mm .:,'.'..'; ••■,■.•■■':-•: ■::%■':•■•■ H o^:!';,:ii^:>:^'';;v;;:::^': ana gg ' ■ Will THE INFLUENCE OF INANITION ON METABOLISM BY FRANCIS GANO BENEDICT WASHINGTON, D. C. Published by the Carnegie Institution of Washington 1907 CARNEGIE INSTITUTION OF WASHINGTON Publication No. 77. IH Zk Z9>t JSorb (§a(timovt (pvtee BALTIMORE, MD., IJ. 8. A. PREFACE. The primary Gbject of this report is to present an accurate statement of the results of experiments on the effect of inanition on metabolism made with a number of men in the chemical laboratory of Wesleyan University, Middle- town, Connecticut. It is believed that physiology and physiological chemistry will be best served by presenting the data accurately and in sufficient detail rather than to sacrifice the protocols for a discussion that at best can be but inadequate. The accumulation of the experimental data, the computation of the results, and the preparation of this report have been made possible only by the fidelity and loyalty of my associates. Mr. T. M. Carpenter, in immediate supervision of the greater number of the respiration calorimeter experiments, has conducted these most wearisome experiments with unusual success. Mr. H. A. Pratt, aside from rendering valuable assistance in the chemical laboratory, has superintended the computations and tabulations, and the entire report has received his helpful editorial criticism. Mr. E. M. Swett acted as physical and chemical assistant and superintended the determinations of the heats of combustion with the bomb calorimeter. Miss Charlotte E. Manning had charge of all the gas analyses, carbon and hydrogen combustions, creatinine determinations, and the analyses of food. Mr. E. E. Fulton made all the determinations of sulphur and phosphorus. Mr. F. P. Fletcher acted as physical assistant in the later series of calo- rimeter experiments. Messrs. J. A. Eiche and E. E. Hartman assisted in the chemical and physical measurements. Mr. W. H. Leslie, Miss A. N". Darling, and Mr. H. C. Morgan have had a large share in the tabulation of the results of the experiments and in the final preparation of the report. The stenographic work was in charge of Miss A. 1ST. Darling, who was ably assisted by Miss M. K. Falsey. Aside from a corps of students, special mention should be made of the assistance in the computations rendered by Messrs. H. L. Knight and F. W. Harder and Misses H. W. Atwater, E. J. Wright, and H. L. Ailing. It is a great pleasure to express my deep sense of obligation to all of these co-workers in the conduct and presentation of the results of experiments that call forth all the patience and accuracy of the trained chemist, physicist, and computer. ii Influence of Inanition- on Metabolism. My thanks are especially due to Dr. A. R. Diefendorf, pathologist of the Connecticut Hospital for the Insane, Middletown, Connecticut, and Prof. Lafayette B. Mendel, of Yale University, New Haven, Connecticut. I have from time to time enjoyed the valuable counsel and advice of both these gen- tlemen. Dv. Diefendorf personally made a very large number of the blood examinations and furnished a statement which has been embodied in this report. Professor Mendel suggested the desirability of determining the creatinine in the samples of urine in the fasting experiments. Some of the determinations he kindly made in the New Eaven laboratory. In this work he was assisted by Mr. 0. E. Closson. F. G. B. Chemical Lauoratoky, Wesleyan University, Middletown, Connecticut, December 21, 1906. CONTENTS PART 1. Page. Introduction 1 Plan and purpose of the experiments here reported 7 Method of investigation 8 Grosser observations 8 Physiological measurements 9 Chemical measurements 12 Respiratory products 15 Physical measurements 16 Recording results and the use of decimals 17 The experimental man 18 PART 2. Statistics of experiments 19 Metabolism experiment No. 59 20 Urine 20 Elimination of water-vapor 30 Elimination of carbon dioxide 31 Oxygen consumed 33 Material katabolized in the body 36 Output of heat 42 Total heat production 46 Balance of energy 51 Relations between oxygen consumption, carbon dioxide elimina- tion, and heat production 52 Respiratory quotient 53 Metabolism experiment No. 68 55 Metabolism experiment No. 69 71 Metabolism experiment No. 70 85 Urine 88 Elimination of water-vapor 90 Elimination of carbon dioxide and absorption of oxygen 90 Material katabolized in the body 90 Output of heat 93 Balance of energy 94 Relations between oxygen consumption, carbon dioxide elimina- tion, and heat production 94 Effect of ingestion of food 94 Changes in body-weight compared with balance of income and outgo 102 Balance of intake and output 103 Metabolism experiment No. 71 1 07 Metabolism experiment No. 72 126 Metabolism experiment No. 73 136 in iv Influence of Inanition on Metabolism. Page tatiBtics of experiments Continued. Metabolism experiment No. 74 L54 Metabolism experiment No. 75 169 Metabolism experiment No. 76 192 Metabolism experiment No. 77 207 A series of 2-day fasting experiments with 7 subjects, metabolism experi- ments Nos. 79-83, 85, and 89 222 Urine 243 "Weight and composition of urine 244 Elimination of water-vapor 245 Elimination of carbon dioxide and absorption of oxygen 245 Elements katabolized in the body 245 Changes in body-weight compared with balance of income and outgo 262 Output of heat 263 Balance of energy 268 Relations between oxygen consumption, carbon dioxide elimina- tion, and heat production 268 Nitrogen metabolism experiments 274 PART 3. Discussion of results 300 Experiments during inanition 301 Body-weight 301 Body temperature 311 Pulse rate 316 Rate of respiration 320 Blood 322 Strength tests 327 Physician's report 334 Subjective impressions and observations regarding general conditions 335 Feces 337 Urine 345 Volume 345 Relation of drinking-water to volume 348 Reaction 351 Specific gravity 351 Total solids 352 Ratio of the total solids to the specific gravity 353 Constituents of total solids 356 Proportions of ash and organic matter in total solids 360 Nitrogen 361 Total nitrogen 362 Excretion of nitrogen on days preceding fast 366 Ratio of total solids to nitrogen 378 Ratio of organic matter to nitrogen 381 Excretion of carbon in the urine 382 Creatinine and creatine 386 Uric acid 395 Contents. v Page. Discussion of results — Continued. Experiments during inanition — Continued. Urine — Continued. Pathological constituents of the urine 397 Sulphur 397 Phosphorus 406 Chlorine 413 Water output 420 Water of feces 421 Water of urine 421 Water of respiration and perspiration 421 Carbon dioxide elimination 438 Relation of carbon in urine to total carbon 444 Oxygen consumption 446 Respiratory quotient 449 Body materials katabolized 452 Protein 453 Fat 461 Glycogen 463 Water 466 Energy 474 Heat elimination 474 Heat production 478 Muscular activity 484 Energy of the urine 490 Energy of katabolized body material 495 Comparison of products of katabolism and heat production 497 Oxygen and carbon dioxide thermal quotients 504 Comparison of all factors of katabolism 508 Energy balance 510 Experiments with food 516 The digestibility of food after inanition 519 Digestibility of food in short experiments 522 Digestibility of food in the nitrogen metabolism experiments. . . . 524 Influence on general physical condition 525 Influence on general metabolism 527 Balance of matter and energy 531 Recovery after inanition 533 THE INFLUENCE OF INANITION ON METABOLISM. PART 1. INTRODUCTION. In the processes of life the transformations of matter with the energy changes dependent upon them are of two markedly different kinds — katabolism or the breaking down of body material, and anabolism, or the construction of body material. Considering the body as consisting not only of the skeleton, muscles, glandular, adipose, and other tissue, but also of the fluids, such as blood and lymph, it is perfectly proper to consider katabolism as that process by which body material is broken clown as a result of oxidation and cleavage. Although under certain conditions of inanition there may be distinct anabolic processes occurring in the body, such as the much discussed formation of fat and sugar from proteid, yet in general it is customary to assume that anabolism follows the ingestion of food. This anabolic process consists in taking certain molecular complexes, such as those in the protein, fat, and carbohydrate of food, and rearranging the fragments of the molecules in such a manner that the re-formed materials are of a molecular composition best suited to or comparable with the structure of the body. While the molecular structures of the protein, fat, and carbohydrate of the food are in general similar to those of the body materials, differences — indeed material differences — are not lacking. Perhaps in no group is this lack of complete uniformity of structure more striking than in the case of protein. Eecent investigation of the cleavage products in the animal and vegetable proteins 1 shows wide differences in the kinds and amounts of the molecular complexes of which these intricate molecules are composed. It is evident, therefore, that the final products of metabolic activity of the living body are the resultants of the anabolic and katabolic transformations which take place. Hence each study of metabolism is a twofold problem in which is considered, first, katabolism, or the actual disintegration of body material; second, anabolism, or the transformation of food materials into body material. Since under normal conditions these two processes are continually occurring side by side and their influences are correspondingly interrelated, it is emi- nently desirable, if possible, so to adjust the conditions of experimentation as to eliminate the complex relations necessarily existing between the processes of anabolism and katabolism. 1 Osborne & Gilbert, Amer. Journ. Physiol. (1906), 15, p. 333. 