-558. Colney Hatch Lunatic Asy- lum, 559. INDEX, . 561-574 INTRODUCTORY REMARKS ON THE DYNAMIC RELATIONS OF FOOD. THE discoveries and inductions of the present age have thrown a new light on the physiology of food. Around us we have to deal with Matter and Force the one a substantive entity, the other appreciable only as a principle of ac- tion. It has long been known that (as cognizable in onr own era) matter can be neither created nor destroyed. It may be variously combined and modified, but it remains the same in essence and un- altered in amount. Force also has more recently been recognized as similarly conditioned; and in order that the bearings of food in relation to this principle may be understood, some preliminary con- siderations explanatory of the views now entertained regarding it are necessary. First, then, we may take it as accepted that, like matter, under present circumstances, force can be neither created nor destroyed. " Ex nihilo nihil fit " and " Nihil fit ad nihilum " form axioms that must be admitted to be incontrovertible. If we except the incon- siderable accession derived from the occasional descent of a meteoric body, the earth's matter remains fixed in amount. It is otherwise, however, with respect to force. Under the form of heat and light, force is constantly being transmitted to us from the sun ; and it is from the force thus derived that, in a manner to be explained further on, life on earth originates and is sustained. In enunciating his doctrine on the " Correlation of the Physical Forces," Grove demonstrated that one kind of force was capable of producing another. His views were first made known at a lecture delivered at the London Institution in 1842. The word "correla- tion" he employed as meaning "reciprocal production in other words, that any force capable of producing another may in its turn 2 18 ' 'INTRODUCTORY REMARKS. be produced by it." The position sought to be established was that heat, light, electricity, magnetism, chemical affinity, and motion, are all correlative, or have a reciprocal dependence that either might produce the others, and that neither could originate otherwise than by production from some antecedent force or forces. Just at this time the same field of inquiry was being investigated by other workers. While Grove was asserting that the great prob- lem awaiting solution in regard to the correlation of physical forces was the establishment of their equivalent of power, or their measu- rable relations to a given standard, Mayer, Joule, and Helmholtz were announcing the actual equivalents themselves. Mayer, of Germany, had the priority in the publication of his researches. As a member of the medical profession he approached the subject through its relation to physiology. In 1842 he pro- pounded, in its full comprehensiveness, the doctrine of the "Con- servation of Force." Nearly at the same time Mr. Joule, of Manchester, discovered the equivalent of heat in mechanical motion. He had been led to prose- cute researches in that direction, with the view of ascertaining the relative value of heat and motion for the advantage of engineering science. He found that what sufficed to raise the temperature of a pound of water one degree Fahrenheit would, under another mode of action, raise 772 pounds a foot high ; or, putting it conversely, the fall of 772 pounds of water from a height of one foot would give rise to an amount of heat sufficient to elevate the temperature of one pound to the extent of one degree Fahrenheit. Thus the mechani- cal work corresponding to the elevation of 772 pounds a foot high, or, what comes to the same thing, one pound 772 feet high, forms the dynamic equivalent of one degree of heat of Fahrenheit's scale. It is necessary to state here that the term " force," when used in a strict sense, is employed under a more limited acceptation now than formerly. Originally it represented what is now distinguished as both " force " and " energy." By " force," in a rigid significa- tion, is understood the power of producing energy ; by " energy " the power of performing work. To give an illustration : powder has force, the cannon-ball energy ; but to speak of the force of the cannon-ball is inexact. I may also remark that the words " actual " and " potential " are in frequent use to qualify the state in which energy is met with. By actual energy is meant energy in an active ON THE DYNAMIC RELATIONS OF FOOD. 19 state energy which is doing work. By potential energy, energy at rest energy capable of doing work, but not doing it. In a bent crossbow there is potential energy^ energy in a state of rest, but ready to become actual, or to manifest itself when the trigger is pulled. Again, actual energy is evolved from the sun. By vege- table life this is made potential in the organic compounds formed. In these organic compounds the energy is stored up in a latent con- dition ; potential energy is reconverted into actual energy when they undergo oxidation during combustion or in their utilization in the animal economy. The doctrine of the " Conservation of Energy " implies that energy is as indestructible as matter, that a fixed amount exists in the universe, and that, however, variously it may be modified, trans- ferred, or transformed in spite of all the changes of which it may be the subject throughout the realm of nature it cannot be created or annihilated, increased or diminished. The doctrine further im- plies that the diiferent forms of energy have their definite reciprocal equivalents ; that so much chemical energy, for instance, will pro- duce so much heat, which is the representative of so much motive power, and so on. The ascertained equivalents of heat and motive power have been already given. Accepted as applicable to the physical forces, the doctrine of the " Conservation of Energy " next began to be applied to living na- ture. Grove in his " Correlation of Physical Forces " (second edi- tion, p. 89), suggested that the same principles and mode of reason- ing adopted in his essay might answer equally for the organic as for the inorganic world, and that muscular force, animal and vegetable heat, &c., might, and one day would, be shown to possess similar definite correlations. He proceeded no further, however, remarking that he purposely avoided entering upon a subject not pertaining to hi> own field of science. At this time the general belief prevailed that the processes going on in the living body were determined by "vitality" or the "vital principle." The physical forces, it was supposed, were overruled in the living body by the vital principle. Without discussing whether we are to admit or deny the existence of this principle as a distinct operating force a question which has been handled by some of the leading men of science of the day we must, I think, concede, as a matter of experience, that in the living organism there are influences 20 INTRODUCTORY REMARKS. at play which have no existence in the dead matter around. Matter which has been impressed with life can produce effects which dead matter cannot. This does not conflict with the extension of the law of the " Conservation of Energy " to living nature. The effects produced may have their origin in the physical forces the living matter forming the medium through which they operate. With ar- tificial appliances force may be made to produce various effects, ac- cording to the nature of the instrument employed. With the same force in operation different kinds of work are performed, according to the character of the machine set in motion. Between the two living matter and a machine there exists an analogy which admits of being followed still further. It is only when in a certain state that matter is capable of forming the medium for the exercise of force in the production of living operations. Modify this state, and though there may be the same matter to deal with, yet it is no longer capable of fulfilling the same office it before performed. So, in the case of an ordinary machine ; it must possess a particular construc- tion before it can form the medium for the operation of force. Dis- arrange this construction, and, although the matter remains un- changed, the application of force is without its proper effect. Thus a disarranged machine may be compared with living matter devital- ized. In both, the capacity of being set in operation by force has existed, and in both that capacity has been lost. Further, it may be said that a machine in working order, but unoperated on by force that is, in a state of rest is like matter possessing vitality, but in a dormant state. Both are ready to move directly the proper force is supplied. Applying the law of the " Conservation of Energy " to living na- ture, the forms of force which we observe in operation are, in the first instance, derived from the sun. When a weight is lifted by the hand it seems a long way off to go to the sun for the muscular force employed in the act. Yet the doctrine of the " Conservation of Energy " justifies, as I will proceed to show, the conclusion that its origin is there. In the first place, the force evolved in muscular action has its source in the material which has been supplied to the body in the form of food. Now, all food comes primarily from the vegetable kingdom, and vegetable products are built up through the agency of the sun's rays. It may be said that the energy contained in these ON THE DYNAMIC RELATIONS OF FOOD. 21 rays, which has been employed in producing the compound, is fixed or rendered latent within it. When a crossbow is bent, the force derived from the muscular action employed in bending it is stored up, ready to be again liberated when the trigger is pulled, no matter whether this be at once or a hundred years hence ; and the force given to the arrow when it is launched is neither more nor less than that which has sprung from the muscular action employed in bend- ing the bow. The same with vegetable products. Their formation is coincident with the disengagement of oxygen from oxidized prin- ciples and the development of combustible compounds. To effect this disengagement the operation of force is required. Now, the force so employed has its source in the heat and light evolved from the sun, and that which is used for the purpose may be said to be- come fixed and to exist in a latent condition to exist stored up in the product, ready to be again liberated on exposure to conditions favorable to oxidation. Thus may these vegetable products be com- pared to a bent crossbow, containing as they do a store of latent force, which may for an indefinite period remain as such, or may be liberated soon after it has been fixed. Whenever liberated, it is no more nor less than the equivalent of the force which has been used in the formation of the product. Our coal-fields represent a vast magazine of force drawn, ages ago, from the sun's rays, and capable at any moment of being set free by the occurrence of oxidation. Vegetable products, then, may be regarded as containing a store of force accumulated from the vast supply continually emitted with the sun's rays ; and, upon the principle of the indestructibility of force, that force which has been applied to unlocking the elements in the combinations from which vegetable products are built up, and to forming the new compound, is contained in such compound in a latent state. Now, as above stated, animals either directly or in- directly subsist upon these vegetable products, and are thence sup- plied by them with accumulated force. By oxidation the force is set free in an active state under some form of manifestation or other. It matters not in what way whether rapidly or slowly, or under what circumstances whether inside or outside the living system, the oxidation occurs ; the result is the same, as far as the amount of the force liberated is concerned, it being implied in the doctrine of the " Conservation of Energy" that it should constitute the equivalent of the solar force originally made use of. This is presuming com- 22 INTRODUCTORY REMARKS. plete oxidation to occur ; but in the processes of animal life, although fully oxidized compounds, like carbonic acid and water, are formed and discharged, yet others, like urea, are expelled in an imperfectly oxidized state, and carry with them a certain amount of latent or unutilized force. Thus it is that the various forms of force manifested in the actions of animal life trace their origin to that emitted from the sun. Plants are media for fixing solar force for converting actual into latent or potential energy. Animals reconvert latent into various forms of actual force. Thus, in the various forms of actual force liberated by the actions of animal life, we have the equivalent of that which has been fixed by plants from the sun. As there is a revolution of matter, so is there a revolution offeree within and around us. In the liberation of actual force a complete analogy may be traced between the animal system and a steam-engine. Both are media for the conversion of latent into actual force. In the animal system, combustible material is supplied under the form of the various kinds of food, and oxygen is taken in by the process of respiration. From the chemical energy due to the combination of these, force is lib- erated in an active state ; and, besides manifesting itself as heat, and in other ways peculiar to the animal system, is capable of perform- ing mechanical work. The steam-engine is supplied with combus- tible material under the form of coal, which differs from our food in representing the result of the vegetative activity of a former instead of the present epoch. Air is also supplied, and from the combina- tion which occurs between its oxygen and the elements of the com- bustible material, heat is produced, which in part is dissipated as such, but in part is applied to the performance of mechanical work. According to Helmholtz, the animal economy, in respect of its capacity to turn force to account in the accomplishment of mechanical work, is a more perfect instrument than the steam-engine. His calculations lead him to conclude that whilst in the best steam- engine only one-tenth of the force liberated by the combustion of its fuel is realizable as mechanical work, the rest escaping as heat, the human body is capable of turning one-fifth of the power of its food into the equivalent of work. There is this, however, to be remarked, that the fuel of a steam-engine is a far less expensive article than the food of an animal being. The animal body, then, may be regarded as holding an analogous ON THE DYNAMIC RELATIONS OF FOOD. 23 position to a machine, in which a transmutation of chemical into other forms of force is taking place. Food on the one hand, and air on the other, are the factors concerned in the chemical action that occurs. It is through the interplay of changes between food and air that the manifestations of animal life, consisting of heat- production, muscular contraction, nervous (including mental) action, and nutritive or formative, secretory, and assimilative action arise. The egesta, or substances dismissed from the system, are metamor- phosed products of the ingesta, or substances entering the system. The elements are the same, in nature and in quantity, in the two cases, but their forms of combination, and, with them, their force accompaniment, are different. The force employed in building up the organic compounds belonging to food is again evolved as they descend by oxidation into more simple combinations, and in the force evolved we have the representative of the active manifestations of animal life. If the products discharged from the system were fully oxidized principles, the force developed in the body would equal that contained in a latent condition in the food. Such, how- ever, is not completely the case, a certain amount of latent force remaining, as has already been remarked, in some of the egesta. The position, therefore, may be formulated thus : The latent or potential force of ingesta equals the force developed in the body plus the force escaping with the egesta. In other words, the unexpended force in the egesta and the force disengaged by the operations of life, and manifested under the various forms of vital activity, equal the force contained in the ingesta. AVhat is required in food is matter that is susceptible of under- \ going change in the system under the influence of the presence of I oxygen. Life implies change, and the manifestations of life are due to the reaction of food, with the derivatives from it, and air upon each other. While in the inorganic kingdom a tendency to a state of rest prevails while the closest affinities tend to become satisfied, and so establish equilibrium, in a manifestly living body rest is im- pnssible. It is true, living organisms of certain kinds may exist in a state of rest, but then there is a suspension of vital manifestations. The state constitutes that which falls under the denomination of " dormant vitality." Animal organisms may exist in it, and the seed of a plant naturally remains for awhile in it. Molecular rest, and, with it, an absence of any show of vital activity prevail. Con- 24 INTRODUCTORY REMARKS, ETC. currently, however, with the manifestations of vital activity molec- ular change change in a particular or prescribed direction occurs. Organic compounds become resolved by the agency of oxygen into more simple combinations, as carbonic acid, water, and urea, and cease to be any longer of service. To maintain a continuance of vital activity fresh organic material is required ; hence the demand for food. But food and the other material factor of life oxygen do not constitute all that is needed. It is further necessary that the two should be brought within the sphere of influence of living matter, in order that the changes may be made to pursue the par- ticular line of direction resulting in the phenomena of life. ON THE ORIGINATION OF FOOD. OUR food is in the first instance