2 Influence of Inanition on Metabolism. Food is usually ingested at more or less regular periods, so tliat the anabolic processes proceed without interruption; but, by withholding food, the anabolic activities may be depressed to such an extent as to make the study essentially one of katabolism. Consequently, studies of the transformations in the body during inanition are of great value and, logically, at least, should precede the studies in which anabolic and katabolic processes are combined. While it may legitimately be considered that the first day without food is not a true fast, i. e., a metabolism in which body material alone is involved, because of the presence of unabsorbed or partially digested food in the ali- mentary tract, it nevertheless seems highly probable that by the time the second day of fasting begins the body is living essentially upon its own substance. Since, however, the retention of fecal matter may result in a more or less prolonged absorption and thus vitiate in a small way the assumption that only preformed body material is being consumed, experiments of more than two days' duration are necessary, and consequently experiments during pro- longed fast should be included in any complete and accurate study of meta- bolism during inanition. Although as a rule man is disinclined to fast more than half a day, many persons have lived with no food whatever for periods as long as 30 or 40 days. It is important here to distinguish between complete abstinence from both food and water and abstinence from food alone. Experiments have shown that life can not be sustained for any considerable period when both food and drink are withheld. It was the popular impression, at least in the early and middle ages, that certain persons were able to subsist upon body material alone for much longer periods of time, extending, indeed, into years. Fasts of more than one day's duration may be divided into six classes. RELIGIOUS FASTING. Fasting as a religious rite has in many recorded instances been prolonged, and more or less complete. The accounts of such fasts are, however, so clouded by superstition and show such a lack of accurate observation that they are without value to science. They served only to maintain popular belief in the ability of some religious ascetics to subsist solely on the eucharist, and of some possessed of devils to abstain from food altogether. Of the numerous recorded instances of this form of complete or partial inanition, many are cited by Hammond2 and Luciani." 'Fasting girls; their physiology and pathology. New York, 1879. In this little volume Hammond refers to cases cited by Gbrres (La Mystique divine naturelle et diabolique, Paris (1861), 1, p. 194); and Wanley (Wonders of the little world, London, 1806, p. 375). * Das Hungern, Leipzig, 1890, p. 70. Introduction. 3 Aside from those directly cormected with, the church and religious orders, numerous instances of prolonged fasting, in general by young girls, are recorded.4 However interesting these and the many similar instances may be to the psychologist and theologian, they can of necessity have no scientific value, and any attempt to discuss them and the degree of probability of the various con- ditions asserted as controlling them would be out of place here. Our present- day knowledge of the processes of metabolism is fixed to such a degree that fasts as prolonged as many of the religious fasts were asserted to have been are inconceivable. It remains, however, to be proven that there may not be instances of suspended animation approximating the hibernation of the cold- blooded animals, in which man may subsist on his own body substance for a period of months. On this point, however, scientific observations are lacking. FASTING OF THE INSANE. A characteristic of many delusions is a revulsion towards food and drink, so marked indeed that in many instances all the skill of the trained psychiatrist is required to combat it. In a large proportion of such cases, artificial feeding must be resorted to. In a number of instances such insane patients have been allowed to fast for a number of days and observations of more or less value have been made upon their general condition of nutrition. In many other instances the fasting had proceeded for a considerable time before the condition of the patient became known to the physician. Hammond 5 cites the case recorded by Esquirol 8 of a person suffering with melancholia who died after 18 days of complete absti- nence; Desportes,7 of a similar patient who lived for two months, consuming only a little water. Francis 8 reports a case of a supposedly bewitched negress who took but two small cups of water during 3 weeks. ITcXaughton " reports a case of a young man who lived with no food, but with water, for 53 days. Other cases are cited by Luciani,10 and it is unmistakably true that fasts of weeks, if, indeed, not months, have been observed in cases of insanity. Modern 4 Hammond refers specifically to the work of Bucoldianus, " De puella quae sine cibo et potu vitam transigit," Parisiis Ann., mdxlii; Citesius, Opuscula Medica, Parisiis, 1639, p. 64; Fabricius, Observationum et curationum chirurgicae, centuria secunda, Genevae, 1611, p. 116; Fowler, A complete history of the Welsh fasting girl (Sarah Jacob), with comments thereon, and observations on death from star- vation, London, 1871. 5 Loc. cit., p. 64. "Des maladies mentales, Paris, 1838, p. 203. 7 Du refus de manger chez les alienes, These de Paris, 1864. 6 New York Medical and Surgical Journal, vol. n, p. 31, cited by Hammond. 0 Copeland's Dictionary of Medicine, vol. i, p. 31, cited by Hammond. 10 Loc. cit., p. 218. 4 Influence of Inanition on Metabolism. methods of treatment preclude such prolonged fasts, since as a rule nutritive enemata and tube-feeding are regularly employed. STARVATION THROUGH ACCIDENT. There are on record many authentic cases of persons in shipwrecks, coal- mine disasters, etc., who have been deprived of food for considerable periods. In a number of these cases u individuals have been rescued after having with- stood fasts of from 14 to 25 days, though in nearly every instance drinking- water, at least in limited amounts, was accessible. PATHOLOGICAL FASTING. Pathological cases involving disturbances of the alimentary tract which preclude the ingestion of food are unfortunately only too common, and since the majority of such cases have been in the hands of intelligent physicians, careful observations have been recorded in many instances. Since the method of treatment involved the ingestion either per os or per rectum of varying quantities of food, such cases are not comparable with complete fasting. Furthermore, the pathological nature of the cases may in many instances have resulted in an abnormal metabolism. FASTING IN HYPNOTIC SLEEP. To this class undoubtedly belong many of the so-called cases of suspended animation which have frequently been observed in the Hindu fakirs, and more especially in recent years in certain cases in which hypnotic suggestion has been of practical use in studying problems of nutrition. The best known instance of the use of hypnosis for this purpose is the case reported by Hoover and Sollman.12 PHYSIOLOGICAL FASTING. Physiological fasting as distinguished from fasting as a result of mental or pathological lesions may be defined as the voluntary fasting of normal subjects. Either for purposes of exhibition or scientific experiment, a number of such fasts more or less prolonged have been made. Giovanni Succi, the professional faster, has made a number of fasts of from 10 to 30 days' duration. During many of these fasts scientists have cooperated to secure more or less complete physiological studies of the effect of fasting on metabolism. Other professional fasters — Cetti, Breithaupt, and Jacques — have been the subjects of experiments Avhich have contributed greatly to our knowledge of the physiology of fasting. One of the most elaborate of such studies is that "Luciani, loc. cit, p. 73; Hammond, loc. cit, p. 62. 12Journ. of Experimental Medicine (1897), 2, p. 403. Introduction. 5 of a 5-day fast made with a medical student, " J. A.," in the Stockholm labora- tory by Johansson, Landergren, Sonden, and Tigerstedt. These and other experiments with men will frequently be referred to in this report. An examination of the various kinds of fasts outlined above shows that a distinction may be made between what may be termed normal and pathological fasting. The first, second, and probably even the third class should be included, with the fourth, under the head of " Pathological fasting." A criticism has been raised 13 as to whether insane patients are abnormal as regards their metabolism. According to Tuczek " there is every reason to believe that in many instances persons mentally unsound have an abnormal metabolism. Admitting that the mental attitude may produce abnormalities in metabolism, it is not at all unreasonable to suppose that in instances where persons are deprived of food through accident the mental strain may likewise result in abnormal metabolism. It is thus seen that only in the case of the hypnotic subjects, professional fasters, and other voluntary subjects, can we have what may be clearly asserted to be normal or " physiological " fasting. STUDY OF FASTING WITH ANIMALS. Eecognizing the difficulty of securing willing human subjects, investigations on the metabolism of fasting animals, usually dogs, have been made. The results obtained are of value to physiology in general, but of less value to human physiology. The metabolism of the dog is not that of man, for the nature of the food is such as to demand markedly different treatment in the alimentary tract. Both the stimuli to secretion and the composition of the digestive fluids are markedly different, and it seems probable that the deductions from the experiments with animals are of questionable value when applied to man. This is perhaps even more noticeable in experimenting in pharmacology than in physiology, yet it is true that marked differences in physiological character- istics are observed between animals and men. While, therefore, in this report the attempt is made to give a complete bibliography of scientific experiments on fasting men, reference in the text to experiments on animals is made only as occasion demands, and there is no attempt to present a complete list of the experiments on fasting animals.15 13Atwater, Eleventh annual report of the (New York) State commission in lunacy (1899), p. 202; Folin, Amer. Jour. Insanity (1904), 60, No. 4, and 61, No. 2. "Archiv fur Psychiatrie u. Nervenkrankheiten (1884), 15, p. 784. 15 F. A. Falck, Beitrage zur Physiologie, Hygiene, Pharmakologie und Toxikologie (1875), has an excellent bibliography of all work on metabolism during fasting completed prior to 1875. Other bibliographies of the subject are given by Weber, Ergebnisse der Physiologie (1902), 1, p. 702; Schaefer, Textbook of Physiology (1898), vol. i, p. 891; Atwater & Langworthy, A digest of metabolism experiments (1898), Bull. 45, Office of Experiment Stations, U. S. Dept. of Agriculture; in the preparation of this report, the excellent Index Catalogue of the Library of the Surgeon-General's Office, U. S. Army, Washington (1880-1904), has been of great assistance. 6 Influence of Inanition on Metabolism. study of fasting with men. While, as was seen in a preceding section, it has not been uncommon for man to fast, efforts to make a scientific study of fasting other than the general observations as to loss in weight, strength, etc., were not made until a com- paratively recent date. Autopsies were occasionally made on the bodies of persons who had died as a result of inanition and the results recorded by attending physicians, but the first scientific experiment to determine the effect of fasting on metabolism in man was not made until 1825, when Lassaigne 1S studied the urea output of an insane patient who fasted 18 days. The first observations on the carbon dioxide elimination of man during inanition were made by Scharling " in 1813. The most extended researches on the metabolism of man during inanition are those made on the professional faster Succi by Luciani," E. Freund and 0. Freund,19 D. Baldi,20 Ajello and Solaro,21 Daiber,21a Brugsch,22 and Tauszk.23 The experiments of Lehman, Miiller, Munk, Senator, and Zuntz,24 on the pro- fessional fasters Cetti and Breithaupt, and the observations of Johansson, Sonden, Landergren, and Tigerstedt 25 on a medical student, included deter- Note. — Since this report was written, three papers reporting the results of a 15- day experiment (March 9-24, 1906) with a professional fasting woman (Schenk) have appeared from the second medical clinic in Berlin. These three papers appeared in the Zeitschrift fur experimentelle Pathologie und Therapie (1906), vol. 3: Gesammt-N- und Aminosaurennausscheidung im Hunger, Dr. Theodor Brugsch and Dr. Rahel Hirsch, pp. 638-645; Die Saurebildung im Hunger, M. Bon- niger and L. Mohr, pp. 675-687; Ueber die Darmfaulniss im Hunger, R. Baumstark and L. Mohr, pp. 687-691. In a private communication Dr. Otto Folin, of the McLean Hospital at Waverley, Mass., has announced the analyses of the urine in a 6-day fasting experiment with an insane man. Dr. Folin also writes that Dr. E. P. Cathcart, of the University of 5*v«*3«%OM* Edinburgh) is at present studying the nitrogenous output of man during inanition. ' Neither of these investigators has as yet published his results. With the cooperation of Dr. A. R. Diefendorf, of the Connecticut Hospital for the Insane at Middletown, Connecticut, the writer has recently completed the analyses of the urine of a fasting insane woman who abstained both from food and drink for 110 hours and from food a total of 161 hours. The results are reported in the American Journal of Physiology, 1907, 18. 16 Jour. d. chim. med. (1825), i, p. 272, cited by Voit (Herman's Handbuch der Physiologie (1881), vi, i, p. 84). Voit also records many other early observations regarding urea output during fasting. 17Annalen der Chemie und Pharmacie (1843), 45, p. 244. 13 Das Hungern, Leipzig, 1890. "Wiener klin. Rundschau (1901), 15, pp. 69-71 and 91-93. * Centralbl. f. klin. Medicin (1889), 10, p. 651. 21 La Riforma Medica (1893), ix, 2, p. 542. "■ Schweiz. Wchschr. Pharm., 34, p. 395. ^Ztschr. f. exper. Path. u. Therapie (1905), 1, p. 419. ^Orvosi hetilap, Budapest (1894), p. 512. "Archiv f. path. Anatomie u. Physiol, u. f. klin. Medicin (1893), 131, Supple- mentheft 1-228. "Skan. Archiv f. Physiologie (1897), 7, p. 29. Introduction. 7 minations of the respiratory gases. Finally reference should be made to the study of fasting metabolism by Sadovyen28 in the Pashutin apparatus at St. Petersburg, in which not only the nitrogen but the carbon dioxide elimination was determined, the 1-day experiment of Likhachev27 in the Pashutin appa- ratus, which was so modified as to measure the heat elimination, and the obser- vations of van Hoogenhuyze and Verploegh ™ on the fasting girl Flora Tosca. PLAN AND PURPOSE OF THE EXPERIMENTS HERE REPORTED. The importance of the study of the fasting metabolism was early recognized in the experiments on the nutrition of man that have been in progress in this laboratory, and the results of three 24-hour experiments and one 48-hour experiment were reported.29 In these experiments the determinations included those of nitrogen, carbon dioxide, water, and heat. In the computation of the results of these experiments, the assumption was made, as is usual in metabolism experiments in which the determination of the oxygen intake is not made, that the amount of carbohydrate in the body remained unaltered. The error of this assumption, especially in experiments on fasting, was only too obvious, but not until provision was made for a direct determination of the oxygen consumed was it possible to secure any definite knowledge regarding the changes in the store of glycogen. The modified form of respiration calorimeter 30 makes it possible to deter- mine not only the metabolism of nitrogenous material, but also the carbon dioxide, water, and heat output and oxygen intake. With the new apparatus and increase in number of determinable factors, a series of experiments to study as completely as possible the metabolism in fasting men was planned. This investigation was made possible by liberal grants of the Carnegie Insti- tution of Washington. According to the plan of the experiments, the respiratory exchange and heat output were made the special subjects of study. Experiments during prolonged fasting were included to note the fluctuations in metabolism from day to day, and since the number of subjects available for long fasts was limited, a series of 2-day fasts with a number of men was included to eliminate the influence of individuality. The rapidity and amount of gain of nitrogenous material in the body consequent upon the ingestion of food after a prolonged fast was studied as a closely related supplementary problem. ^Trudi Russkavo obshtshestva okhraneniya Narodnavo Zdravia (1888), 12, pp. 13-76, St. Petersburg. 27 Dissertation, St. Petersburg (1893). »Zeit. f. physiol. Chemie (1905), 46, p. 440. 28 Experiments on the metabolism of matter and energy in the human body, 1900- 1902. W. O. Atwater & F. G. Benedict. Bull. 136, Office of Experiment Stations, U. S. Department of Agriculture (1903), Washington, D. C. 30 Carnegie Institution of Washington Publication 42 (1905). 8 Influence of Inanition on Metabolism. METHOD OF INVESTIGATION. In any study of metabolism, the greater the number of series of simultaneous observations on the same person, the more nearly does the study approximate completeness. The number of factors which may be determined is limited by the conditions surrounding an experiment, and while the use of the respiration calorimeter enables us to secure data heretofore not attainable, the nature of the apparatus is such as to preclude many physical and psychical obser- vations which might have been obtained on a subject living outside the chamber of the respiration apparatus. The observations made in these studies naturally divide themselves into four classes — grosser or general observations, and physiological, chemical, and physical measurements. The following sections enumerate the observations themselves and the methods employed in obtaining them. GROSSER OBSERVATIONS. In all of the instances of prolonged fasting which have been observed by scientists, certain grosser observations have been made which deal for the most part with the physical appearance, loss in weight, and general mental and physical condition. Such observations were likewise made in connection with the series of experiments here reported, although, as has been pointed out above, they were made, at times, with considerable difficulty. Body-weight. — By means of the special form of weighing apparatus described in an earlier report,31 accurate observations on body-weight were possible in all save the first experiment. Use is made of the fluctuation of the body- weight not only to note the condition of the subject but also as a check upon the accuracy of the determinations of intake and output. The total weight of income and the total weight of outgo being known, the difference between them may be determined and should represent body substance gained or lost; this could be readily checked by means of the weights recorded by the special apparatus for obtaining body-weight. Examination by physician. — In a study of fasting involving abstinence from food for a number of days, it was deemed important to have careful examinations made of the subject from time to time by a skilled physician; consequently it was arranged to have Dr. John E. Loveland, of Middletown, Connecticut, make careful examinations of all the subjects of these experi- ments, and also to examine the subjects of long experiments on each day of the experiments proper. The observations of the physician were for the most part confined to those obtainable through a glass window and by the telephone, although he could make use of the continuous record of body temperatures obtained during the experiments. Several attempts were made to determine 31 Carnegie Institution of Washington Publication 42 (1905). Method of Investigation. 