derived from the vegetable king- dom. Dumas at one time said, "L'animal s'assimile done ou detruit des matieres organiques toutes faites; il n'en cree done pas." But, as he afterwards admitted, this is not the case. The animal, it is true, is constantly consuming or destroying organic substances, and is incapable of forming them from the inorganic principles, but sup- plied with organic matter, organic compounds of various kinds are constructed. Mulder's discoveries in 1838 led up to the doctrine that the albu- minous compounds of plants and animals agree in composition and properties, whence it was inferred that the animal simply took the compound produced by the plant and made it a component part of its own bod^. Liebig was the first to maintain that animals pos- sessed the power of forming one kind of organic compound out of another. A warm controversy was at one time carried on upon this point, turning particularly upon the formation of fat. While, on the one hand, it was held by Liebig that, in the animal system, fat could be formed from sugar, Dumas and Boussingault maintained, on the other, that whatever fat was found in an aninal being was derived through its food from without. From the researches ini- tiated by this dispute it became incontestably established that Liebig was right, and the French chemists were ultimately compelled, even on the evidence of the results obtained by themselves, to abandon the doctrine they had advanced. A moment's consideration will further suffice to show that one kind of albuminous compound is capable of being-constructed from others. In the young mammal subsisting solely on milk, it is to the casein that we must look for the source of fibrin and albumen ; and in the animal feeder, secreting milk, the casein produced is de- rived from the fibrin and albumen. Gelatin, moreover, has no ex- 26 ORIGINATION OF FOOD. istence in vegetable food. At the present day we may waive the discussion of this matter, it being now established that none of these nitrogenous principles enter the system under the form in which they occur in food. They are all converted, during the performance of digestion, into a certain principle (albuminose), which is the princi- ple that is absorbed, and that is subsequently transformed by the assimilative power of the animal into the various compounds met with. The position, then, is this, that animals are not simply consumers of organic compounds, but are capable of exerting a constructive action as well. They must, however, be supplied with organic mat- ter previously formed, and thus the capacity that really exists is that of transforming one organic compound into another. All organic matter has its primary source in the vegetable kingdom, from which kingdom, it follows, all our food must directly or indirectly be de- rived. The vegetable feeder goes directly for its food to the vege- table kingdom. The animal feeder is equally dependent upon the products of the vegetable kingdom for its pabulum. But it obtains it only at secondhand, so to speak, or in an indirect manner, its food consisting of the flesh of animals which have themselves been nour- ished upon vegetable products. Now, it is only under exposure to the action of the sun's rays that plants will grow, and hence it is to the influence of these rays that we must refer the production of food in the first instance, and the primary source of all life upon our earth. It has already been shown how the energy emitted from the sun, under the forms of heat and light, is capable, through the medium of the plant, of disengaging oxygen from its combination with car- bon and hydrogen in carbonic acid and water, and leading to the formation of re-oxidizable compounds ; and how the energy evolved from the re-oxidation of these compounds, whether by combustion or within the animal system, represents or forms the equivalent of that employed in effecting their construction. What an immeasurable amount of force to be, and to have been, emitted from the solar centre ! It is true that it must possess a store of heat altogether unrealizable by comparison with anything cogniza- ble around us ; for it has been shown by recent investigations with the spectroscope that iron and other metals, which cannot by any known method of heat application be converted into the gaseous INFLUENCE OF THE SOLAR FORCE. 27 state upon our earth, exist in that state around the sun. It is true, also, that the sun is a body of almost inconceivable magnitude. To give the simile of Helmholtz, 1 " Its diameter is so great that if you- suppose the earth to be put into the centre of the sun, the sun itself being like a hollow sphere, and the moon going about the earth, there would be a space of more than 200,000 miles around the orbit of the moon lying all interior to the surface of the sun. 77 But when we come to consider that in that small pencil of rays which has im- pinged upon our earth at a distance of nearly 95,000,000 miles from the sun, that, under the view now held by philosophers, has been contained all the energy or source of power which has been fixed by plants, and much besides that has escaped being so utilized, we can- not help being struck by the immensity of the store of power accu- mulated in the sun. Geology teaches us that at an early epoch in the history of our globe this solar influence must have manifested itself to a much stronger degree than it does even at the present time. The vast coal-beds forming a portion of the earth's crust have origi- nated in vegetable growth. During the carboniferous era, which comprises the period of this coal-formation, the atmosphere was probably laden with carbonic acid and humidity to a much greater extent than at the present day. But it is to the solar energy that we -must look as the source of the luxuriant vegetation which evi- dently flourished at that time, and which must have existed in the Arctic regions as well as in the lower latitudes, since coal-deposits are there found. It has been already stated that it is only under the influence of the force contained in the sun's rays that organic compounds are built up by the agency of the plant; and it is found to be the green parts only of plants those where chlorophyll exists that effect the decomposition of carbonic acid and water fixing the carbon and hydrogen and liberating the oxygen. This operation, it is the dis- tinctive function of the plant to perform, and it fails to be carried on when the influence of light is absent and unless chlorophyll is pres- ent. Under these conditions absence of light and chlorophyll oxygen is absorbed and carbonic acid liberated instead, just as occurs in the animal. I have been informed that it is known to florists, as 1 "Lectures on the Conservation of Energy," Medical Times and Gazette, vol. i, p. 415, 1864. 28 ORIGINATION OF FOOD. the result of practical observation, that in the case of the variegated- leaved geranium, a slip that may happen to be possessed of white leaves only will not grow alone like other slips. The absence of chlorophyll explains the non-capacity to effect the changes necessary for growth. The solar beam is composed of rays possessing different properties and different degrees of refrangibility, and the question has been raised what part of the solar spectrum exerts greatest power over vegetable growth? The colored rays produced by passing a pencil of light through a prism are arranged in the following order : Violet, Yellow, Indigo, Orange, Blue, Ked. Green, The greatest illuminating power of the spectrum, is in the brightest yellow rays, whilst the greatest heating power is in rays below the red, and therefore less refrangible than any of the colored rays ; while the greatest chemical power power of effecting chemical change is in the rays at the other extremity of the spectrum, namely, the violet, and still more the invisible rays just above. Draper, from experiments conducted in 1843, states that on causing plants to effect the decomposition of carbonic acid in the prismatic spectrum, he found the yellow rays by far the most effec- tive. The relative power of the various colored rays he asserts to have been' as follows : Yellow, Blue, Green, Indigo, Orange, Violet. Ked, In opposition to the conclusion arrived at by Draper, it is affirmed by others that it is to the blue and violet rays that must be referred the maximum power of effecting the decomposition of carbonic acid through the medium of the plant. Helmholtz 1 says: "The obser- vations upon vegetable life have shown that plants can grow only 1 "Lectures on the Conservation of Energy," Medical Times and Gazette, vol. i, 1864, p. 473. ACTION OF VEGETABLE LIFE. 29 under the influence of solar light, and that as long as solar light, and principally the more refrangible parts of solar light, the blue and violet rays, fall upon the green parts of plants, the plants take in carbonic acid and exhale oxygen." He further remarks that in ex- erting this influence these rays are completely absorbed ; for it can be shown that solar light which has passed through green leaves in full development is no longer capable of exerting any chemical in- fluence. I have spoken of light as a factor in the construction of organic compounds by the plant. The elements of which these organic compounds consist are drawn from the inorganic kingdom, and chiefly, as Liebig pointed out, from carbonic acid, water, and am- monia principles which all exist to a greater or less extent in the atmosphere, and from the atmosphere are to a large extent, if not entirely, derived. In the case of the low vegetable organisms which become developed in moist situations as a green layer on the barren surface of rocks and stones, the elements required for their growth must have been derived solely from the atmosphere. In the case of the higher organisms, however, the elements of growth are drawn from the soil as well as the atmosphere. Humus, which forms the constituent of the soil which supplies these elements, consists of the decaying remains of organic products. But it is not as organic matter that humus serves as food to the plant ; that is, it is not the organic matter itself that is utilized. It is, on the other hand, as a source of carbonic acid and ammonia, principles resulting from its decomposition, that it owes its position in relation to the alimenta- tion of plants. The stages passed through in the history of vegetable life leading to the provision of a fitting supply of food for animal existence may be thus represented : Beginning, let us say, with a barren surface of rock, which may have been freshly exposed to the atmosphere from some subterranean, volcanic, or other agency, the germs of low vege- table organisms settling upon it, extract from the atmosphere their elements of growth. Passing through their term of life they die, and fresh ones spring up and similarly live and die. So the process goes on, higher and higher forms making their appearance. The decaying remains of this primitive growth incrust what was a barren surface with a layer of earth or mould, in which ultimately the highest plants find a suitable position for taking root and growing. 30 ORIGINATION OF FOOD. Thus clothed with vegetation, a fit locality is provided for the sup- port of animal life, animal beings finding in the vegetable products now existing the necessary material for their subsistence. It may be mentioned here that there is one class of vegetable organisms the Fungi which seems to occupy an exceptional posi- tion, and to resemble animals in being dependent upon organic prod- ucts for their growth. It is possible, however, that the seeming appropriation of organic matter may be more apparent than real, and that the dependence upon organic matter may arise from a specially large and constant supply of carbonic acid and ammonia being required as a condition of growth. Still, it must be said that this class of vegetable organisms is not dependent for growth upon light like the others, has no green surfaces for decomposing carbonic acid, and, in fact, instead of absorbing carbonic acid and setting free oxygen, does as is the case with animals, precisely the reverse. These are strong arguments in favor of growth from an appropri- ation of organic compounds ; but there is this to be remarked, that the growth in question occurs only where decay is going on, and there is nothing, at all events, to show that any other than organic compounds in a state of decomposition can be made use of. The chief elements of the various organic compounds built up by the agency of vegetable life are carbon, hydrogen, oxygen, nitrogen, sulphur, and phosphorus ; and the following may be regarded as the sources from which they are derived. In the above enumeration carbon is mentioned first as being the element which occurs by far the most extensively in organic nature. Large as is the quantity of carbon entering into the composition of organic substances, the main, if not the entire source from which it is derived is the carbonic acid in the atmosphere. According, how- ever, to Saussure, the amount of carbonic acid contained in air is not, as a mean, more that one part, by volume, in two thousand ; but then it must be remembered that it is constantly being generated, not only as a product of animal life, but from various processes carried on around us. Now, it appears that the leaves and other green parts of plants are continually absorbing this carbonic acid, and, with the aid of light, effecting its decomposition, the oxygen being exhaled and the carbon detained and applied to the production of organic substances. Whilst it is only by the leaves and green surfaces that carbonic acid ACTION OF VEGETABLE LIFE. 31 is decomposed and oxygen liberated, it is probable that its absorp- tion is not limited to those parts, but that some enters through the roots, this being derived from the. process of decomposition going on in the organic matter of the soil, and from the carbonic acid carried down from the atmosphere with the rain. Striking as it may seem, yet there are sufficient grounds for be- lieving that the vast store of carbon contained in forests, of whatever extent we may encounter, has been derived in the manner above mentioned. Geological investigations render it almost certain that at one time the atmosphere was far richer in carbonic acid than it is now, and that vegetation also was proportionately more luxuriant. The absorption of carbonic acid and exhalation of oxygen which take place in plants under the influence of light constitutes, then, a process of alimentation. The reverse process the absorption of oxygen and exhalation of carbonic acid a process which forms one of the principal phenomena of animal life occurs also to some ex- tent in plants, and stands out unconcealed during the night, when from the absence of light there is no decomposition of carbonic acid and liberation of oxygen going on. It also occurs as the result of certain operations of plant life, as, for instance, during germina- tion, flowering, and fruiting. Hydrogen and oxygen are supplied to an unlimited extent to plants under the form of water. In the production of the carbo- hydrate group of organic compounds; that is, compounds such as starch, sugar, dextrin, gum, cellulose, &c., in which carbon is united with hydrogen and oxygen in the proportion to form w r ater, it is possible that water is directly assimilated, although this is by no means an ascertained fact. In a large number of other compounds, however, it is evident from their composition that for water to serve for their production its elements must undergo separation. The oleaginous compounds, for instance, chiefly consist of carbon and hydrogen. The amount of oxygen present is very much less than that required to form water with their hydrogen. For this element to be appropriated a deoxidation must occur, and it is believed that some of the oxygen exhaled by the plant under the influence of light has its source, not only in carbonic acid, but likewise in water. Although plants are freely surrounded with nitrogen this ele- ment forming the large constituent it does of the atmosphere, yet it is not from the atmosphere that the nitrogen of organic matter is 32 ORIGINATION OF FOOD. derived. The researches of Saussure and Boussingau.lt have demon- strated that plants are incapable of appropriating the free nitrogen of the atmosphere and elaborating it into organic matter. Liebig's view, and it is one which is by common consent indorsed, is that the nitrogen of organic matter is derived from ammonia. This able chemist was the first to show that ammonia is a constant constituent of the atmosphere. It is true that the quantity in which it is present is so small that it cannot be recognized except by abstraction from a large volume of air. It may be removed and its quantity determined (" On the Estimation of Ammonia in Atmospheric Air/ 7 by Horace T. Brown, " Proceedings of the Royal Society," vol. xviii, p. 286) by passing a given volume of air through water slightly acidulated with sulphuric acid. It is also susceptible of recognition in rain-water, where it exists under the form of carbo- nate. Ammonia, like carbonic acid, forms a product of the decom- position of organic matter. The nitrogen of organic matter, indeed, is returned to the inorganic kingdom under the form of ammonia. Thus in humus we have a source of ammonia which, doubtless, com- bines with some of the carbonic acid also generated, and in this state is in great part dissipated into the atmosphere. The great volatility of the product would lead to this result. Diffused through the at- mosphere it would "be abstracted by rain and snow, and in this way carried back to the earth, to be brought in contact with the roots of plants, through which its absorption is supposed to be effected. Ac- cording to Liebig, ammonia enters the vegetable organism in com- bination with carbonic or sulphuric acid, while, according to Mulder, the combination is with the acids he describes as existing in humus. Nitrogen forms an element which is of the highest importance to vegetable as well as to animal life. It is not only necessary that it should enter into the constitution of vegetable substances so that animals may obtain a supply of it with their food, but it forms an indispensable element in relation to the molecular changes of the plant as well as of the animal. Wherever living changes are carried on nitrogenized matter is present. The proclivity of this to change forms one of its most characteristic qualities, and the changes it undergoes induce changes of a definite kind in other matter which per se has a tendency to remain at rest. Thus, in nitrogenized mat- ter we have, as it were, the requisite starting-point for the various changes which result in the phenomena of life. FORMATION OF ORGANIC COMPOUNDS. 