9 the blood pressure by means of the sphygmomanometer. Since the obliteration of the radial pulse is necessary for the successful determination of the blood pressure, poor success attended its use. In order to enable the attending physician to note the character of the pulse, a provision was made for attach- ing a thin rubber diaphragm to the wrist of the physician and then inclosing this in the outer end of the food aperture in such a manner that no air could enter or escape. On opening the inner end of the food aperture, the subject could then place his hand in such a manner that the physician could obtain the radial pulse. This procedure, unfortunately, could not be used simulta- neously with the sphygmomanometer. It is further to be regretted that the technique of this operation was not completely developed before it was first used, and hence certain slight errors due to leakage of air were unavoidable on one or two occasions. By means of the analysis of the air in the chamber at the end of each day, however, it was possible to correct for the amount of the leak with great accuracy. PHYSIOLOGICAL MEASUREMENTS. The same difficulties which attended the making of grosser observations on the subject prevented a thorough series of physiological measurements, such as could be obtained with a subject moving freely about the laboratory. It is very much to be regretted that the observations actually made could not have been amplified and a greater amount of data secured. Fortunately, however, the study of Succi by Luciani was especially complete in observations of this nature, and inasmuch as Succi's fasting experiment lasted for 30 days, it is fair to assume that the data obtained by Luciani could not be greatly amplified by any data secured in an experiment of such short duration as 7 days. Measurements of body temperature. — In spite of the difficulties of making physiological observations on a subject inclosed in a respiration chamber, it was possible to secure the body temperature of the subject with great accuracy by means of a rectal thermometer, fully described elsewhere.32 By means of this thermometer, which is on the principle of a bolometer, the variations in electrical resistance of a fine platinum wire inclosed in a pure silver tube and inserted several centimeters in the rectum can be observed at will. While the majority of our subjects have experienced no difficulty in wearing this electrical resistance thermometer for a considerable period of time, the subject of the longer fasting experiments found this difficult and it was only during certain experimental days that he could use the thermometer and insure us normal results. It was deemed inadvisable to urge the use of the thermometer for fear of causing the subject discomfort which might produce abnormal results. In 32Archiv f. d. ges. Physiol. (1901), 88, p. 492; Carnegie Institution of Washing- ton Publication 42, p. 156 (1905). 10 INFLUENCE OF INANITION ON METABOLISM. a number of cases where the rectal thermometer was not worn, his temperature was taken, usually in the mouth, by means of a clinical thermometer. Pulse rate. — As a measure or an index of the degree of internal muscular activity, the pulse and respiration rates are of great value, and consequently in any series of observations which include total heat production it is desirable to have a continuous record, if possible, of the pulse rate and the respiration rate. In the longer fasting experiments here reported, the subjects made observations of their own pulse rate, sometimes as frequently as every half hour. Each subject was instructed to count the radial pulse for upwards of 2 minutes, noting the count during one minute and using the second minute as a check on the first count. Obviously this gave no pulse records during sleep, and this omission seemed so serious that an apparatus for indicating the pulse rate during the whole day was secured. Through the kindness of Prof. W. T. Porter a Fitz pneumograph83 was obtained. This pneumograph was adjusted around the chest of the subject and a stout-walled transmission tube used to connect the pneumograph with a glass tube in the inner door of the opening through which food is passed into the chamber. A flexible rubber tubing connected the tube in the inner door with one in the outer door of this compartment, and finally a rubber tube connected the tube in the outer door with a tambour. In spite of this long transmission, a distance of some 3 or 4 meters, the form of tambour used gave excellent results. Prof. W. B. Cannon, of the Harvard Medical School, kindly came to Middletown and assisted in adapting this apparatus to the conditions of experimentation. It was ascertained that when the pneumograph was adjusted carefully over the apex beat of the heart the pulse rate could be very accurately noted. In the earlier tests a curve was traced on smoked paper. It was found later that the curve was unnecessary, since the vibrations of the pointer of the tambour were so marked that the observer could count the respirations without difficulty. This apparatus served its purpose admirably and has resulted in our obtaining many observations of pulse rate during sleep that would have been otherwise unobtainable. During certain of the fasting days, and especially during certain periods of the experimental day, the strength of the pulse seemed to fall off to such an extent that the minor vibrations of the pointer, due to the transmission of the pulse waves, were so small as to be recognized with difficulty. Furthermore, special precautions were necessary to make sure that the pneumograph remained in approximately the position in which it was originally adjusted. For this purpose, two or three straps of canvas were placed over the shoulders to keep the pneumograph from slipping down. Each subject wore the pneumograph continuously throughout the whole "Journ. Experimental Medicine (1896), 1. Method of Investigation. 11 experiment and very little, if any, discomfort was experienced from having it about the chest, even during the hours of sleep. Respiration rate. — In addition to the work of circulation, the work of respi- ration forms a considerable portion of the total amount of internal muscular work and consequently any accurate data regarding the rate of respiration are of importance in all measurements involving energy transformation. Various attempts have been made previously in this laboratory to secure the rate of respiration of subjects during experiments. By the use of a stop watch the attempt has been made to count the respirations by looking through either the glass window in the front of the chamber or the glass doors of the food compartment. But little success attended these attempts, for the subject moved so frequently that it was impossible to count the risp and fall of the chest for any great length of time. Furthermore, the subject very soon became aware of the fact that the respiration rate was being counted, and the difficulties of counting the rate of respiration when the subject knows that the count is being made are only too well understood by physicians. The pneumograph is ideally adapted for giving the desired data. Indeed, it was extremely fortunate that with the same piece of apparatus two such valuable factors could be measured. When the curve of the respiration was drawn upon smoked paper, there was obviously a superimposed curve of a much smaller amplitude showing the pulse. Singularly enough, it was found that during the waking period, when the subject was more or less actively moving about, more difficulty was experienced in obtaining the respiration rate than the pulse rate; for while the slight vibration of the pointer for each pulse beat could, as a rule, be detected by the observer, the grosser vibrations due to the rise and fall of the chest were frequently masked by fluctuations due to body movements. This was particularly true during the period when meals were eaten. On the other hand, during the night and at times when the pulse rate became very feeble and almost impossible to recognize, it was practically always possible to obtain the respiration rate. Strength tests. — The importance of securing data regarding the effect of inanition on strength has been recognized heretofore. The observations made by the subject of the experiments of Johansson, Landergren, Sonden, and Tigerstedt,34 in which the subject noted the length of time he could suspend himself on the arm, were of much value. Unfortunately, with the subject of the prolonged fasts here reported, tests of strength were not included, as the multiplicity of other observations made it difficult to include these in the daily routine. In the later group of experiments, made on different subjects, a series of strength tests was made with the Tiemann hand dynamometer before, during, and after the close of the fast. Inasmuch as the element of fatigue is S4Skan. Archiv f. Physiol. (1897), 7, p. 31. 