33 The four elements which have been referred to, viz., carbon, hydro- gen, oxygen, and nitrogen, form by far the chief constituents of or- ganic compounds, but sulphur and phosphorus are also present, to a small extent, bound up with the other elements in certain organic principles. Sulphur, for example, is met with in casein, and both sulphur and phosphorus in fibrin and albumen. The probable source of these elements is sulphates and phosphates, the acids of the salts undergoing deoxidation through the medium of the operations carried on in the plant, in the same manner as occurs in the case of carbonic acid. As yet I have been referring merely to the source of the elements entering into the constitution of the^organic compounds produced by plants, and upon this point it may be considered that our infor- mation is pretty definite. The precise mode, however, in which these elements are combined or elaborated into the infinite variety of organic compounds existing is quite another matter, and one which (it must be conceded) belongs as yet only to the domain of hypothesis. The point has been the subject of many laborious re- searches, conducted by some of the most distinguished observers, but, in spite of these attempts to elucidate it, we have at present little or nothing beyond conjecture to deal with. It may be fairly surmised, however, that the production of the higher compounds is effected step by step, or by a series of transition stages, and not by a direct or immediate union of the elements entering into their com- position. Whatever the exact changes that ensue, there can be no doubt that they proceed in a definite and precise order. In organic nature we know that change induces change, and the change first set in motion in the act of growth may be regarded as starting the changes which produce the various organic compounds met with. Bodies in contact with changing matter are within the sphere of influence of a metabolic or metamorphosing force, and to the opera- tion of this force is to be ascribed much that occurs as the result of living action. It is the formation of organic compounds which constitutes the special province of the plant to effect in relation to the production of food. Food, however, to fulfil the requirements of animal life, must contain certain mineral or inorganic as well as organic prin- ciples a supply of the former being quite as indispensable as a supply of the latter. But we need not concern ourselves about a 3 34 ORIGINATION OF FOOD. separate supply of mineral matter. The productions of nature wisely contain in combination all that is wanted. It happens that, besides being furnished with carbonic acid, water, and ammonia, for the formation of organic compounds, plants require for their growth a supply of saline principles. These they draw from the surrounding soil, and a portion of the advantage accruing to vegetable growth from the employment of manure is owing to the mineral matter it contains, and which is thereby given to the soil. In appropriating mineral matter as an element of nutrition, the plant exercises a selective action. It is found, for instance, that some of the saline compounds belonging to the soil, and not others, are present, that they are present in different proportions as regards each other, and to a different extent in different parts of the plant. Min- eral matter holds, in fact, a definite relation to the component parts of a plant, and probably enters into some sort of combination with the organic constituents. Thus, in vegetable products we find not only the organic, but like- wise the inorganic matter we require ; and, in taking up and apply- ing mineral matter as it does to its own purposes of growth as well as forming organic compounds, the vegetable organism contributes in a complete manner towards the supply of what is wanted for ani- mal nutrition. A reciprocal relation, however, it must be observed, in reality exists between what is supplied and what is wanted. We are as much adapted to the appropriation of the food supplied to us as our food is adapted to our wants. Were we not so adapted existence would be impossible for us. In nature all things are mutually adapted to each other. In what has been said about the production of food by the vege- table kingdom for animal subsistence, it is seen that animals and plants stand in direct antagonism to each other, as regards the re- sults of the main operations of life. Plants draw their food from the inorganic kingdom, and produce organic compounds. Animals find their food in these organic compounds, and, in applying them to the purposes of life, reconvert them into inorganic principles. In the appropriation of inorganic matter, as food, plants absorb car- bonic acid, and set free oxygen. Animals, in their consumption of organic matter, absorb oxygen and give out carbonic acid. Thus, animal life and vegetable life stand in complemental relation to RESULTS OF ANIMAL AND VEGETABLE LIFE. 35 each other, and it is in accordance with the requirements for the persistence of living nature upon the surface of our planet that it should be so. If the operations of animal and vegetable life pro- ceeded in one and the same direction only, the effect would be a gradual alteration of the chemical arrangement of matter, until a state of things was arrived at unfit for the further continuance of life. Under the existing order of things animals and plants in such a manner neutralize each other's effects upon surrounding matter that they balance each other's operations, and thereby maintain a state of uniformity. THE CONSTITUENT ELEMENTS OF FOOD. OF the various elements known to exist in nature only a limited number enter into the constitution of living bodies. The following is a list of those found as constituents of the human body. The first four, namely, carbon, hydrogen, oxygen, and nitrogen, exist in far larger quantity than any of the others. As for those which occur towards the end of the list, they are present only in exceedingly minute quantity, if, indeed, they are invariably present. It is more than doubtful if they are to be regarded as essential constituents. Carbon, Hydrogen, Oxygen, Nitrogen, Sulphur, Phosphorus, Chlorine, Sodium, Potassium, Calcium, Magnesium, Iron, Fluorine, Silicon, Manganese, Aluminium, Copper. The food being the source from which the elements forming the constituents of the body are derived, it follows that food must con- tain all the elements which are there met with. No article can, as food, satisfy the requirements of life that fails to comply with this condition. ALIMENTARY PRINCIPLES: CLASSIFICATIONS, CHEMICAL RELATIONS, DIGESTION, ASSIMILATION, AND PHYSIOLOGICAL USES. ALTHOUGH it is necessary that our food should contain the ele- ments that have been enumerated and contain them in such pro- portion as to furnish the requisite amount of each to the system yet it is not with these elements as such that, from an alimentary point of view, we have to deal. It is only in a state of combination that the elements are of any service to us as food ; and, as has been already mentioned, the combination must have been formed by the agency of a living organism the combination must, in other words, constitute an organic product. Xow, taking the different organic products which nature affords us as food, we find that they may, by analysis, be resolved into a variety of definite compounds. These constitute what are known as "alimentary principles," in contradistinction to "alimentary sub- stances," or the articles of food as supplied to us by nature. In a scientific consideration of food it is necessary to speak first of the alimentary principles. It is only, indeed, by looking at it through its constituent principles that we are in a position to dis- cuss its physiological bearings, and I will begin by pointing out the most convenient division and classification to be adopted. Popularly, the ingesta are looked upon as consisting of food and drink, the one supplying us with solid, the other with liquid, matter. Superficially, this appears a natural and convenient mode of primary grouping, but in a physiological point of view it is completely worth- less. " Food " and " drink " constitute terms referring only to the particular state in which an article for consumption may happen to 38 ALIMENTARY PRINCIPLES. exist viz., whether it is in a solid or a liquid form. What is drunk, for instance, and this holds good particularly in the case of milk, may be rich in food or solid matter, and in the food we consume there is invariably a large proportion of liquid matter. Physiologically, then, the separation of the ingesta into " food " and " drink " is unsuitable. The two material factors of life are food and air ; and food may be considered as comprising that which contributes to the growth and nutrition of the body, and, by oxida- tion, to force-production. Regarded in this comprehensive light, food embraces both solid and liquid matter ; and the primary natural division is into organic and inorganic portions that is, combinations of elements producible only through the agency of life ; and chemi- cal combinations drawn simply from the mineral kingdom and in- corporated with the others. The inorganic portion of food consists of water and various saline principles. The organic portion may be subdivided into compounds of which nitrogen forms a constituent, and compounds from which it is absent ; in other words, into nitrogenized and non-nitrogenized compounds. The non-nitrogenized alimentary principles are com- posed of the three elements carbon, oxygen, and hydrogen, vari- ously united together, whilst the nitrogenized likewise contain these three elements, but, in addition, nitrogen ; and, for the most part, sulphur, or sulphur and phosphorus as well. Liebig, regarding the nitrogenized and non-nitrogenized principles as contributing to quite distinct purposes in the animal economy, re- ferred to them as forming the basis of a physiological classification. The former he looked upon as destined for appropriation towards the growth and maintenance of the components of the body, and therefore he called them " plastic elements of nutrition." The latter he regarded as simply designed for undergoing oxidation, and, in this way, for serving as a source of heat. These he termed " ele- ments of respiration ;" but the expression, it must be said, does not properly convey what is meant, and Dr. R. Dundas Thomson sug- gested that the term " calorifiant " should be employed instead. " Calorifacient," however, is a more appropriate word, and by gen- eral consent has been adopted. It stands to reason that for the growth and repair of the various textures of the body, as these have nitrogen forming an essential ingredient of their constitution, nitrogenized compounds must be CLASSIFICATION OF FOOD. 39 supplied ; but, from what is now known, it must also be said that these compounds are likewise susceptible of application to heat pro- duction. They are truly, indeed, " histogenetic," or tissue-forming materials, but, by the separation of urea (which is known to occur in their metamorphosis in the animal system), a hydro-carbonaceous compound is left, which may be appropriated to heat production. It may be asserted, in fact, that there is sufficient to show that the ni- trogenized principles in reality subserve both purposes in the animal economy. In fat, again, we have a non-nitrogenous principle, and one belong- ing, therefore, to the calorifacient group. There is every reason, however, to believe that fat is essential to tissue-development. It seems to .be intrinsically mixed up with nitrogenized matter in the animal textures. Certainly, it may be said to be directly applied towards the formation of adipose tissue. Fat, therefore, takes rank as a nutrient no less than as a calorifacient principle. Hence, Liebig's definition is not to be accepted in a rigid sense. Although nitrogenized principles constitute true "elements of nutri- tion/' yet it neither follows nor appears likely that they are limited to this purpose. Fats are undoubtedly important calorifacient princi- ples, and cannot per se supply what is required for tissue-develop- ment; they, nevertheless, take part in the process. According to our current views, which will be discussed more fully further on, fats are also concerned, in a manner not previously suspected, in muscular force-production. Taking all these considerations into account, Liebig's classification loses the scientific force it was originally sup- posed to possess. The subdivision of the organic portion of food, however, into nitrogenized and non-nitrogenized groups is still prac- tically and physiologically convenient. Prout proposed a classification which arranged food in four groups of principles, viz.: (1) the aqueous, (2) the saccharine, (3) the olea- ginous, and (4) the albuminous. This classification, it will be seen, fails to include saline matter, which, as already said, forms an element indispensable to nutrition. The saccharine and oleaginous groups comprise non-nitrogenized principles, while the albuminous comprehends the nitrogenized. The classification that will be adopted in this treatise is one which involves no expression of physiological destination, but is based on 40 ALIMENTARY PRINCIPLES. the chemical nature of the principles. It is first assumed that food falls naturally into Organic, and Inorganic, divisions. Next, that the organic is subdivisible into- Nitrogenous, and Non-nitrogenous ; and further that the non-nitrogenous is naturally and conveniently again subdivisible into Fats and Carbohydrates: the former consisting of carbon and hydrogen in combination with only a small amount of oxygen ; the latter of carbon with oxygen and hydrogen always in such relation to each other as to be in the exact proportion to form water. To this latter group belong such principles as starch, sugar, gum, &c. It must be observed that there are a few principles which do not strictly fall within either of the preceding groups. Such, for in- stance, as alcohol, the vegetable acids, and pectin or vegetable jelly. Alcohol occupies an intermediate place between the fats and carbo- hydrates, whilst the others are even more oxidized compounds than the carbohydrates in other words, contain a larger amount of oxygen than is required for the conversion of their hydrogen into w^ater. These principles are hardly of sufficient importance, in an alimentary point of view, to call for their consideration under a dis- tinct head, and they will therefore be spoken of in connection with the carbohydrates. Having said thus much upon the classification of the alimentary principles, I shall next speak of them in relation to their respective physiological bearings, taking the groups in the following order : 1. Nitrogenous principles. 2. Hydrocarbons or Fats. 3. Carbohydrates. 4. Inorganic materials. THE NITROGENOUS ALIMENTARY PRINCIPLES. NITROGEN enters largely into the composition of the animal body. It therefore requires to be freely supplied from without. Although living in an atmosphere about four-fifths of which consist of nitro- gen, yet it is not from this source (though the question was formerly entertained) that our supply of nitrogen is drawn. Nitrogen, to be available for us, must be supplied in a state of combination. It is not, indeed, with nitrogen in the form of an element that we have anything to do in the question of alimentation, but only with com- pounds containing it, and such compounds, it may be said (as re- gards animal alimentation), that have been produced under the influence of life that is, compounds which answer to the name of " organic." Organic nitrogenous matter, then, and not nitrogen, is what we require to have supplied to us, and what alone we have to deal with physiologically. Such nitrogenous matter must, therefore, constitute an essential ingredient of our food, and we find that it there exists under various chemical forms. Chemists recognize several well-defined compounds amongst the nitrogenous matter found in different articles of food. Besides these, there may be some nitrogenous matter which is still susceptible of being used, but which has not yet been specialized, and which in an analysis would fall among the extractives. This, however, cannot be sufficient in amount to be of much significance. If we look at the nitrogenized alimentary principles which have been made known, some are characterized by yielding protein when subjected to the action of an alkali and heat, whilst from others no protein is similarly to be procured. The former comprise the albu- minous group, and are often referred to as the protein compounds ; the latter constitute the gelatinous principles. 42 ALIMENTARY PRINCIPLES. When the discovery of protein was first of all made by Mulder, the substance was regarded as forming the base or radical of the albu- minous principles. It contains the four elements carbon, hydrogen, oxygen, and nitrogen ; and each of the albuminous principles was regarded as simply resulting from the combination of the supposed base with different quantities of sulphur and phosphorus, or sulphur only. It must be stated, however, that there is nothing to show that protein really exists in the compounds from which it is to be obtained. It can be regarded only as a product of the chemical process to which it is necessary to subject the compounds to obtain it. Looked at in this light, it constitutes a chemical and not a physiological principle. It, therefore, has no direct physiological bearing, but it nevertheless serves to link together certain important physiological compounds. The albuminous or protein compounds comprise such as albumen, fibrin, and casein, with some others, which form modifications of these. Albumen may be looked upon as the most important representa- tive of the protein group. It consists of the four elements carbon, oxygen, hydrogen, and nitrogen, with the addition of some sulphur and phosphorus. As it is met with in animal productions, it is in such intimate union with fatty, alkaline, and earthy matter, that it is with some difficulty separable from them. It varies to some extent in its behavior, as it is obtained from different sources. The albumen of the blood, for instance, does not agree in all respects with the albumen of the white of egg. One of the most striking properties of albumen is its coagulability upon the application of heat. It, therefore, exists under two states, viz., soluble and coag- ulated albumen. Albumen may be regarded as the pabulum in the blood from which the different animal tissues are evolved. That it can afford per se the nitrogenous matter required for nutrition is proved by its being the principle in the egg from which are developed the nitro- genous tissues of the chick. Fibrin is characterized by its property of undergoing spontaneous coagulation. It is composed of the same elements as albumen, but contains a larger proportionate amount of sulphur, and also a rather larger quantity of oxygen. The fibrin of muscular tissue is not NITROGENOUS COMPOUNDS. 43 identical with the fibrin of blood. It will be referred to under the head of syntonin. Casein forms the protein compound of milk. It is distinguisha- ble from fibrin by not undergoing spontaneous coagulation, and from albumen by not being coagulable by heat, and by being thrown down by organic acids which do not precipitate albumen. Besides the four elements carbon, oxygen, hydrogen, and nitrogen it contains sulphur, but no phosphorus. It is remarkable for the large quantity of phosphate of lime which it is capable of holding bound up with it, and the tenacity with which it retains it. There is, it should be stated, a little uncertainty regarding the chemical constitution of casein. By some it is regarded, not as a simple, but as a compound body a body composed (in reality) of a combination of two or more others. Besides these well-known protein compounds there are modifica- tions of them which have been particularized by chemists, and the following may be referred to as connected with the subject of food. Vitellin is the name given to the modified form of albumen which exists in the yolk of the egg. There are certain points in which this substance comports itself differently with reagents from ordinary albumen. Globulin is the albuminoid matter existing in the fluid contents of the blood-corpuscle. It is there intimately associated with, but, nevertheless, quite distinct from, the coloring matter. The same principle is also found in the crystalline lens of the eye. Different opinions have been expressed regarding the true position it holds. Lecanu looked upon it as identical with albumen ; Simon with casein; while Lehmann remarks that he would be disposed to place it by the side of vitellin, if the elementary analysis were not opposed to that view. Syntonin, or muscle-fibrin, was first recognized as a distinct sub- stance from blood-fibrin by Liebig. Among the properties in which it differs from blood-fibrin one of the most striking is its ready solubility in a weak solution of hydrochloric acid. Over blood- fibrin a watery solution of hydrochloric acid has no solvent power. It causes it only to swell up. The protein compounds have as yet been referred to only as they occur in animal productions. But vegetable productions also con- tain compounds which, in the language of Liebig, are not only simi- 44 ALIMENTARY PRINCIPLES. lar to, but absolutely identical with, the albumen, fibrin, and casein of the animal kingdom. Vegetable albumen is contained in wheat and the other seeds of the cerealia. The juices of most vegetables, such as turnips, carrots, cauliflower, cabbage, &c., yield more or less precipitate with heat by virtue of its presence. It is also found in considerable abundance in association with vegetable casein in the oily seeds, such as almonds, nuts, &c. Vegetable fibrin, like albumen, is also found in the cereal seeds. It remains behind when flour is washed with a stream of water for the extraction of gluten. The albumen, starch, &c., are carried away with the water, and a tenacious mass is left, which is known as crude gluten. It is not this which constitutes vegetable fibrin, but vege- table fibrin forms a portion of it. By means of boiling alcohol the crude material obtained as above is resolved into two portions. The one which is dissolved consists of gluten and casein, whilst that which remains is vegetable fibrin. Vegetable fibrin also exists in the juice of the grape and most vegetables. Vegetable casein can be obtained from peas, beans, and other legu- minous seeds, and is sometimes specially denominated legumin. It also exists, with albumen, in the almond and suchlike oily seeds. The gelatinous principles constitute nitrogenous compounds, but do not yield protein like the compounds that have just been referred to. They comprise gelatin and chondrin, and are obtainable only from animal products : gelatin from bone and other structures containing fibrous tissue, and chondrin from cartilage. The most striking property they possess is that of their aqueous solution gelatinizing upon cooling. It is gelatin which forms the basis of soups. Besides carbon, hydrogen, oxygen, and nitrogen, as constituent elements, a small amount of sulphur appears also to be present. They contain no phosphorus. The question has been raised, and largely discussed, as to whether gelatin and chondrin exist in the tissues, or are formed in the pro- cess of obtaining them, viz., the prolonged boiling of the tissue in water. On looking at the chemical properties of gelatin, we notice that it forms an insoluble compound with tannic acid. Now, it is well known that a structure which yields gelatin, on being soaked in a solution of tannic acid, gives rise to the formation of the compound mentioned. It is this, indeed, which forms the basis of leather, a DIGESTION OF THE NITROGENOUS PRINCIPLES. 45 fact which is strongly in favor of gelatin really existing as a constit- uent of the animal body. It has been stated that the gelatinous principles which have fallen under consideration are to be obtained only from animal products. No nitrogenous compound of the kind is met with in vegetable ma- terials. The jelly yielded by fruits and some other vegetable sub- stances is quite a different article. It consists only of the three ele- ments carbon, hydrogen, and oxygen, and is known chemically as pectin and pectic acid. All the nitrogenous principles must undergo digestion before they can enter the system. Digestion, in fact, is simply a process which has for its object to fit substances for absorption into the system ; and the nitrogenous principles are in a state to resist absorption, cer- tainly to any material extent, until they have been liquefied and transformed by the agency of digestion. Beyond being mechanically comminuted or reduced to a more or less finely divided state in the mouth, our nitrogenous food under- goes no change until it reaches the stomach. In this organ it is brought into contact with a secretion, the gastric juice, which has the effect of dissolving and transforming it into a principle which pos- sesses the important property of being highly diffusible, and thereby readily transmissible from the alimentary canal into the bloodves- sels. With all the nitrogenous alimentary principles the result is the same. They each, under the influence of the gastric juice, lose their characteristic properties and become converted into the highly soluble and diffusible product referred to. Mialhe was the first to recognize this product of the digestion of the nitrogenous principles, and gave it the name of albuminose. Peptone is the name which has since been applied to it by Lehmann. Mialhe held that the substance obtained by the digestion of the pro- tein bodies was identical with that obtained from the gelatinous principle's. This would bring the latter into precisely the same po- sition with regard to nutrition as the former. Although our knowl- edge about the precise extent of capacity of gelatin as an article of nutrition cannot be looked upon as complete, yet the information before us justifies the inference that it does not possess the same capabilities as an albuminoid substance. If such be true, the prod- 46 ALIMENTARY PRINCIPLES. nets of digestion of the two cannot be completely identical, however much they may resemble each other in their general properties. It has been stated that, by the action of the stomach, the various principles composing our nitrogenous food lose their characteristic properties, and become converted into a substance which has received the designation of peptone from one, and album inose from another. Fibrin is dissolved, and is not susceptible of again solidifying. Al- bumen in a fluid form is not precipitated, as has been asserted, and then redissolved, but simply transformed. Albumen in the solid or coagulated state is dissolved, and fails to be again coagulable. Casein is first rendered solid, or curdled, and then redissolved. It is now no longer susceptible of being thrown down. Gelatin is liquefied, and cannot again be made to gelatinize. No matter from what principle a digested product or peptone has been obtained, the following are the characters which are found to belong to it. It is soluble to the highest degree in water, and it signifies nothing whether the liquid is in the acid, neutral, or alka- line state. It is not precipitable from its aqueous solution by heat. It is soluble in dilute alcohol, but absolute alcohol precipitates it. It is an uncrystallizable substance, devoid of odor and almost of taste. In a physiological point of view its most important property is the high degree of diffusibility it enjoys. It is designed for re- moval from the alimentary canal by absorption, and, by possessing the property referred to, a physically favorable disposition exists for the accomplishment of what is wanted. The nitrogenous alimentary principles, then, on reaching the stom- ach, are fitted for absorption by undergoing transformation into a highly soluble and diffusible substance. The change, we know, is wrought by the secretion of the stomach, although the precise modus operandi cannot be explained. There are two indispensable ingredi- ents of the gastric juice, viz., pepsin (a neutral nitrogenized princi- ple) and an acid. Pepsin is a secretory product, peculiar to, and therefore obtainable only from, the stomach. About the acid there is nothing peculiar, and different views have been held regarding the kind of acid that is naturally present. With the combination of pepsin and acid a liquid is obtained which dissolves nitrogenous matter in the same manner out of as within the stomach. Accord- ing to Lehmann, it is only hydrochloric and lactic acids and these, the same authority affirms, give the acidity to the natural secretion LIBRARY COLLEGE OF AGRICULTURE Btrkley. OF THE NITROGENOUS PRINCIPLES. 47 which yield an energetic digestive fluid with pepsin ; but, accord- ing to my own experiments on artificial digestion, other acids, such as the phosphoric, sulphuric, citric, and so on, will equally answer the purpose. From the above statements it follows that the solution of nitro- genous food in the stomach is effected by the action of a liquid which owes its virtue to the presence of a couple of principles pepsin and an acid. The action of this liquid is favored by the elevated temperature belonging to the body, and also by the move- ment to which the contents of the stomach are subjected by the ac- tion of the muscular fibres with which the walls of the organ are provided. As it is reduced to a fluid state the food is forced on into the upper bowel. Chyme is what this product of gastric digestion is called. Besides nitrogenous matter in a dissolved state, it con- tains a portion suspended in a finely divided form which has not yet undergone solution, and likewise, in the same state, those con- stituents of the food which resist the solvent action of the stomach. The nitrogenous matter which has escaped from the stomach in an undissolved state is submitted to a further digestion in the intes- tine. This may be shown by direct experimental observation. And it is not by a continued action of the gastric juice which passes on with the food in its course, but by an action exerted by the secre- tions poured into the intestine itself. It has been stated that the pres- ence of an acid forms an indispensable factor in gastric digestion. The chyme as it passes on from the stomach is strongly acid. It con- tains nitrogenous matter which has not yet undergone solution, and also gastric juice whose power (it may be inferred) has not become exhausted. So far, we have conditions which suffice for a continu- ance of the process carried on in the stomach. It happens, however, that on reaching the small intestine the chyme encounters alkaline secretions. The pancreatic juice is, to a marked extent, alkaline, and so is also the intestinal juice. The bile likewise contains a quantity of alkali in feeble combination, and easily taken by the gastric juice acid. Thus it happens that the chyme becomes more or less neutralized as the small intestine is being traversed. As the result of observation, in fact, I have noticed that by the time the lower part of the ileum is reached the intestinal contents may be found to present a neutral or even alkaline reaction. In this way, through contact with the secretions poured into the intestine, the 48 ALIMENTARY PRINCIPLES. energy of the unexhausted gastric juice contained in the chyme is destroyed, and whatever solution of nitrogenous food now occurs must be due to another agency. Let us, therefore, inquire into the effect which the various secre- tions, as they become incorporated with the chyme, are capable of producing. First, as regards the intestinal juice. This fluid, it is evident, possesses some solvent influence upon nitrogenous matter. Bidder and Schmidt ascertained by experiment that meat and coagulated albumen contained in a muslin bag undergo, on being placed in the empty small intestine, in which the bile and pancreatic juice are prevented by a ligature from descending, in from four to six hours' time, a considerable amount of digestion. In an experiment per- formed by myself, in which the hind legs of a frog that had been separated from the body, were introduced into the empty small in- testine, secured by a ligature from the descent of secretions from above, I found, after the lapse of six hours, the legs partially di- gested a portion of the skin, for example, having been dissolved away, the muscles underneath it separated, and some of the bones, to a slight extent, exposed. Next, as regards the pancreatic juice. Besides its other offices in the animal economy, this liquid acts upon and dissolves nitrogenous matters, as appears from the following considerations. In 1836, Purkinje and Pappenheim asserted that the pancreas contained a principle capable of exerting a digestive action upon the nitrogenized elements of food. This statement attracted little attention, and soon dropped out of notice. More recently Lucien Corvisart, of Paris, having reopened the subject, proved, by a series of experiments, that the pancreas, as one of its functions, supple- ments the action of the stomach, and, after a copious meal, contrib- utes to digest those nitrogenous matters which have escaped the stomachic digestion. As far as the result is concerned, the two kinds of digestion, he states, coincide, each leading to the produc- tion of albuminose. While acidity, however, is a necessary condi- tion to digestion in the case of the gastric juice, the pancreatic secre- tion, it is affirmed, possesses the power of acting equally well, what- ever the existing reaction whether acid, neutral, or alkaline. In support of his doctrine, Corvisart has adduced three sets of experimental results. ACTION OF PANCREATIC JUICE. 