12 Influence of Inanition on Metabolism. also important, record was kept of all the individual tests. In some of the earlier experiments a trial was made first with the right hand, then with the left hand, again with the right, and then with the left, alternately, until five or six tests with each hand had been made. In the later series five tests were made with the right hand and five with the left. The data secured furnish evidence as to the rapidity of the onset of fatigue as affected by inanition. Blood examination. — Some of the earlier observations 3° indicated noticeable morphological changes in the composition of the blood, and it seemed desirable to obtain from the subject of these experiments such data as was possible. Accordingly, during some of the prolonged fasts, a number of blood exam- inations were made. The blood examination consisted of a study of smears and the determination of hemoglobin and the number of leucocytes and ery- throcytes per cubic millimeter. During the period before fasting and that after the subject left the calorimeter, no special difficulty was experienced in making these examinations. Since, however, in most of the long fasting experiments, the fasting concluded and the experiment with food began while the subject was still inside the respiration chamber, great difficulty was experienced in securing samples of blood for these tests. In order to obtain blood samples during the calorimeter experiments, a device similar to that arranged for the examination of the subject's pulse by the physician was used. This consisted of a thin sheet of rubber, through the center of which a small hole was cut. A thimble was inserted in this hole and the rubber diaphragm fastened to a frame made by removing the glass from an extra outer door to the food compartment. When this frame was closed, it made an air-tight closure. The subject was instructed to open the inner door of the food com- partment, insert his middle finger into the thimble, and then give a sudden thrust. In this manner the finger was accessible to the observer outside of the calorimeter and no air could enter, as the sheet rubber closed tightly about the finger. Obviously this procedure is open to the objection that more or less constriction is placed upon the finger and consequently there may be an abnormality in the blood taken in the sample. As a matter of fact, during the long fast it was extremely difficult to get any blood for the tests and, conse- quently, they are very incomplete. chemical measurements. The chemical measurements in connection with the metabolism experiments included the chemical analyses of the food, feces, urine, and the respiratory gases. The technique of these examinations has been given in detail elsewhere." It may be briefly referred to here. M Tauszk, loc. cit. so Carnegie Institution of Washington Publication No. 42 (1905); U. S. Dept. of Agr., Office of Experiment Stations, Buls. 44, 63, 69, 109, 136, and 175. Method of Investigation. 13 Food, feces, and urine. — Although the object of the experiments here reported was primarily the study of metabolism during fasting, there was opportunity to study the effect of fasting on metabolism when food was given after fasting ; hence the chemical examination of food was included in many of the studies here reported. The chemical determinations made in these experiments included water, ash, nitrogen, carbon, organic hydrogen, ether extract, sulphur, and phosphorus of food and feces. In addition to the above the examination of the urine included determinations of chlorine, phosphoric acid, sulphuric acid, creatine, and creatinine. Sampling and weighing for analyses. — In so far as possible, the determi- nations were made on fresh material. This was always the case with foods. With feces a preliminary drying was necessary. This drying was done in a vacuum desiccator. Urine was delivered from carefully calibrated pipettes. Water. — While formerly the water was determined by noting the loss in weight of the substances when heated at about 100° C, experience in this laboratory has shown that this method is open to criticism. Accordingly, water determinations in this series of experiments were made by the vacuum method.37 Ash. — Ash was determined by charring a sample of the dried material, extracting the charred mass with water to remove the more volatile salts, igniting the residue, and evaporating the extract with the residue at very low red heat. The determination of ash has by no means the scientific accuracy of the other determinations made in connection with these experiments, since the composition of the material weighed as ash may vary noticeably with the method of treatment. Arrangements could not be perfected in time to make the determinations of the calcium, magnesium, and potassium, but the crude ash determination is not without value. Nitrogen. — Nitrogen was determined by the Kjeldahl method, mercury being used to facilitate oxidation and potassium sulphide to destroy the mercur- ammonium compounds in the distillation. The distillation was carried out in a special form of still.38 The accuracy of the nitrogen determinations was very frequently checked by testing one or more materials of known nitrogen content, such as ammonium sulphate, ammonium ferrous sulphate, urea, and. uric acid. In such tests the digestion was carried out with the addition of 1 gram of pure sugar, so that the conditions might be identical with those of the regular analyses. Carbon and hydrogen. — The carbon was determined by the Liebig method, with such modifications as have been found to be practicable from extended "Benedict & Manning, Am. Jour. Physiol. (1905), 13, p. 309. 38 Jour. Amer. Chem. Soc. (1900), 22, p. 259. 14 Influence of Inanition on Metabolism. experience in this laboratory.30 In the later experiments, the determinations of the carbon in the urine were made by the electrical method of Morse.40 The accuracy of these methods of analysis was frequently tested by burning known substances, such as cane sugar or urea. Ether extract. — The ether extract (crude fat) of food was made by the usual method, by extracting 2 grams of the material (previously dried for the determination of water) in a paper filter pocket and weighing the crude extract thus removed. Feces receive special treatment in the analysis as discussed beyond. Phosphorus and phosphoric acid. — Total phosphorus in food, feces, and urine was determined by fusion with sodium peroxide and subsequent conversion to magnesium pyrophosphate. The modification suggested by Dubois,41 which includes the addition of a small amount of sodium carbonate to the fusion mixture, was found to be very satisfactory. Phosphoric acid in urine was determined, as is usual, by titration with uranium acetate, potassium ferrocyanide being used as an indicator. Sulphur and sulphuric acid. — The sulphur was determined by fusion with sodium peroxide and sodium carbonate in nickel capsules, precisely as in the case of phosphorus. The sulphuric acid in the fused mass was precipitated as a sulphate by barium chloride and water. The combined inorganic and ethereal sulphates in the urine were determined according to the method of Folin,43 which consists of heating urine with hydrochloric acid and potassium chlorate and precipitating the sulphate as barium sulphate. An attempt was made to obtain data regarding the neutral sulphur (S03) in urine by the difference between the sum of the inorganic and ethereal sulphates (S03) and the total sulphur (S03). The researches subsequently carried out by Folin 43 on the difficulties of obtaining the precipitates of barium sulphate may tend to lessen the value of the results obtained. Creatine and creatinine. — The significance of the excretion of creatine and creatinine, as pointed out by Folin,44 led to the determination of these com- pounds in the urine of the fasting subjects. The method of Folin for deter- mining creatine and creatinine has been employed in this laboratory. This depends upon the color reaction, noted by Jaffe, produced by the addition of a solution of creatinine to an alkaline solution of picric acid. The depth of color is compared with a half normal solution of potassium bichromate. From the standard color of the normal solution the amounts of creatinine in unknown solutions can readily be obtained. Chlorine was determined in urine by the Volhardt method. 89 Benedict, Elementary organic analysis, Easton, Pennsylvania, 1900. "Amer. Chem. Jour. (1905), 33, p. 591. "Jour. Amer. Chem. Soc. (1905), 27, p. 729. uAmer. Jour. Physiol. (1905), 13, p. 52. "Jour. Biol. Chem. (1906), 1, p. 131. "Amer. Jour. Physiol. (1905), 13, p. 83. Method of Investigation. 15 BESPIEATOBY PRODUCTS. For any study of the transformations of matter in the body, complete knowledge of the respiratory products is of the greatest importance. In the experiments here reported, determinations were made not only of the carbon dioxide and of the water-vapor eliminated through the lungs and skin, but likewise of the oxygen absorbed. The apparatus and methods have been described in detail in a previous publication." Water. — The determination of water, in brief, is as follows : The air-current which leaves the respiration chamber and which contains the water-vapor is caused to pass through a previously weighed vessel containing strong sulphuric acid. The water is retained by the acid and the increase in weight of the vessel indicates the amount of water absorbed during each experimental period, which is usually of from two to three hours' duration. Since there may be noticeable fluctuations in the amount of water in the residual air in the chamber, absolute determinations of the amounts of moisture in this air are made at the end of each period, the differences between the amounts at the beginning and end of each period giving data for correcting the absolute water elimination for that period. Carbon dioxide. — The determinations of carbon dioxide are made by con- ducting the air-current from the respiration chamber after it has been freed from water, through vessels filled with soda lime. The carbon dioxide is completely absorbed and provision is made for collecting and weighing the quantity of water lost from the reagent as the dried air passes through it. The soda-lime vessels, as well as the last sulphuric acid-containing vessel, are weighed at the beginning and end of each period. Fluctuations in the quantity of carbon dioxide residual in the air of the chamber are allowed for by analyses immediately after the end of each period and thus the actual carbon dioxide production can be computed accurately. Oxygen. — The determination of oxygen is somewhat more complex. This gas is inclosed in a highly compressed form in steel cylinders, from which it is admitted to the ventilating air-current. The loss in weight of the cylinder at the end of each period indicates the amount of oxygen admitted. Corrections are made for the slight amounts of nitrogen present in the gas. In addition to the amount of oxygen so admitted, however, there may be very marked fluctuations in the oxygen in the volume of air residual in the chamber, and, consequently, analyses of this residual air are also necessary to ascertain any alterations in the oxygen content. Actual determinations of oxygen from ab- sorption by potassium pyrogallate are made at the end of each day. A system of computation which includes the determination of the apparent volume of the air in the chamber at the end of the period, the amount of carbon dioxide in Carnegie Institution of Washington Publication No. 42 (1905). 