49 First. If the pancreas of an animal be taken when its active principle is at its maximum of quantity and quality, that is, from the fourth to the seventh hour after digestion has begun, and it be tln-n finely cut up and infused for an hour in twice its volume of water at a temperature of 20 Cent. (68 Fahr.), and the infusion be at once experimented with, it will be found, he asserts, to possess a power of dissolving the nitrogenized alimentary principles, and converting them into albuminose; and this, with no evidence of pu- trefaction being perceptible, provided the experiment be stopped at the end of four or five hours, in which time, under a temperature of about 100 Fahr., the pancreatic principle will have effected all that it is capable of doing. Secondly. The pancreatic juice obtained during life from the duct of the gland is found to be capable of acting, he affirms, as a power- ful solvent on the nitrogenized alimentary principles, when the requisite precautions are taken in conducting the experiment: the juice, that is to say, must be obtained from the fourth to the seventh hour after the ingestion of food, at which time it is charged to its maximum degree with the pancreatic principle; and must also be experimented with immediately after its collection. It dissolves, Corvisart says, fibrin more quickly and more largely than albumen. The, heat being maintained between 42 and 45 Cent. (108 and 113 Fahr.), a specimen of pancreatic juice of ordinary energy dis- solves, it is stated, if the mixture be agitated every quarter of an hour, all that it is capable of taking up of fibrin in two or three hours at the most, and of solid albumen in four or five hours; the experiment, up to this time, being attended with no evidence of ordinary decomposition, while, at a subsequent period, ordinary de- composition is found to set in. Thirdly. Azotized substances introduced into the duodenum when pancreatic juice is flowing into it are found to be dissolved, notwith- standing the gastric juice and bile are precluded from entering by applying a ligature to the pylorus and bile-duct. It is necessary to state that the evidence derivable from the last experiments must not be taken for more than it is really worth,, viewed in relation to pancreatic juice per se. The bile and the gastric juice may, it is true, have been prevented entering the duo- denum, and thereby precluded from contributing to the effect, but it 4 50 ALIMENTARY PRINCIPLES. is impossible to exclude from operation the secretions of Brunner's and the other glands of the duodenum. My own experiments with the pancreatic juice at first inclined me to think that the effects producible on nitrogenous matter through the agency of the pancreas were rather like those which result from putrefaction than from true digestion. On re-performing the experiments, however, I obtained results which certainly appeared to indicate that some digestive action had been at work. For example, upon operating with the pancreatic infusion, taken conformably with the instructions of Corvisart, I found that frogs 7 hind legs (which, according to my experience, con- stitute one of the most, if not the most, sensitive and distinct tests of digestive action) were, upon some occasions, softened, so that the flesh broke down under very slight pressure, without any evidence of ordinary putrefaction being apparent. The effect, however, was not to be compared with what is observed with artificial gastric juice, and ordinary decomposition is quickly prone to occur, which is not the case in experiments with gastric juice. Whatever the power actually enjoyed by the pancreatic juice in this direction, the chief point of interest to us as regards the subject of food is not whether this or that secretion, poured into the intes- tine, will dispose of nitrogenous matter, but whether nitrogenous matter really undergoes digestion in the intestine; and, thus framed, it will be presently seen that the question admits of being answered in a very positive manner. The bile forms another secretion, which becomes incorporated with the alimentary matter after its exit from the stomach. There is nothing, however, to show that this fluid possesses any solvent power over the nitrogenized principles of food. Remarks have been made upon the action of the secretions taken individually, but as regards the subject of food, the point of great- est interest to us, as has been already said, is what occurs within the intestine when all the secretions are allowed to enter. Experiment shows that there is a very powerful solvent action exerted, and, as I can state from personal investigation, a few hours suffice for nitro- genous matter, introduced directly into the upper part of the small intestine, to be completely digested. With reference, therefore, to the digestion of nitrogenous matter, the intestine may undoubtedly .be regarded as performing a part supplementary to that of the PRODUCTION OF ALBUMINOSE. 51 stomach. Besides its other functions, it serves to complete the di- gestion of whatever nitrogenous alimentary matter may have escaped the digestive action of the stomach, and it may be remarked that the same result namely, the production of albuminose or peptone occurs as when the solution has been effected in the stomach. Reviewing the stages that are passed through preliminary to the appropriation of nitrogenous matter within the system, we have seen that, through the agency of the stomach and of the intestine, it un- dergoes conversion into a principle which, from its diffusible nature, is readily susceptible of absorption, and it is in this form, viz., as albuminose, that the various nitrogenous alimentary principles reach the circulation. The conversion, of the nitrogenous alimentary matters into albu- minose is necessary, it has further to be remarked, not only as a process preparatory to absorption, but also as fitting them for subse- quent application to their proper destination. It cannot absolutely be affirmed that no absorption whatever occurs without previous conversion into albuminose; but this much is certain, that the amount so absorbed must be very trifling, and it can be shown that if they directly reach the circulation in any quantity they visibly pass off without being applied to the purposes of the economy. Bernard was the first to demonstrate that the albumen of egg, reaching the circulation without having previously undergone diges- tion, quickly passes from the system into the urine. If introduced directly into one of the bloodvessels, or even if injected into the sub- cutaneous tissue, it rapidly betrays its presence in the urine. This I can attest from my own experience. Both after injection into a vein and into the subcutaneous tissue the albumen of egg, as I have often seen, is soon recognizable in the urine. It has also been observed that a meal consisting largely of eggs, particularly if taken after prolonged fasting, has been followed by the appearance of albumen in the urine. Here, apparently, it has happened that some albumen has reached the circulation without having undergone the usual conversion ; and, as when experimentally injected, has been thence discharged with the urine. Hence it may be concluded, not only that egg-albumen and blood-albumen differ strikingly from each other in a physiological point of view, but that egg-albumen, as such, is not fitted for entering the circulation. The conversion of 'albumen into albuminose, therefore, not onlv 52 ALIMENTARY PRINCIPLES. bears on the facility of absorption, but on the adaptability for subse- quent application in the system. The process of metamorphosis, in fact, is required, not only with a view to adaptability for absorption, but to subsequent fitness for utilization in the system. Casein and gelatin I have found 1 comport themselves in the same manner as albumen, namely, pass off from the system with the urine when directly introduced into the circulation. The injection of three ounces of milk into a vein was observed in an experiment to be followed by the appearance of casein in the urine. The injec- tion of 100 grains of isinglass, dissolved in two and a half ounces of water, also so charged the urine with gelatin as to give rise to the formation of a firm, solid jelly on cooling. Thrown oif as they thus are from the system, albumen, casein, and gelatin are evidently not adapted for direct introduction into the circulation. Fibrin, on account of its solidity, cannot be similarly experimented with. Digestion, in its case also, is an indispensable condition to its introduction into the circulation. In respect, in- deed, of all these principles, it may be said that their metamorphosis in the digestive system is needed as a preliminary step to their capa- bility of appropriation in the body, and their application to the pur- poses of life. We have followed the nitrogenous alimentary principles to the stage of albuminose. The precise nature of what next ensues is not yet known. There can be little or no doubt as to the progress from albuminose to the albumen of the blood, but as to what next occurs we have no data to show. With the ultimate products that are formed we are acquainted, but the steps of metamorphosis are as yet beyond our knowledge. The chain we have hitherto followed now wants one or more links, which we have as yet no means of dis- covering. As regards the seat of metamorphosis we have also no information of a precise nature to deal with, but we may, neverthe- less, hazard the surmise that the liver is the viscus in which albu- minose, like other nutritive matters absorbed from the alimentary canal, mainly, if not entirely, undergoes metamorphosis. The va- rious nitrogenous principles of the body must be primarily derived 1 " Gulstonian Lectures (1862) on Assimilation and the Influence of its Defects on the Urine," Lancet, vol. i,' p. 574, 1863. USES OF NITROGENOUS MATTER. 58 from it ; but, whether by direct transformation into them, or by- passing through the stage of albumen, we have not the means of deciding. That albumen is susceptible of metamorphosis, however, into the other principles, we know, from its forming in the egg the pabulum whence the various nitrogenous principles of the young bird take their origin. Instead of wandering farther into the domain of conjecture as to the subject of metamorphosis, let us now turn our attention to the purposes fulfilled by the nitrogenous principles as alimentary matter. First in importance is the supply of material for the development primarily, and for the renovation secondarily, of the tissues. Wherever vital operations are going on, there nitrogenous matter is present, forming, so to speak, the spring of vital action. Although non-nitrogenous matter contributes in certain ways towards the maintenance of life, yet it is nitrogenous matter which starts and keeps in motion the molecular changes which result in the phe- nomena of life. Nitrogenous matter, it may be said, forms the basis, without which no life manifests itself. Life is coincident with molecular change. In non-nitrogenous matter the elements of the molecule are not, of themselves, prone to change ; whereas in the molecule of nitrogenous matter there exist a greater complexity of grouping among the elements, and these cohere so loosely, or are so feebly combined, as to have a constant tendency to alter or to re- group themselves into simpler combinations. By this change in the nitrogenous, change is induced in the contiguous non-nitrogenous molecule, and, occurring as the whole does in a definite or prescribed order, the phenomena of life are produced. Nitrogenous matter, in this way forming the instrument of living action, is incessantly being disintegrated. Becoming thereby effete and useless, a fresh supply is needed to replace that which has fulfilled its office. The primary object of nitrogenous alimentary matter may thereupon be said to be the development and renovation of the living tissues. We have seen that nitrogenous matter forms an essential part of living structures. It holds the same position in the case of the ftecretions. These owe the active properties with which they are endowed, chiefly, if not entirely, to a nitrogenous constituent. This is drawn from the blood by the glands just as it is drawn by the 51 ALIMENTARY PRINCIPLES. tissues ; and on passing from the blood it is modified or converted, by the agency of the gland, into the special principle encountered. Nitrogenous matter is thus as essential to the constitution of the active secretions as it is to the tissues ; and, as the amount of the secretions required is in relation to the general vital activity, a cor- responding demand for nitrogenous matter is created. I now come to treat of nitrogenous matter in relation to force- production. The dependence of muscular and nervous action upon oxidation of the respective tissues is one of the many doctrines which have emanated from the inventive intellect of Liebig. According to the view propounded, nitrogenous matter alone constitutes the source of muscular and nervous power. The tissues being consumed in the exercise of their functional activity or the manifestation of their dynamic properties, fresh nitrogenous matter is alleged to be needed to replace that which has served for the production of power. Thus viewed, nitrogenous matter has been regarded as not only applied to nutrition and to the formation of the nitrogenous constituents of the active secretions, but also to the restitution of the loss incurred by the production of power. What wonder, then, if with all these purposes to fulfil, the nutritive value of food should have been measured, as it latterly has been, by the amount of nitrogenous matter it contains ? Liebig's doctrine was at once accepted, and until recently has been looked upon as expressing a scientific truth. Like many other of its author's views, its plausibility was such that no one ventured to question its soundness. Gradually, however, experimental in- quiry began to invalidate it, and the reactionary move has advanced till Traube has been led to express himself in directly opposite terms regarding the source of muscular and nervous power. According to this authority, for instance, the organized or nitrogenous part of a muscle is not destroyed or consumed in its action. The resulting force is affirmed to be due, instead, to the oxidation of non-nitroge- nous matter the muscle merely serving as a medium for the conver- sion of the generated force into motor power. The point has at- tracted much attention of late, and researches of an elaborate nature have been conducted with regard to it. Let us see the position in which these researches have placed it. NITROGENOUS MATTER AND MUSCULAR ACTION. 55 The argument representing the question to be solved may be thus expressed : Does the force evolved by muscular action proceed from destruction of muscular tissue? If so, nitrogenous matter would be needed to replace the loss incurred, and the result would be equiva- lent to nitrogenous matter through the medium of muscle being applied to the production of motor power. Now, if muscular action is coincident with the destruction of muscular tissue, there must, as a product of the destruction, be a nitrogen-containing principle eliminated. The elements of the compounds that have served their purpose in the economy do not accumulate, but are discharged from the system under certain known forms of combination. The nitro- gen, therefore, belonging to a consumed nitrogenous structure should be recognizable in the effete matters thrown off from the body. Nay, more : as the force developed by muscular action cannot arise spontaneously as it can be produced only by transmutation from another force the destruction of muscular tissue (which through the chemical action involved supplies the force), should be in pro- portion to the amount of muscular work performed, and the nitrogen contained in the excreta in proportion also to the amount of muscu- lar tissue destroyed. Now, in proceeding to measure the extent of tissue metamorphosis by, the nitrogen eliminated, it is necessary, in the first instance, to be sure of our data regarding the channels through which nitrogen finds its exit from the body it is necessary, that is to say, to ascer- tain whether nitrogen escapes with the breath and perspiration, as was at one time asserted, as well as by the alimentary canal and the kidneys. We have no accessible means, it must be stated, of deter- mining in a direct way whether nitrogen passes off by the lungs and skin. Our conclusions have to be based upon comparing the nitro- gen ingested with that encountered in the urine and alvine evacu- ations. Formerly, it was said that a deficiency in the latter existed, and it was put down to loss by pulmonary and cutaneous elimina- tion. Barral, for instance, only detected half the nitrogen of the food in the urine and faeces, and thence inferred that the remainder was discharged with the breath and perspiration. In opposition to this, however, several trustworthy observers (amongst whom may be named Voit, Ranke, Haughton, and Parkes), aided by the im- proved methods of analysis introduced by modern experience, have recovered within a very close approach all the nitrogen of the food 56 ALIMENTARY PRINCIPLES. from the urinary and intestinal excreta. Dr. Parkes's observations are especially worthy of reliance, and he confidently asserts that it may be looked upon as established, that an amount of nitrogen is discharged by the kidney and intestine equivalent to that which enters with the food. Admitting this to be the case, we have only to look to the products that escape from these two channels for the information that is wanted about the discharged nitrogen in relation to the question before us. Next comes the determination of the relation respectively held by the urinary and intestinal nitrogen to the point under consideration. It has long been known that the chief portion of the escaping nitrogen is to be met with in the urine. Lehmann, for instance, found, whilst subsisting on a purely animal diet (eggs), that a daily average of 30.3 grammes (467 grains) of nitrogen entered his system, and that a daily average of 24.4 grammes (376 grains) was dis- charged by the urine. Here, therefore, it was ascertained that an amount equal to five-sixths of the ingested nitrogen escaped by the kidneys. But more recent and precise evidence has been afforded by a series of very carefully conducted observations made upon two sol- diers by Dr. Parkes. 1 The observations extended over sixteen consecutive days, and the results not only bear on the ingestion and egestion of nitrogen generally, but likewise show that the great bulk of outgoing nitrogen is to be met with in the urine. The men were both of almost precisely the same weight at the end of the time as at the beginning, so that the ingoing and the outgoing matter must have been closely balanced. They were subjected to varying con- ditions of rest and exercise, but consumed exactly the same allow- ance of food every day. The nitrogen in the food taken during the sixteen days amounted to 313.76 grammes; and from the urine of one of the men (distinguished as S.) there were recovered 303.660 grammes, and from that of the other (distinguished as B.) 307.257 grammes. Thus the amount of nitrogen discharged from the kid- neys was, in the case of S., only about ten grammes, and in that of B., six grammes less than that admitted with the food. The alvine evacuations were collected and analyzed only upon three occasions. Taking the mean of the results then obtained as representing the 1 Proceedings of the Royal Society, June 20th, 1867. MUSCULAR ACTION AND ELIMINATION OF NITROGEN. 