16 Influence of Inanition on Metabolism. this air, the amount of water-vapor, and the amount of nitrogen enables the amount of oxygen at the end of each experimental period to be computed with great accuracy, and thus obviates the necessity of making the actual analyses with potassium pyrogallate at the end of each period. In general, the agreement between the computed oxygen content and that found by analysis is very satisfactory. The loss in weight of the cylinder, therefore, corrected for storage or loss cf oxygen from the air in the chamber, furnishes data for the computation of the oxygen absorption during any given experimental period. The measurements of respiratory gases in these experiments include the cutaneous respiration as well as that of the lungs. The nitrogen of perspiration is also determined in many instances. PHYSICAL MEASUREMENTS. The physical measurements incidental to experimenting on the transfor- mations of matter and energy in the body consist generally of two kinds, first, the measurement of the potential energy of the food, feces, and urine, and secondly, the measurement of the heat elimination from the body. In certain classes of experiments, where external muscular work is performed, measure- ments of the heat equivalent of external muscular work are made. The present series, however, consists entirely of rest experiments. Potential energy of food, feces, and urine. — As a result of many experi- ments of Eubner, Stohmann, Berthelot, and others, it is possible to compute with reasonable exactness the energy equivalent of food, feces, and urine. Improvements in the technique of the use of the bomb calorimeter, however, have been adopted in this laboratory, and consequently the potential energy of food, feces, and urine were, in every instance, actually measured by means of the calorimetric bomb,46 in that a dried portion of the material was burned in oxygen under a pressure of 20 atmospheres. In the fasting experiments the determinations were confined to those of the heats of combustion of the urines, and a great deal of experimental work has been done upon this subject. The most elaborate research on the heat of combustion of urine is that published by Farkas and Korbuly.47 Our experience has been that the most concordant results have been obtained by drying in vacuo 10 to 15 cubic centimeters of urine with 50 mg. of salicylic acid and burning the dried mass. The details of this investigation are not completed. They will be published elsewhere. Unquestionably, the heat of combustion of urine is low rather than high, since it is almost impossible to avoid the loss of ammonia or the conversion of urea into ammonium carbonate, both of which result in a loss of energy. The method outlined above gives results which, in our judg- 40Atwater & Snell, Jour. Amer. Chem. Soc. (1903), 25, p. 7. "Archiv f. ges. Physiol. 1904), 104, pp. 564-607. Method of Investigation. 17 ♦ ment, are at present the best that can be obtained, although it is well recog- nized by physiologists that the determination of the heat of combustion of urine is one of the most perplexing problems with which we have to deal. In preparing samples of food and feces for combustion, it is necessary first to partially dry the substance, as otherwise the combustion would be very un- satisfactory. In this partial drying, loss of organic matter, and consequent loss of potential energy, should be avoided. Our experiments with vacuum desiccators have again shown the wisdom of drying physiological preparations and food materials at room temperature in a vacuum rather than at a tempera- ture much above that of the body, i. e., the water-bath or steam-oven. Heat elimination from the tody. — The chamber of the respiration apparatus is provided with appliances for measuring heat. These appliances have been described in detail in a previous publication.48 The respiration chamber, which consists essentially of an air-tight copper box, is surrounded with alternate layers of air, zinc, and wood, so as to minimize the radiation. The heat pro- duced by the subject is brought away by a current of cold water flowing through a copper pipe, to which a large number of disks are soldered to increase the absorbing surface. Arrangements are made, by heating and cooling the air surrounding the chamber, to render the walls of the chamber adiabatic. By noting the rise of the temperature of the water in passing through the cham- ber, and the mass of the water, the amount of heat brought away by the water- current can readily be computed. The heat required to vaporize the water given off in the air-current is obtained by multiplying the number of grams of water vaporized by the factor 0.592. Complete tests of the respiration calorimeter. — Burning known weights of ethyl alcohol in the chamber furnishes a means of checking the accuracy of the respiration calorimeter for measuring water and carbon dioxide output and oxygen intake, as well as heat elimination. A large number of such check tests have given very satisfactory results. In general the quantities actually deter- mined differ from the calculated amounts by less than 1 per cent. RECORDING RESULTS AND THE USE OF DECIMALS. In the numerous computations involved in experiments of this nature it is frequently difficult to determine early in the calculation just what degree of refinement in the mathematical calculations is warranted by the accuracy of the experimental process from which the data are derived. Such decision has been withheld in every instance till as late as seemed necessary, in some in- stances undoubtedly too late, and hence the results are at times expressed with one or two decimal places that are not of real significance. It is to be noted, however, that in the final summary and balance tables supernumerary figures are in general omitted. 48 Carnegie Institution of Washington Publication No. 42 (1905). 2 18 Influence of Inanition on Metabolism. the experimental man. In computing the changes in body-weight, the moisture given ofE from the man, the changes in body temperature, and consequently fluctuations in total heat produced, and various other factors of a similar nature, theoretically, the man should be without clothing in the chamber; but this is neither prac- tical nor comfortable, and hence the body-weights were, in most instances, taken with the man plus union suit and stockings. This was considered our experimental man. If the body temperature fell 1°, it was assumed that the union suit and the stockings likewise fell 1°. The error seemed almost impossible to avoid, especially with the other discomforts attending on a fasting experiment. However, the exact statement of the conditions in which the weights and measurements were taken is here recorded. It should be said that the average weight of union suit plus stockings has been found to be 483 grams. PART 2. STATISTICS OF EXPERIMENTS. The experiments here reported consisted of studies in metabolism during inanition, both on the actual days of the fast and also on days following fast, when food was ingested. Furthermore, in order to throw light on the regain- ing of nitrogen lost from the body during fasting, two nitrogen metabolism experiments, lasting 25 and 14 days, respectively, were made. The fasting ex- periments lasted from 2 to 7 days. The experiments with food following fast were of from 1 to 3 days' duration. The subjects were, with the exception of S. A. B., all students in Wesleyan University, nine young men, in good health. S. A. B., with whom the longer experiments were made, was a young "masseur," who had made several fasts prior to his arrival in Middletown with a view of studying his daily losses in weight. A list of the experiments made in connection with this research is given in table 1. Table 1. — Duration and character of experiments. Metabolism experiment number. 59 68 69 70 71 72 73 74 75 76 Nitrogen metabolism No. 1 77 Nitrogen metabolism No. 2 79 80 81 82 83 85 89 Date. Dec. 18, 19, 20, 1903 Apr. 27, 28, 1904 Dec. 16, 17, 18, 19, 1904 Dec. 20, 21, 22, 1904 Jan. 7, 8, 9, 10, 1905 Jan. 11, 1905 Jan. 28, 29, 30, 31, Feb. 1, 1905. . Feb. 2, 3, 4, 1905 Mar. 4, 5, 6, 7, 8, 9, 10, 1905 Mar. 11, 12, 13, 1905 Mar. 14, 15, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, 29, 30, 31, Apr. 1, 2, 3, 4, 5, 6, 7, 1905 Apr. 8, 9, 10, 11, 1905 Apr. 