57 daily average, and calculating from this for the sixteen days, the quantity of nitrogen discharged from the bowels amounted in S. to 25.8 grammes, and in B. to 17.2 grammes, thus somewhat exceeding the difference between the ingested nitrogen and that excreted in the urine, or giving, in other words, rather more nitrogen discharged than nitrogen ingested. The nitrogen discharged from the bowels may be said to have been found to form, upon an average, from about one-eighth to one- twelfth or one-thirteenth of the total nitrogen voided. Owing its origin, as it does, to the nitrogen belonging to the undigested food on the one hand, and that contained in the unabsorbed intestinal se- cretions on the other, it is constantly liable to incidental variation. There is this, also, to be remarked, that the nature of its source ex- cludes it from possessing any relation to the question under consid- eration. We have, therefore, only the urinary excretion to look to as forming the channel through which the exit of nitrogen, resulting from the metamorphosis of nitrogenous matter in the system, takes place ; and observation has shown that in the human subject it .is mainly under the shape of urea that the escape occurs. What, now, is the state of the urine in relation to rest and exer- cise? If muscular disintegration forms the source of muscular work, the quantity of urinary nitrogen ought to increase in propor- tion to the amount of muscular work performed. Lrhmann, imbued with Liebig's views, as his writings show, speaks of there being an actual increase in the elimination of urea in proportion to muscular exercise, and yet he gives it as the result of observation upon himself that, while under ordinary circum- stances he passed about 32 grammes (493 grains) of urea in the twenty-four hours, the quantity passed after severe bodily exercise was upon one occasion 36 grammes (555 grains), and upon another 37.4 grammes (577 grains) only this insignificant disparity to cor- respond with the difference in the amount of muscular work per- formed. Voit experimented upon a dog, and determined the amount of urea voided during rest and the performance of mechanical work, in association with abstinence and a regulated diet of meat. The work imposed upon the dog was running in a treadmill. The re- sults, both during abstinence and feeding, exhibited no material ex- cess in the urea voided during work over that voided during rest. 58 ALIMENTARY PRINCIPLES. Dr. E. Smith, also, in his observations on the elimination of car- bonic acid and urea during rest and exercise, found, in the case of the prisoners at Cold bath Fields, that, in the absence of food, the labor of the tread wheel did not, to any material extent, increase the nitrogen discharged under the form of urea. Like others have done, he noticed a distinct relation between the urea discharged and the food ingested. At the same time he regarded and this was several years ago, when our knowledge stood in a very different po- sition from what it does now the relation between the urea and muscular work as far less established then than it had been held to be for some time before. The theory that muscular work is dependent on, and proportioned to, the destruction of muscular tissue by oxidation, received its de- cisive blow from the now celebrated observations of Drs. Fick and Wislicenus, professors of physiology and chemistry respectively at Zurich. 1 These experimentalists subjected themselves to a measur- able amount of work by ascending a mountain of an ascertained height. They argued that if the work performed be due to destruc- tion of muscular tissue seeing that the nitrogenous product of de- struction is discharged in great part, if not entirely, with the urine the collection of the urine, and the determination of its nitroge- nous contents ought to show the amount of nitrogenous matter de- stroyed. Again, as the mechanical work to be performed must be represented by an equivalent of chemical action to produce it, the destruction of nitrogenous matter, as measured by the nitrogen ap- pearing in the urine, ought to accord with the amount of work per- formed. To simplify the experiment, the food consumed by the ex- perimentalists consisted solely of non-nitrogenous matter; so that the nitrogen appearing in the urine might be derived exclusively from that belonging to the system. Drs. Fick and Wislicenus chose for ascent the Faulhorn, near the lake of Brienz, in the Bernese Oberland, a steep mountain of about 2000 metres (6561 feet) above the level of the lake, and furnished with hotel accommodation on the summit, enabling them to rest overnight and make the descent next day. On the 30th of August, between 10 minutes past 5 in the morn- ing and 20 minutes past 1 in the afternoon, the ascent was made. 1 "On the Origin of Muscular Power," by Drs. Fick and Wislicenus,- Philo- sophical Magazine (Supplement), vol. xxxi, 1866. FICK AND WISLICENUS 'S EXPERIMENTS. 59 From the noon of the 29th no nitrogenous food had been eaten by the experimenters, their diet consisting solely of starch and fat (taken in the form of small cakes), and sugar as solid matter; and tea, beer, and wine as drink. After ascending the mountain, Drs. Fick and Wislicenus rested, and took no other kind of food till 7 in the evening, when they partook of a plentiful repast of meat and its usual accompaniments. They began to collect their urine for examination from 6 P.M. of the 29th that is, six hours after the commencement of their non- nitrogenous diet. The urine secreted from this time till ten minutes past 5 A.M. of the 30th, when the ascent began, was called the " be- fore-work " urine. The urine secreted during the ascent was called the "work" urine; and that from 1.20P.M. to 7 P.M. (from the completion of the ascent to the cessation of the non-nitrogenous diet), the "after-work" urine. Finally, the urine secreted during the night spent on the Faulhorn up to half-past 5 A.M. was also col- lected, and denominated "night" urine. Each specimen was measured, and both the quantity of urea and the absolute amount of nitrogen contained in it determined. For the object before us it will suffice to confine our attention to the ni- trogen ; and the quantity of this element secreted per hour (calcu- lated from the amount contained in the respective specimens and the time passed in secretion), stood thus, for the several periods : Quantity of Nitrogen excreted per hour. Fick. Wislicenus. Grammes. Grammes. Before work, 0.03 0.61 During work, 0.41 39 After work, 0.40 040 Night, 0.45 0.51 A glance at these figures shows the agreement that existed in the two cases. The result proved that, whilst the nitrogenous excretion was related to the food ingested, it was not so to muscular action. Less nitrogen, it is noticeable, was voided during the "work" and " after-work " than during the " before-work " period, and this was plainly attributable to the absence of nitrogenous food from the diet. During the night, after the meal of mixed food, there was an increase, greater in Wislicenus's than in Fick's case ; but the one 60 ALIMENTARY PRINCIPLES. meal did not bring the amount of nitrogen up to the point at which it stood shortly after the commencement of their abstinence from nitrogenous food. The conclusion, then, that in the first place may be drawn from this experiment is that muscular work is not accompanied by the increased elimination of nitrogen that might be looked for if it re- sulted from the oxidation of muscle. But let us inquire whether the disintegation of nitrogenous matter which actually occurred dur- ing the " work " and " after-work " periods, as measured by the nitrogen excreted, would account for the generation of an amount of force equivalent to that expended in the work performed. Knowing that the nitrogenous matter of muscle contains say, in round numbers 15 per cent, of nitrogen, it is easy to calculate to how much muscular tissue the excreted nitrogen was equivalent; and taking the muscular tissue thus represented, an approximate, if not an absolute, estimate can be given of the amount of mechanical work which its oxidation would be capable of performing. The height of the ascended mountain, likewise, being known, the amount of muscular force actually employed in raising the weight of the body to the summit can also be definitely expressed. We have, therefore, these data supplied : 1st. From the nitrogen excreted the amount of nitrogenous mat- ter oxidized : 2d. The amount of force that this oxidation would generate; and 3d. The expenditure of force required to raise the bodies of the experimenters to the height they reached. Now, if the work performed were due to the oxidation of muscle, the second factor ought to equal the third ; that is, the force produ- cible from the muscle oxidized ought to be equivalent to the force that was expended. The results of the calculation, however, show, as will be presently seen, that the force expended considerably ex- ceeded the amount derivable from the nitrogenous matter consumed. Nor is this all. Besides the force expended in simply raising the body-weights of the two men to the elevation reached, there would also be occurring, during the performance of the work, an expendi- ture of muscular power in keeping up the circulation, in respiratory action, and the other life-processes. The calculations on these points have been carefully worked out by Fick and Wislicenus ; and though MUSCULAR WORK AND MUSCLE OXIDATION. 61 the data for the process are scarcely precise enough to warrant our regarding the results as scientifically exact, still they may be ad- mitted as affording a basis for a safe general conclusion to be drawn. We are also told that wherever a doubt existed about the data, figures were taken as favorable as was allowable to the old hypoth- esis, which referred the source of power to muscular oxidation. In giving the conclusion furnished, it is not necessary to intro- duce the details of the calculation. It will suffice to say that sum- marily stated the result of the calculation showed that the measured work performed during the ascent exceeded by about one-half in Fick's case, and more than three- fourths in that of Wislicenus, the amount which it would be theoretically possible to realize from the amount of nitrogenous matter consumed. It has been shown by Professor Frankland 1 that the results of Fick and Wislicenus in reality afford stronger evidence than they have contended for. Fick and Wislicenus were obliged to estimate the force-value of the nitrogenous matter, shown by the nitrogen in the urine to have been destroyed in the system, from the amount of force known to be producible by the oxidation of its elements, be- cause the actual determination for the compound itself had not been made. Professor Frankland, however, has since experimentally as- certained with the calorimeter the amount of energy or force under the form of heat evolved during the oxidation of a given quantity of nitrogenous matter as the oxidation occurs within the living system, where, it must be remembered, on account of the nitrogen carrying off the elements that are associated with it in urea, some carbon and hydrogen escape being consumed. Frankland's results give as the actual amount of energy producible from the nitrogenous matter consumed in the bodies of the experimentalists about half the quan- tity they had reckoned in their calculations. Thus, the results tell so much the more in Fick and Wislicenus's favor. Frankland con- siders, taking all points into consideration, that scarcely a fifth of the actual energy required for the accomplishment of the work per- formed in the ascent of the mountain could have been obtained from the amount of muscle (nitrogenous matter) that was consumed. As- suming, therefore, the foregoing conclusions to be entitled to credence, 1 " On the Origin of Muscular Power," Lond. Philos. Magazine, vol. xxxii, 1866. 62 ALIMENTARY PRINCIPLES. the doctrine which ascribes muscular action to oxidation of muscular tissue becomes utterly untenable. Dr. Parkes has conducted, in a most careful manner, a series of investigations on the influence of rest and exercise, under different diets, upon the effete products of the system, and, more particularly, to test the accuracy of the results arrived at by Fick and Wislicenus. He says, " Although these results [Fick and Wislicenus's] are sup- ported by the previous experiments of Dr. Speck, who has shown that, if the ingress of nitrogen be restricted, bodily exercise causes no or a very slight increase in the elimination of nitrogen by the urine, it appeared desirable to carefully repeat the experiments, not only because the question is one of great importance, but because objections might be, and, indeed, have been, reasonably made to the experi- ments of Professors Fick and Wislicenus, on the ground that no sufficient basis of comparison between periods of rest and exercise was given, that the periods were altogether too short, and that no attention was paid to the possible exit of nitrogen by the intestines." Dr. Parkes's experiments were conducted upon perfectly healthy soldiers, men who, when steady and trustworthy, as were the sol- diers made use of, form, as Dr. Parkes observes, highly suitable subjects for experiments of the kind, their regularity in diet and oc- cupation, and their habits of obedience, affording a special guarantee for the precision with which they will carry out the instructions given. There can, indeed, be little or no doubt, from the harmony observable all through, that the results furnish as exact and reliable information as can be hoped to be obtained. The total nitrogen contained in the urine was determined, as well as the urea; and by -this step more conclusive evidence is supplied than by the simple determination of urea, as had only been done in the experiments of Fick and Wislicenus and others: obviously so, because it might be said that nitrogen escaped (as is really to some extent the case) in other forms than that of urea. The experiments consisted of two series, and extended, in each case, over several successive days. In the first series' a comparison is instituted of the products of excretion during rest and exercise under a non-nitrogenous diet. In the second 2 the same comparison 1 Proceedings of the "Royal Society, Jan. 1867, vol. xv, No. 89. 2 Id. vol. xvi, No. 94. DR. PARKES'S EXPERIMENTS ON ELIMINATION OF NITROGEN. 63 is made under a fixed diet, containing an ordinary admixture of ni- trogenous and non-nitrogenous food. In drawing conclusions regarding the destruction of muscle from the nitrogen eliminated, it is, of course, of the first importance that that the whole of the voided nitrogen should be presented to our notice. Dr. Parkes is convinced, from his experiments, that no nitrogen escapes either by the breath or perspiration, but that it is all to be found in the excreta from the kidneys and bowels. The nitrogen discharged by the bowels forms a comparatively small and varying proportion, and being derived from the undigested food and the unabsorbed digestive secretions, has no bearing in reference to the point before us. There remains, therefore, only the urinary nitrogen to consider as a measure of the tissue-metamorphosis occur- ring in the system. Thus prefaced, let us now see what light is thrown upon the matter under consideration by Dr. Parkes's experi- ments. For the sake of simplicity notice will only be taken of the total urinary nitrogen voided, as this gives in a more reliable man- ner than the urea the information that is wanted. The men forming the subjects of the first series of experiments are distinguished as S.'and T. T. was a much smaller man than S. (S. weighing 150 and T. 112 Ibs.), and it will be observed that he, throughout, passed a less amount of urinary nitrogen. He did not consume quite so much food; and as it was found that he discharged rather more nitrogen from the intestine, it may be assumed that he did not so fully digest and absorb what he ingested. For six days the men were kept upon an ordinary mixed diet, and pursued their customary occupation. The urine was collected and examined during four out of the six days, and the following is the mean amount of the total nitrogen passed per diem : Mean urinary nitrogen per diem. Mixed diet, with customary occupation, . During the following two days the diet was restricted to non- nitrogenous food, consisting of arrowroot, sugar, and butter. The only nitrogen ingested and this may be regarded as too insignifi- cant to require being taken into account was in the tea the men were allowed to drink, it being thought desirable not to deprive them of this beverage. Throughout the two days they remained as 64 ALIMENTARY PRINCIPLES. much at rest as was practicable ; they were allowed to get up, but not to leave the room. Mean urinary nitrogen per diem. Non-nitrogenous diet, with rest, . . - { ' J' 176 ra mes ' The men were now put back, for four days, upon a mixed diet, with customary occupation, just as at the beginning of the experi- ment. Mean urinary nitrogen per diem. Mixed diet with customary occupation, { ^ 12.988 grammes. ( I. 11.095 u Next, they were restricted again for two days to the same non- nitrogenous food as before, but this time it was accompanied with active walking exercise. During the first day the distance walked was 23} , and during the second 32} miles. The diet, it is stated, satisfied hunger, and there was no sinking or craving for other kinds of food. Mean urinary nitrogen per diem. Non-nitrogenous diet, with active exer- f S. 8.971 grammes. ei*e, ........ \ T. 8.034 To complete the experiment four more days were passed under observation with the ordinary mixed diet, accompanied by ordinary exercise. Rather more nitrogenous food was taken during these four days succeeding the two days' active exercise than during the four days succeeding the two days 7 rest, the men feeling more hun- gry after the " work " period than after the period of " rest." The mean for T., it is mentioned, is for three days instead of four, one analysis having failed. Mean urinary nitrogen per diem. Mixed diet with customary occupation, . f ^ J^l grammes. From this series of results we find that there was no material variatioh in the amount of urinary nitrogen discharged during the two days when a distance of 56J miles was walked, as compared with the two days spent in as complete. a state of rest as possible, on both occasions restriction to non-nitrogenous food being enjoined. Comparing both these periods, however, with those in which nitro- DR. PARKES'S EXPERIMENTS ON ELIMINATION OF NITROGEN. 65 genous food was taken, we recognize a marked exemplification of the well-established fact that diet, on the other hand, exerts a striking influence over the amount of nitrogen eliminated with the urine. During each of the non-nitrogenous diet-periods the quantity of nitrogen eliminated was considerably less than during the others ; it is also noticeable that the influence of the non-nitrogenous food was extended into the subsequent ordinary diet-periods, less nitrogen being voided during these than at the commencement of the experi- ment, before any restriction from nitrogenous food had been imposed. This point, however, will be further alluded to hereafter. In the second series of experiments the amount of nitrogen elimi- nated was determined under the conditions of rest and exercise, com- bined with a mixed diet. One of the two men, S., was the same who had been made use of in the former experiment; the other, B., was a fresh man, weighing 140 Ibs., and therefore nearer in size to S., who weighed 150 Ibs., than T., of the former experiment, who weighed 112 Ibs. During the sixteen days over which the observations ex- tended each man took precisely the same allowance of food in the twenty-four hours the. food consisting of weighed quantities of meat, bread, potatoes, and the other constituents of an ordinary mixed diet. For the first four days the men pursued their custom- ary employment. The next two days were passed in rest. Then followed four days of ordinary employment ; after this, two days of active exercise ; and finally, four days again of ordinary employment. The amount of nitrogen eliminated by the kidneys during the several periods is shown in the following table : Ordinary employment (mean of four days), . Rest (mean of two days), Ordinary employment (mean of four days), . Active exercise walking on level ground, 24 miles the first day, and 35 the second (mean of two days), Ordinary employment (mean of four days), . Urinary nitrogen per diem. S. 17.857 grammes. B. 18.502 S. 19.137 B. 19.471 S. 17.612 B. 18485 S. 19.646 B. 19.959 S. 21.054 B. 20.092 In these results it will be seen there is nothing to sanction the doctrine that the source of muscular power resides in the destruction 6 66 ALIMENTARY PRINCIPLES. of muscular tissue. In two persons subsisting on an identical and unvarying daily diet, and subjected to varying conditions of muscu- lar exertion, we find nearly the same quantity of nitrogen elimina- ted during two days' hard walking as during two days of rest. It is curious, and also, it must be owned, does not appear explicable, that during the periods of both rest and active exercise the daily amount of nitrogen eliminated was in excess of that eliminated during the first two periods of ordinary employment, the figures at the same time for the associated periods respectively agreeing very closely with each other. In the third period of ordinary employ- ment that is, after the two days of walking exercise the nitrogen voided was greater in quantity than at any other time. Such excess, however, did not amount to anything particularly marked. Comparing in detail the nitrogen eliminated during the correspond- ing portions of the two-day periods those of rest and active exer- cise Dr. Parkes observes, with respect to the results furnished, "On the first day of exercise, the nitrogen in each man fell below the cor- responding day of rest by 1.626 and 1.131 grammes. In the next twelve hours, which were almost entirely occupied in exercise [this period extending from 8 A.M. to 8 P.M.], the diminution was still greater, being 2.498 and 1.225 grammes, which would be equivalent to 5 and 2J grammes for twenty-four hours. In the last twelve hours [8 P.M. to 8 A.M.] of rest after work, the elimination increased greatly, so that 5.142 and 3.331 grammes more were excreted than in the corresponding rest period." Seeking to reconcile his results in relation to muscular action, Dr. Parkes observes : " It appears to me that we can only express the facts by saying that a muscle during action appropriates more nitrogen than it gives off, and during rest gives off more than it appropriates." But must we, I would suggest, look only to the muscles for the source of the variation in the amount of nitrogen discharged in these experiments ? The results, in the first place, conclusively showed that the nitrogen eliminated forms no measure of muscular work performed, and hence it may be inferred as a corollary that muscular work is not a result of muscular destruction. But, taking the vari- ation in the voided nitrogen that was observable, independently of that occasioned by diet, why should we seek its source exclusively in the muscles? On looking at the several daily amounts discharged, I remark the DR. PARKES'S EXPERIMENTS ON ELIMINATION OF NITROGEN. 67 existence of instances in which considerable variation occurs within the periods themselves. Thus, during the first day of the first period, when the men were engaged in ordinary employment, B. dis- charged" 20.4 17 grammes of nitrogen, and during the third day only 17.090, a difference approaching to 3J grammes. Again, during the last period, which was also spent in ordinary employment (it will be remembered the daily diet was the same throughout the experiment), the urinary nitrogen voided by both men stood as follows : s. B. First day, . . 21.25 grammes. . . 20.25 grammes. Second day, . . 19.942 " . . 19.273 " Third day, . . 23.488 " . . 19.248 " Fourth day, . . 19.536 " . . 21.597 " On the third day, it thus appears, S. discharged nearly four grammes of nitrogen in excess of that on the fourth, and about 3 J in excess of that on the second. No corresponding fluctuation, it will be remarked, was observable in the case of B. Here, then, are marked variations in the elimination of nitrogen without a variation of muscular action. In a more recently performed experiment 1 Dr. Parkes's results show, with a fixed daily ingress of nitrogen, a variation in the daily exit .amounting in the extreme to seven and a half grammes. Now, we know that the nitrogen of the urine is derivable from the metamorphosis of the nitrogenous ingesta within the system. It is true the food taken was every day the same throughout the ex- periment that has been forming the subject of consideration, but it does not follow that the rate of metamorphosis was every day simi- larly identical. Doubtless, like other processes of life, it is influ- enced by various internal conditions. We know also, as the result of observation in the case of starvation, that, notwithstanding an absence of ingoing nitrogen, an elimination of this element still con- tinues, and that the nitrogen eliminated is drawn from the nitroge- nous principles of the body, belonging alike to the solids and fluids. There is a general waste or loss occurring, and the only difference noticeable is that the loss goes on with different degrees of rapidity in the different parts of the system. In the muscles it certainly occurs somewhat more rapidly than elsewhere, but this is all. With 1 Proc. Koy. Soc., March, 1871. 68 ALIMENTARY PRINCIPLES. these considerations before us it appears to me that we are taking an unjustifiably narrow view in looking only to the muscles to account for the variation in question in the voided nitrogen. Exercise can- not fail to influence the processes going on in the system generally, as well as in the muscles, and, in accounting for the results observed, instead of limiting ourselves, with Dr. Parkes, to the assertion that " we can only express the fact by saying that a muscle during action appropriates more nitrogen than it gives off, and during rest gives off more than it appropriates," I think what we ought rather to say is, that during exercise the system appropriates more nitrogen than it gives off, and during rest gives off more than it appropriates. Voit, however, disputes the reality of exercise producing any in- fluence over the elimination of nitrogen, and has taken exception to some of Dr. Parkes's experiments, on the ground, more particularly, that the daily ingress of nitrogen could not be kept sufficiently stable. This has elicited from Dr. Parkes a further series, the results of which are recorded in the " Proceedings of the Royal Society " for March, 1871. In these it appeared that there was no change in- duced, either at the time or afterwards, by a moderate amount of ad- ditional exercise under a mixed regulated diet ; but, under a non- nitrogenous diet, the increase in the nitrogen on the following clay to the performance of a hard day's march was exceedingly striking. The non-nitrogenous diet was continued through five successive days. During the first three it was associated with the ordinary work of a soldier ; on the fourth, with a march of thirty-two miles, carrying a weight of 43 \ Ibs. ; and on the fifth with rest. As the ordinary re- sult of abstinence from nitrogenous food, the eliminated urinary ni- trogen underwent a steady decrease during the first four days ; on the fifth, however, it showed a marked ascent, the amount being then in considerable excess of that discharged on the first. In the "New York Medical Journal" for October, 1870, Dr. Austin Flint, Jr., records the result of the examination of the urine secreted during the performance of, perhaps, an unprecedented amount of muscular work within the space of time occupied. A Mr. Weston, set. 32, of medium height, and weighing ordinarily 122 Ibs. without his clothes, celebrated as a pedestrian of the United States, undertook to perform the astonishing feat of walking one hundred miles in twenty-two consecutive hours. The feat, it ap- pears, was accomplished within the time namely, in twenty-one WESTON'S WALKING FEAT AND ELIMINATION OF NITROGEN. 69 hours and thirty-nine minutes. The food consumed during the period was taken in small quantities at short intervals, and con- sisted of between one and two bottles of beef essence, two bottles of oatmeal gruel, and sixteen to twenty raw eggs, with water. He drank a little lemonade and took water very frequently, but only in quantity sufficient to rinse his mouth. While walking the last ten miles he took, it is further stated, two or three mouthfuls of charn- pagne, amounting to about three fluid ounces, and about two and a half fluid ounces of brandy in ten-drop doses. The head and face were sponged freely at short intervals, and the food and drink were taken mainly on the walk, which was conducted within a covered inclosure. The urine passed during and at the completion of the walk meas- ured 73 J fluid ounces, and presented the specific gravity of 1011. According to Dr. Flint's analysis it contained 424f grains of urea. Now, 500 grains form about the average daily quantity of urea dis- charged under an ordinary mixed diet; and as the diet during the performance of the pedestrian feat was rich, as the account shows it to have been, in nitrogenous matter, the quantity of urea, apart from any other consideration, was even less than might have been ex- pected. And yet, on the strength of a comparison with another examination of the urine conducted three months later, when only 191 grains of urea are stated to have been discharged in the absence of exposure to muscular exertion, Dr. Flint argues that muscular ex- ercise notably increases the elimination of urea. To take a solitary result of so exceptional a kind as the discharge of 191 grains of urea in the twenty-four hours, and use it as a ground of comparison for reasoning upon, as Dr. Flint has done, is surely to violate all rules of sound induction, and it is to be hoped that we shall not find the observation quoted by writers as bearing out what Dr. Flint has contended for. During November, 1870, Mr. Weston undertook another pedes- trian feat, and this time a very elaborate examination was made of the ingesta and egesta, and of various conditions of the body, by Dr. Flint and a staff of associates. The results- are recorded in detail in the " New York Medical Journal" for June, 1871. The feat proposed was to w r alk 400 miles in five consecutive days, and upon one of the days 112 miles were to be walked in twenty-four con- 1 secutive hours. Mr. Weston commenced the undertaking on the 70 ALIMENTARY PRINCIPLES. 21st of November. The examination of the ingesta, egesta, &c., had been conducted for five days before; it was also carried on during the five days of the walk, and continued for five days after- wards. Thus, the results for three periods before, during, and after the walk were obtained. The subjoined tabular representa- tion will give a summary view of the leading points noted. The walk was undertaken over a measured track marked out in the form of a parallelogram, within a large covered space namely, the Em- pire Skating Rink in New York. It appears that Mr. Weston failed this time to accomplish the feat he had attempted ; distance walked during the five days amounting to 31 7 J miles, and the greatest dis- tance on any one day to 92 miles. DR. FLINT'S observations on the effects of the five-day pedestrian feat performed by MR. WESTON. Weight of body (uude). Before the walk. Ibs. First day, . . 120.5 Second, . . 121.25 Third, . . . 120. Fourth, . . 118.5 Fifth, . . . 119.2 During the walk. First day, . . 116.5 Second, . . 11625 "Dhird, . . . 115. Fourth, . . 114. Fifth, . . . 115.75 After the walk. First day, . . 118. Second, . . 120.25 Third, . . . 120.25 Fourth, . .. 123.5 JPifth, . . . 120.75, Tem- perature. Fahr. 99 7 98.4 980 99.1 99.5 95.3 94.8 96.6 96.6 97.9 986 98.4 99.3 98.8 97.5 Pulse. 75 73 71 78 93 93 109 68 80 76 73 70 78 76 Miles talked. 15 5 5 15 1 80 48 92 57 40.5 2 2 2 o Nitrogen in ingesta. Grains. 361.22 288.35 272.27 335:01 440.43 151.55 265.92 22861 144.70 383.04 385 65 499.10 394.83 641.71 283.35 Excess or Nitrogen deficiency in in t-gesta. nitrogen egested. Grains. Grains. 323.26 37.96 301.18 -f 12.83 330.36 + 58.09 300.57 34.44 320.06 120 37 357 10 -f 205.55 370.64 -f 104.72 397.58 + 168.97 348.53 -f 203 83 332.77 50.27 29570 8995 358.81 140.29 409 87 + 15.04 382.89 258.82 418.49 -h 135.14 Dr. Flint still holds to his former opinion, and looks upon the above results as showing, to use his own words, that " excessive and prolonged muscular exertion increases enormously the excretion of nitrogen, and that the excess of nitrogen discharged is due to an increased die-assimilation of the muscular substance/ 7 Let us accept Dr. Flint's estimates of the ingoing and outgoing nitrogen. It is true, during the first four days of the walking pe- 71 riod the exit of nitrogen was in considerable excess of the entrance ; but why should this be referred specially and exclusively to muscu- lar disintegration ? There was during these four days a progressive decline in the weight of the body, the loss reaching a little over 5 Ibs. From the account given, considerably less solid food was taken then, than before and after. There existed a state of marked disturbance of the bodily functions, as shown by the depression of temperature and elevation of pulse ; but little sleep was obtained ; and on the third day, when an attempt was made to walk the 112 miles in twenty-four consecutive hours, drowsiness, it is stated, pre- vailed to such an extent that it was found impossible to make the necessary time to accomplish what had been intended. On the fourth day Mr. Weston actually broke down for a time altogether, becom- ing dizzy, staggering, and at last failing to be able to see sufficiently to turn the corners of the track. Xow, apart from the fact that a marked deviation from the physi- ological state existed when the results, upon which the conclusions are based, were yielded, is there anything in the results to show that in reality we have more to deal with than simply a consumption of nitrogenous material within the system beyond the supply for the time from without? Taking the figures throughout, there is not much more to be seen than a difference occasioned by a falling off in the amount of nitrogen ingested during the first four days of the walk ; and it is well known that when the ingesta do not furnish what is wanted for meeting the expenditure going on (as during in- anition), the resources of the body are drawn upon, and the nitroge- nous matter existing in the various parts both solids and fluids wastes or yields itself up as well as the rest. On the fifth day, after a prolonged sleep, which appears to have restored the flagging pow- ers, the previous relation was reversed. The food ingested afforded more than enough to meet the requirements. There was a gain of If Ib. in body-weight, and, according to the figures, the nitrogen discharged fell short by 50.27 grains of that which entered, notwith- standing a walk of forty miles