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 1905 Oct. 13, 14, 1905 Oct. 27, 28, 1905 Nov. 21, 22, 1905 Nov. 24, 25, 1905 Dec. 5, 6, 1905 Dec. 9, 10, 1905 Jan. 10, 11, 1906 Subject. Dura- tion. Days. B. F. D. 3 A. L. L. 2 A. L. L. 4 A. L. L. 3 S. A. B. 4 S. A. B. 1 S. A. B. 5 S. A. B. 3 S. A. B. 7 8. A. B. 3 S. A. B. 25 S. A. B. 4 S. A. B. 14 H. E. S. 2 C. R. T. 2 A. H. M. 2 H. C. K. 2 H. R. D. 2 N. M. P. 2 D. W. 2 Character of experiment. Fasting. Do. Do. With food. Fasting. With food. Fasting. With food. Fasting. With food. Free selec- tion of food. Fasting. Free selec- tion of food. Fasting. Do. Do. Do. Do. Do. Do. 19 Metabolism Experiment No. 59. This experiment began December 18, 1903, and continued without inter- ruption for three days. The subject, B. F. D., was a student in Wesleyan University, of athletic temperament, and active in his movements. Since it is customary for each student at the university to have his body measurements taken, very complete records of the physical characteristics of the subject were available. The an- thropometric records are given herewith. Measurements of B. F. D. — Date, Oct. 20, 1903. Age, 22 years. Weight kilograms.. 67.3 Height centimeters . . 170.3 Length of — Sternum do 138 Navel do 98.7 Pubis do 80.7 Sitting do 94.1 Knees do 43.3 Shoulders, elbow do 34.3 Elbow to tip do 44.2 Arm reach do ... . 107.1 Right foot do 26 Left foot do 25.8 Girth of— Head do 55.2 Neck do 37.3 Chest- Depressed do ... . 85.4 Inflated do 94.5 Normal do ... . 91 At 9th rib, full do 85 At 9th rib, depressed .. do 77.8 Waist do 77.5 Hips do 96 Right biceps do 32.2 Left biceps do ... . 31.9 Right arm do 28.9 Girth of— Left arm centimeters Right elbow do . . Left elbow do . . Right forearm do . . Left forearm do . . Right wrist do . . Left wrist do . . Right thigh do . . Left thigh do . . Right knee do . . Left knee do . . Right calf do. . Left calf do.. Right instep do . . Left instep do . . Breadth of — Head do. . Neck do. . Shoulders do . . Chest do. . Waist do . . Hips do. . Depth of — Chest do. . Abdomen do . . 28.7 25 25 28.6 28.2 16.1 16.5 57 56 38 37.2 38 37 25.3 24.5 15.2 11.3 37 27.7 25.2 31 16.8 19.3 No especial preliminary preparation was made by the subject for this experiment, though in some of the later experiments the question of defecation during an experimental period was eliminated by the use of enemata for the removal of fecal matter. Moreover, although fasting experiments of one or two days' duration had previously been made in this laboratory, no regular routine for this type of experiment had been decided upon. The preparation of a regular program to be followed each day was therefore deemed inadvisable. It was definitely provided, however, that the subject was so to regulate his muscular movements as to have the same amount of activity, 20 Metabolism Experiment No. 59. 21 as nearly as possible, on all days of the experiment, and that the times for urinating, rising, weighing, and retiring should be regular. Since previous experience had shown that greater accuracy was obtained in the analyses of the respiratory gases if during the waking hours the subject took special precau- tions to remain as quiet as possible for the half hour preceding the end of each experimental period, he was cautioned to spend this last half hour of each period in sitting quietly in the chair, reading or writing. The subject entered the calorimeter chamber during the early evening of December 17. The bedding, urine bottles, feces can, and such books as the sub- ject wished, together with all the articles too large to pass through the food aperture, had been placed in the chamber before the large plate-glass window which closes it was sealed in place. In addition to a union suit of underwear, stockings, a light sweater, and trousers, the subject wore a pair of heavy-soled shoes primarily designed to prevent the conduction of heat from the soles of the feet to the metal floor of the chamber. Subsequent experience showed that these special shoes are un- necessary. After entering the respiration chamber, the subject adjusted the rectal ther- mometer and prepared for his three days' stay in the calorimeter by placing his bedding, books, and other articles in convenient positions on the shelves. Shortly before 11 p. m. he retired for the night, and at 1 a. m. the first pre- liminary period began. By this time the calorimeter chamber had reached temperature equilibrium and the preliminary air analyses had been made. The absorbing vessels used to purify the air were changed at the end of each experimental period, i. e., every 3 hours. The regular air analyses were made at 7 a. m. daily. Routine. — The general routine followed during this experiment was as fol- lows : At 7 a. m. the subject rose, dressed, and weighed himself on a platform scale inside the chamber, and then spent the rest of the day sitting or lying down. A considerable portion of each day was occupied with reading. Urine was collected every 3 hours until 10 p. m., after which it was not collected again until 7 a. m. the next day. The muscular movements were restricted to moving about the chamber, caring for the excreta, opening and closing the food aperture, telephoning, and dressing and undressing. Notes from diary. — No plans for definitely testing the effect of the stay in the respiration chamber on the mental or physical condition of the subjects had been made, since no detrimental results were anticipated. Yet it seemed desirable to obtain some notes concerning the psychical and physical condition of the men as recorded by themselves. For this reason each subject was in- structed to keep a diary, in which he was at liberty to record anything he chose 22 Influence of Inanition on Metabolism. The following are extracts from the diary of B. F. D. during this experiment. They include everything relevant to his physical and mental condition. Notes from diary. Dec. 17, 1903: Entered calorimeter at 10h35m p. m. Dec. 18, 1903: 7 a. m. Arose. During the night I did not sleep very well, though this morning I feel rested and quite re- freshed. My strange surroundings were the cause of my not sleeping well. Do not feel uncomfortable and am not hungry. 10 a. m. Feeling a little drowsy and have a slight pain in my head. llh55m a. m. Am feeling all right and think my head is a little better. lh50m p. m. Feeling better; my head is clearer. I could eat, but do not feel any distress. 2h55m p. m. My head is aching a little. 4h10m p. m. Am at present feeling very well; do not feel hungry and head feels better. 5b20m p. m. At present I am not at all hungry and feel better than any time during the day. 5h50m p. m. Attempted to defecate. 7b15m p. m. Am now feeling well and am not at all hungry. 9 p. m. Beginning to feel a little sleepy. Dec. 19, 1903: 7 a. m. Arose. 7h15m a. m. Slept very well last night and did not wake until 6h35m a. m. Do not feel hungry, and in general feel well, except that my head does not seem as clear as usual. 8h10m a. m. My head feels much better and clearer; feel so well that I have tried to sing. Not at all hungry. 9h50m a. m. Have been sitting since 7 and my head is tired; have been reading and studying. 10M0m a. m. Head now feels much better. llh45m a. m. Feel very well. lh5m p. m. My hands feel a little cold, though I am very comfortable and my head feels clear. 2h5m p. m. Am feeling well, except a little drowsy. 2h10m p. m. Have been thinking of some nice things to eat and find that my appetite responds with double force. 3 p. m. Do not feel like doing anything; my ambition seems to be leaving me. Otherwise I feel very well. My head is all right at present. 4h10m p. m. Feeling very well and quite rested. 6h25m p. m. Feeling well, but am tired of studying and am lonely. Sh27m p. m. Have been writing to pass away time and feel the better for doing it. Feel very well. Dec. 20, 1903: 7 a. m. Arose. 7h20m a. m. Slept pretty well during the night, but woke at 5 and stayed awake until 7; did not sleep as soundly nor as well as the night be- fore. (I want to say for yesterday that whenever I would study hard my head would begin to ache a little. Then when I would rest or do a little reading or writing for amusement my head would soon clear up and again feel fresh. This was especially noticeable when I studied geometry. Yesterday morn- ing, also, there was a slight taste in my mouth, not noticeable this morning.) 7h30m a. m. Am feeling well and strong, but rather dull. Sh5m a. m. Attempted to defecate. 9 a. m. Am feeling well; no pain in head. 10h8m a. m. Have been lying quiet for an hour; am feeling very well, about the same as though I were lying in bed in my room. 10h47m a. m. When I read of food my ap- petite is stimulated. llb32m a. m. Lying down and reading; feeling very well. Am hungry. 12h15m p. m. Hunger is not painful. 4h35m p. m. Feeling very well. 8h40m p. m. Feeling very well; have had no pain in my head to-day. Have felt better to-day than during the past two days. I0h34m p. m. Am feeling well, but slight- ly tired. Dec. 21, 1903: 7h5m a. m. Slept until 2h30m this morn- ing, when I awoke and remained awake until some time after 6; then slept until 7. When I awoke at 5 yesterday morning and at 2b30m this morning I was very warm, too warm to sleep. Am feeling good; my head is clear and I am not hungry. My mouth is a little dry. Metabolism Experiment No. 59. 