and a half was performed. The distance walked during the five days amounted to 31 7| miles, and the excess of nitrogen eliminated during the time, over that in- gested, appears to have been 633 grains. Presuming, for sake of argument, this to have represented the nitrogen of muscle disin- tegrated in the accomplishment of the work performed, we have be- 72 ALIMENTARY PRINCIPLES. fore us the data for ascertaining how far the force producible in this way would correspond with the expenditure that must have occurred. According to Mulder's analysis, albuminous matter contains 15.5 per cent, of nitrogen. Reckoning from this proportion, 633 grains of nitrogen will correspond with 4083 grains of dry albumen, and the composition of the nitrogenous matter of muscle is closely anal- ogous. Now, the force producible from the oxidation of albumi- nous matter has been experimentally ascertained by Frankland; and, as it occurs within the body, the oxidation of 4083 grains of dry albumen would give rise to the evolution of an amount of power equal to lifting 1540 tons one foot high. Here we have one side of the question the amount of work ob- tainable from the nitrogenous matter presumed to have undergone disintegration as muscular tissue and so far the information in our possession may be regarded as sufficiently authentic to enable us to frame a reliable conclusion. As regards the work accomplished, we may assume, with Professor Haughton, that the force expended in walking or progressing on level ground is equal to that required to lift one-twentieth of the weight of the body through the distance traversed. The distance walked amounted to 317J miles, and if we take the weight of the body and clothing at, say, 120 Ibs., this will give the performance of an amount of work equal to lifting 4490 tons one foot high, or about two-thirds more work than the oxida- tion of the nitrogenous matter representing the 633 grains of nitro- gen could accomplish. And, in this calculation, only the external work has been taken into consideration. There is, in reality, also a considerable amount of internal work constantly being performed viz., that employed in keeping up the circulation, in respiration, and in various other essential actions of life. I have entered thus minutely into the question of the elimination of nitrogen in relation to muscular work because it bears in so forcible and direct a manner upon the question immediately before us viz., the uses to which the nitrogenous alimentary principles are applied in the system. Briefly represented, the position of the niat- ter may be said to be this : Many years ago it was asserted by Liebig that muscular action involved the destruction of muscular tissue. The plausibility of the doctrine, and the readiness with which the views of its author were then received, must be considered as having led to its being at once RSUM ON NITROGENOUS FOOD AND MUSCULAR ACTION. 73 generally accepted as though it formed a scientific truth, although, in reality, only constituting a speculative proposition, unsupported by anything of the nature of proof. It was further argued that, if muscular action involved the destruction of muscular tissue, the ex- cretion of the nitrogenous product of destruction urea ought to be in proportion to the amount of muscular work performed. This seemed to follow as a necessary sequence, and the one being accepted, the other was taken for granted also. Thus, notwithstanding the absence of anything in the shape of proof, we find physiologists rea- soning and writing as though the doctrine had been actually proven. If the theory of Liebig were true, we should have to look upon nitrogenous alimentary matter as forming, through the medium of muscular tissue, the source, and the only source, of muscular power. The renewal of muscular tissue for subsequent oxidation in its turn, and evolution of muscular force, would thus constitute one of the functions of nitrogenous alimentary matter; and, on its supply would, accordingly, depend our capacity for the performance of muscular work. It is only lately that the doctrine has been submitted to the test of experiment, and with what result the foregoing account of the researches of various observers has shown. Even Liebig 1 has now come to assert that muscular action is not attended by the produc- tion of urea. He admits that the question as to the source of mus- cular power has been complicated by an inference which has proved erroneous, and for which he acknowledges himself as responsible the inference, namely, that muscular work is represented by the metamorphosis of muscular tissue, and the formation of urea as a final product. While admitting this much, however, Liebig still looks to changes in the nitrogenous constituents of muscle as the source of muscular power. He assumes the presence in muscle of nitrogenous substances in a much higher state of tension than syn- tonin and albumen, and to these he refers the performance of mus- cular work, taking shelter under the proposition that it is due to the liberation of the tension thus presumed to have been accumulated in them during their formation. The application of food to the genesis of muscular power will form 1 Proceedings of the Koyal Bavarian Academy of Sciences, 1869. Pharma- ceutical Journal, 1870. 74 ALIMENTARY PRINCIPLES. the subject of further consideration hereafter, when we reach the head of non-nitrogenous matter. Suffice it here to reiterate that muscular action is not to be considered as the result of muscle-destruction, as was formerly supposed, and hence, that nitrogenous matter is not applied through muscle in the manner hitherto maintained to the development of muscular force. Thus much, from the evidence before us, may be said, but, at the same time, common experience seems to show that a plentiful supply of nitrogenous matter in the food tends to increase the capacity for the performance of muscular work. If, however, it does so in any other way than by supplying material for nutrition and the secretions, and so contributing to the production of a fully nourished and vigorous state of the system, we have no data before us to indicate how. Let me next draw attention to the application of nitrogenous matter to force-production by the direct utilization of the carbon and hydrogen it contains. Liebig's doctrine, which, until recently, has formed the accepted one on this point, was that nitrogenous food, to be turned to account for force-production, must pass through the condition of Jiving tissue. This brings us back to the discussion that has preceded, with the addition that our nitrogenous food must perform work as tissue to enable it to be susceptible of application to force, or say heat-production. Thus, in his work on "Animal Chemistry," at page 60, Liebig says, "The flesh and blood con- sumed as food yield their carbon for the support of the respiratory process, whilst the nitrogen appears as uric acid, ammonia, or urea. But, previously to these final changes, the dead flesh and blood be- come converted into living flesh and blood, and it is, strictly speak- ing, the carbon of the compounds formed in the metamorphosis of living tissues that serves for the production of animal heat." Again, at page 77, we find " Man when confined to animal food respires like the carnivora at the expense of the matter produced by the metamorphosis of organized tissues ; and just as the lion, tiger, and hyena, in the cages of a menagerie, are compelled to accelerate the waste of their organized tissues by incessant motion, in order to fur- nish the matter necessary for respiration, so the savage, for the very same object, is forced to make the most laborious exertions and go through a vast amount of muscular exercise. He is compelled to consume force merely in order to supply matter for respiration." NITROGENOUS FOOD AND HE AT -PRODUCTION. 75 Once more, in speaking of the derivation of urea from the metamor- phosis of nitrogenous matter, he says, at page 144 : " There can be no greater contradiction with regard to the nutritive process than to suppose that the nitrogen of the food can pass into the urine as urea, without having previously become part of an organized tissue." Liebig's idea, then, upon this point is very precise. He considers that nitrogenous matter may contribute towards heat-production, but that it must first pass into the condition of tissue before it can do so, and that it is in the wear and tear of tissue that occurs the splitting up of the compound, so as to lead to the production of urea for excretion on the one hand, and the liberation of carbon and hy- drogen for oxidation on the other. The facts which have been already adduced suffice to refute this doctrine. Indeed, it may be considered as now abundantly proved that food does not require to become organized tissue before it can be rendered available for force-production. But Liebig, himself, in language not less precise than that which he at first employed, has recently 1 given utterance to words which directly contradict his original view, inasmuch as he now asserts that muscular work and the production of urea bear no immediate relation to each other, and that among the products formed as the result of muscular action, urea certainly does not even constitute one. If the elimination of urea, as has been shown, is not related, as was formerly supposed, to muscular action, it is, on the other hand, in a very direct manner influenced by the food ingested. As far back as 1854, Messrs. Lawes and Gilbert, in opposition to the views then prevailing, showed by the results obtained in their observations on the feeding of cattle, that the nitrogen in the urine is related to that in the food, and not to the muscular work ; and, since then, the concurrent testimony of numerous observers, as has been already pointed out, may be held as completely establishing this position. Lehmann's well-known experiments upon himself strikingly illus- trate the extent to which this influence is manifested. The results he obtained were as follows : While living on a purely animal diet, namely, almost exclusively on eggs, Lehmann passed 53.2 grammes (820 grains) of urea in the twenty-four hours as the mean of twelve observations. 1 Proceedings of the Royal Bavarian Academy of Sciences, 1869. 76 ALIMENTARY PRINCIPLES. Upon a mixed diet the urea amounted to 32.5 grammes (501 grains), as the mean of fifteen observations. Upon a vegetable diet the urea given as the mean of twelve ob- servations was 22.5 grammes (347 grains). And, lastly, upon a purely non-nitrogenous diet (fat, milk-sugar, and starch), he voided, as the mean of three observations, only 15.4 grammes (237 grains) of urea. It is thus seen that upon an animal diet, which is the richest in nitrogenous matter, the voided urea more than doubled that elimi- nated upon a vegetable diet, while the amount of urea voided upon a mixture of the two kinds of food held an intermediate position. When no nitrogenous matter was ingested the urea was at its mini- mum. What was then passed would be derived from the metamor- phosis of the nitrogenous matter belonging to the blood and the other constituents of the system. Some experiments of Schmidt show, also, in accordance with the results obtained by Lehmann, that the amount of urea passed is re- lated to the quantity of food ingested, the nature of it remaining the same. Schmidt found that a cat excreted the following relative amounts of urea to body-weight under the consumption of different amounts of meat : J'aily amount of Daily amount of urea excreted meat eaten. per kilogramme body-weight. 44.188 grammes. . . . |. 2.958 grammes, 46.154 " ..... 3.050 75.938 5.152 " 108.755 " 7.663 " From these results it may be computed that a cat, living on a flesh diet, discharges by the kidneys on an average 6.8 parts of urea for every hundred parts of meat consumed. The great bulk of the nitrogen belonging to the food ingested thus passes out of the system in the form of urea. If all escaped in this way the quantity of urea discharged would amount to (say) 7.88 per cent, of the weight of the meat ; the nitrogen contained in 100 parts of flesh corresponding with that contained in 7.88 parts of urea. There were, then, 6.8 parts of urea produced instead of the 7.88 parts, which may be spoken of as representing the actual equivalent, as far as contained nitrogen is concerned, of 100 parts of flesh. METAMORPHOSIS OF NITROGENOUS FOOD. 77 Lehmann, from his observations on himself, asserts that as much as five-sixths of the nitrogen of the ingested food were found in his urine under the form of urea. For example, while living upon a purely animal diet, consisting of thirty-two eggs daily, he ingested about 30.16 grammes of nitrogen, and, in the urea voided, dis- charged about 25 grammes of nitrogen. The discharge of urea being thus proportioned to the amount of the nitrogenous matter ingested, it follows that nitrogenous matter must undergo metamorphosis of such a nature within the system as to lead to the production of urea. Further, it may be said that this metamorphosis must take place rapidly, as it is found that the effect upon the excretion of urea quickly follows an alteration in the food ingested. Lehmann, for example, again drawing from his observa- tions on himself, noticed, in the morning after he had lived exclu- sively on animal food, that his urine was so rich in urea as to throw down a copious precipitate of the nitrate, on the addition of nitric acid. In Dr. Parkes's observations, also, upon the two soldiers S. and T. before referred to, the alterations in the food ingested speedily influenced the amount of urea escaping. These men were, first of all, kept for four days upon a regulated mixed diet; next for two days upon a non-nitrogenous diet; then, again, for four days upon a mixed diet ; afterwards for two days on a non-nitrogenous diet ; and, lastly, for four days on a mixed diet. S., during the first four days, on the mixed diet, passed 35 grammes of urea as the daily mean. During the first day of the non-nitrogenous diet he passed 20, and during the second, 13.52 grammes. Resuming the mixed diet, he passed, on the first day, 20.67 grammes of urea; on the second, 25.68 grammes ; on the third, 26.29 ; and on the fourth, 29.67. Changing, again, to the non-nitrogenous diet, he passed on the first-day 19.12, and on the second, 15.00 grammes of urea. On the next four days, the diet being a mixed one, he passed during the first day, 20.8 ; the second, 26.36 ; the third, 28.32 ; and the fourth, 30.10 grammes of urea. With T. (a much smaller man than S.) the mean, for the first four days of mixed food, was 25.92 grammes of voided urea. During the next two days, upon non-nitrogenous food, he passed, on the first day, 17.3; and on the second, 12.65 grammes. On the following four days, upon a diet of mixed food, he voided 14.40 grammes the first day ; 23.00 the second ; 25.20 the 78 ALIMENTARY PRINCIPLES. third ; and 22.99 the fourth. During the next two days, resuming the n on -nitrogenous diet, he voided 16.00 the first day, and 13.20 grammes the second. With the return to a mixed diet during the following four days the urea stood at 23.00 on the first ; 24.36 on the second; 24.57 on the third ; and 21.36 grammes on the fourth. Although conducted for settling another point, it will be seen that these observations very clearly and consistently throughout show that the production and elimination of urea are speedily affected by the ingestion of nitrogenous matter. With the view of obtaining more precise information regarding the time required for the metamorphosis of nitrogenous matter to occur and lead to an increased elimination of urea, Mr. Mahomed, whilst formerly assisting me in my laboratory, carried out, with laudable zeal and self-denial, two series of experiments upon him- self, the particulars of which I will introduce here. It may be men- tioned that he was 22 years of age, 6 feet in height, and list. lllb. in weight. The method of procedure had recourse to was to diminish the elimination of urea by limiting in one experiment, and withholding in the other, the introduction of nitrogenous matter, and then note within what space of time the ingestion of nitrogenous matter showed its effects upon the urine. The first experiment was commenced on April 16th, 1871. Mr. Mahomed had been previously living upon an ordinary mixed diet, and took his dinner of mixed food, as usual, at 1.30 P.M. From this time he restricted himself to rice, arrowroot, butter, sugar, and tea. Rice was allowed, that he might not suffer too much privation, and as being one of the least nitrogenous of the natural food prod- ucts. The diet was continued throughout the 17th, and at 8 A.M on the 18th, four eggs purposely to supply nitrogenous matter were eaten. This was the only deviation from the diet of the preceding day, so that an opportunity was given for the urea to be again at a low point on the following morning, when a meal consisting mainly of meat was taken. Subjoined is a representation of the results ob- tained, arranged in a tabular form : METAMORPHOSIS OF NITROGENOUS FOOD. 79 r ^3 05 . a A c to OS z 01 OQ rt 5? > -5 ^ Sb '0* 3 J c ^ to g 1 -Ul oo O $ to S Ci r^ t< rrt $ '? i S -i 's - 3 M C o o g & * I s:? * b c S S a g' J 3 1 ] jr * ^ - 0} c g O ' ' ' t S . ' ' ' r -\ * a s s s ^ s a a a a ass?, g - C ^ iO I s ** OS CO ^ O CO OS 1-H -^ CO O CN TJH TfH 1^ O _i co ^ o -H i^ co o co co c^ "^ id co c* 1 c - o g c ; M CO CO iO i COCO'^tlC^Ji-O OS O -< CO OS M< O r-i t^ CO ? - r r CO CO OS i CNS^CNCOCO OCO^^CO lOCOTHCO'*^ ||5 ill CO CO CO CN CO O 10 10 10 O O r- t~- i- CM c-i t>- iO O CO CO COCOi-H 11 | CCOOO 5>l t- -^ -^ -< rjl id id oi ^ co' Tji co' co co* *** tc a "& a . a a a a a . a | . rf ^ ^ 1 s ri a qooq O^PHO^ ^H "* CO ' CO'-OCO^H llj-ll! Ill +j >-H O CO i i j r^ v *O OO v "^ 3333 33^33 o -^O^OO H^OOOOO J3+a*J-4J j3-u*J*j^j*J 1 asss sa^ss 'S s c s ' s '5 s s s a' a 8, oooo oocoo CO COO COOOOO a OOr-.Tt