23 NOTES TAKEN AFTEB EXPEBIMENT. When I came out of the calorimeter, at first I felt rather weak, but think this was caused more by being inactive for 3 days than from weakness. Walked to my room, washed, and then walked a short distance. Felt rather hungry. Ate 1 slice of toast and drank 1 glass of milk; it tasted very good. This made me feel better and stronger and I walked back to my room and shaved. My hands were not unsteady. Attended recitations at 8h10m a. m. and 9 a. m., but felt rather sleepy. At 10 o'clock ate 2 slices of bread and drank a small glass of milk; afterwards felt stronger. Attended recitations till 1 p. m. Felt rather sleepy. At 1 o'clock went to dinner and ate 3 slices of toast and drank 2 glasses of milk. After dinner, took a slow walk in the open air for over 2 hours. This, though it made my legs a little tired, seemed to refresh me and made me feel much better. Studied from 4h30ra until 6, when I went for supper. Was hungry and ate all that I wanted, including milk, toast, stewed meat, and 2 dishes of apple-sauce. After supper felt strong and well enough to study until 10h45m, when I stopped and took a hot bath and went to bed. Slept very soundly and very well for 8 hours; after that felt no effects of my fast and was strong and well. Body movements. — Although the subject endeavored to secure uniformity of body movements from day to day, a partial check upon his records of his muscular activity was attempted by securing a more or less complete record of grosser body movements as observed by the physical assistant sitting at the window in the front of the calorimeter. These movements were recorded, often very informally, on the sheets which are used for recording the thennometric observations from which the total heat is computed. These observations of grosser movements were supplemented by the record kept by the chemical assistant of the number of times the food aperture was opened and closed. A list of all recorded body movements follows. In it the imperative mood has been used for the sake of brevity. Movements of subject. — Duration, three days, from Bee. 18, 7 a. m., to Dec. 21, 7 a. m., 1903. December 18. A. M. p. M. A. M. 9h48m lie. lh00m move about. 7h04m rise, urinate, tele- 9 52 sit, telephone. 1 02 urinate. phone. 9 54 lie. 1 05 food aperture. 7 14 weigh. 10 04 rise, telephone. 1 28 read. 7 22 write. 10 06 move about, uri- 1 45 food aperture. 7 28 take pulse. nate, sit. 1 48 write. 7 32 read. 10 15 food aperture. 2 02 telephone. 7 36 telephone. 11 06 food aperture. 2 22 change position. 7 46 read. 11 18 lie. 2 28 read. 7 48 telephone. 11 48 restless. 2 32 stop reading. 7 54 telephone, drink. 11 50 sit. 3 04 lie, read. 8 00 food aperture.1 11 54 telephone. 3 22 stop reading. 8 12 rise. 11 56 stand. 3 28 sit, take pulse. 8 14 dress. 11 58 sit, telephone. 3 32 food aperture. 8 18 read. P. M. 3 34 restless. 9 14 telephone. 12h 10m food aperture, 3 38 read. 9 18 lie, read. read. 3 50 telephone. 1 This term is used in this and all similar tables hereafter without further remark to indicate that the subject goes to the food aperture, opens it, passes out whatever he wishes, takes in anything designed to be received, and closes the food aperture. 24 Influence of Inanition on Metabolism. Movements of subject. — Continued. December 18 (cont.) A. M. P. M. P. M. 4h05m food aperture. 8h 8 24m 26 write, drink. 7" 24™ 7 36 \ move about. 4 26 telephone. 8 28 write. 7 44 read. 4 36 food aperture. 8 34 lie. 8 14 write. 4 44 read. 8 40 sit. 9 02 food aperture. 5 44 undress. 8 44 write. 9 08 telephone, sit. 5 52 attempt to defe- 8 48 drink. 9 12 food aperture. cate. 9 08 write. 9 14 drink. 5 54 rise, move vigor- 9 14 stop writing. 9 16 read. ously. 9 22 read. 9 34 lie. 5 58 dress. 9 40 telephone. 10 00 rise, urinate. 6 08 lie, read. 9 44 read. 10 02 wralk. 6 28 write. 10 04 urinate, food aper- 10 06 sit, read. 6 46 lie. ture. 10 20 food aperture. 7 02 rise, urinate, tele- 10 06 lie, read. 10 42 telephone. phone. 10 22 stop reading. 10 54 move, stand. 7 04 move about. 10 34 rise, weigh. 10 56 undress, etc. 7 06 food aperture. 10 36 write. 11 00 retire. 7 08 sit. 10 44 read. December 20. 7 12 recline. 10 58 telephone. A. M. 7 20 read. 11 04 stop reading. 7h 00m rise, telephone. 7 54 telephone. 11 12 read. 7 01 urinate. 7 56 sit. 11 16 recline, read. 7 04 walk, dress. 8 00 move about. 11 20 sit. 7 10 weigh, telephone. 8 06 8 24 8 44 sit. 1 £ — 11 26 recline, read. 1 XV 7 12 lie. recline, read. p. 12" M. 02m food aperture. 7 15 7 16 food aperture, move about. 8 58 take pulse. 12 08 recline, read. 7 18 7 24 sit. write. 9 04 9 08 read. sit. 12 1 36 00 lie. rise. 9 12 read. 1 01 urinate. 7 34 drink. 9 52 recline. 1 08 lie. 7 46 take pulse. 10 00 rise. 2 08 sit. 7 48 stand. 10 02 urinate. 2 20 read. 7 50 food aperture. 10 04 sit, drink, read. 2 36 lie. 7 58 move about. 10 06 telephone, food ap- 2 40 read. 8 04 move vigorously. erture. 2 58 sit. 8 08 sit. 10 08 sit, read. 3 06 read. 8 10 write. 10 48 rise, open bed. 3 26 stop reading, take 8 16 read. 10 56 telephone. pulse. 8 26 recline, take pulse. 11 00 retire. 4 04 urinate, food aper- 8 34 read. December 19. ture. 9 00 sit, write. A. M. 4 06 sit. 9 06 recline. 7h 00™ rise. 4 08 food aperture. 10 04 sit, food aperture. 7 01 urinate. 4 12 read. 10 06 sit. 7 02 telephone, dress. 4 18 stand. 10 08 write. 7 10 weigh. J 22 sit. 10 12 recline, read. 7 14 sit, write. 4 A 26 read. 10 32 stop reading, rest 7 20 walk about, food aperture. 4 4 32 44 recline, take pulse. 10 36 on elbow, write, read. 7 22 fold bed. 4 52 telephone, food ap- XV OVJ 7 28 7 34 write, read. 5 04 erture. sit. P. M. 12h 10m move about. 7 38 write. 5 12 read. 12 18 telephone. 7 48 telephone. 5 44 recline. 12 22 sit at table. 7 52 read, write. 6 06 sit. 12 25 food aperture. 8 02 take pulse. 6 30 stand. 12 30 lie. 8 10 sit. 7 02 telephone, urinate. 1 04 rise, urinate, sit. 8 12 telephone, food ap- 7 04 food aperture. 1 06 write. erture. 7 12 stand. 1 14 move about. Metabolism Experiment No. 59. 25 Movements of subject. — Continued. December 20 (cont.) p. M. P. M. P. M. 4" 36m stand, food aper- 7b 44™ sit. lh32m write. ture. 8 04 food aperture. 1 34 telephone. 4 40 sit, read. 8 08 drink. 1 44 take pulse. 5 04 read. 8 10 rise, sit. 1 48 recline, read. 6 08 recline. 9 08 rise. 2 18 sit. 7 00 rise, telephone. 9 24 lie, read. 2 24 recline, read. 7 01 urinate. 10 02 sit, rise, urinate 2 58 take pulse. 7 02 walk, sit. 10 14 telephone. 3 24 sit, take pulse. 7 10 food aperture. 10 30 food aperture. 3 28 lie, read. 7 12 sit. 10 50 rise. 4 04 sit, read, write. 7 16 rise, sit. 10 54 undress. 4 05 food aperture. 7 40 rise. 11 00 retire. Pulse. — B. F. D. took his pulse throughout the experiment at intervals of from 30 to 45 minutes from 7 a. m. to 11 p. m. daily. The pulse was counted for 2 minutes and the average recorded in his diary. The records are tabulated below. Ptilse rate 0 f subject in metabolism experiment No. 59 (December, 1903). Rate. Rate. Time. Time. Dec. 18. Dec. 19. Dec . 20. Dec. 18. Dec. 19. Dec. 20. 7h20m a.m. . . . 61 "63 31>00n>p.m 56 63 66 7 30 64 . . . . 3 30 54 60 66 7 50 . . . . 64 4 10 59 61 67 8 05 . . 57 4 40 55 55 67 8 35 . . . . 53 5 20 56 , , 64 8 50 61 64 5 45 . . 60 9 00 . . 64 6 00 53 68 9 25 57 . . . . 6 25 . 9 60 , . 9 40 , . 64 . . 6 40 58 t t 10 05 55 , . 61 7 05 54 56 64 10 45 55 63 61 7 45 , # 63 64 11 15 52 56 , , 8 00 55 , . , , 11 32 , , 65 8 27 57 , , 11 45 55 57 # . 8 40 65 12h15mp.m. . . . , . . . 62 9 10 46 63 56 12 30 52 , . . , 9 40 47 55 , , 1 05 55 54 61 10 05 , , 50 51 1 50 57 62 66 10 34 50 , , 59 2 20 55 65 10 51 55 1 At 7 a.m. Dec. 21, pulse=67. Drinking-water. — No attempt was made in the experiments here reported to insist upon complete abstinence by the subject, and consequently drinking-water was allowed whenever desired. A bottle filled with water and previously weighed was passed to the subject through the food aperture. The temperature of the water was taken immediately before it entered the chamber. The amount of water consumed was determined in all cases by deducting from the weight of the bottle plus water at the time it entered the calorimeter the weight of the bottle plus water when it left the chamber. This method takes account of the 26 Influence of Inanition on Metabolism. small quantities that are necessarily vaporized from the mouth and neck of the bottle. However, for the purpose of heat measurement it is immaterial in this form of calorimeter whether a gram of water is vaporized in the body of the subject or from the neck of the bottle. In this experiment it is not surely known at what definite times water was drunk, or the actual amount consumed at any given time, but the total amount for each 24 hours is known, and from the above data the periods during which the water was consumed and the amount of water have been estimated. Since, as will be seen later, in computing the total heat production for any given period, the amount of drinking-water consumed during that period is a not unimportant factor, these estimates of this amount were necessary. Save on the first day, no water was consumed after the subject retired and the amounts consumed were not far from the same each day, ranging from a maximum of 1360.2 grams on the second day to 1188.0 grams on the last day. The water furnished was that from the city supply and no allowance was made for salts or organic matter. The analyses of the water as furnished by the State Bacteriological Laboratory shows in parts per million 56 of total solids, 0.028 free and 0.278 of albuminoid ammonia, and 1.90 of chlorine. The amount of water consumed per period and the total amount per day are given in table 2. Table 2. — Record of water consumed — Metabolism experiment No. 59. Date. Period during which water was consumed. Total for day. 7 to 10 a.m. 10 a.m. to 1 p. m. i to 7 p. m. 7 to 10 p.m. 10 p. m. to 1 a.m. 1903. Dec. 18-19 Dec. 19-20 Dec. 20-21 , . Grams. 400.0 400.0 400.0 Grams. 400.0 400.0 388.0 Grama. 400.0 390.7 Grams. 169.5 400.0 Grams. 142.5 Grams. 1342.5 1360.2 1188.0 Ueine. In order to show the rate of elimination of nitrogenous material during each 24 hours of the fast, the urine was collected in 3-hour periods from 7 a. m. to 10 p. m. It was deemed inadvisable to awaken the subject to collect the urine in 3-hour periods during the night, as previous experience had shown that the subject found it difficult to go to sleep again. The weight in grams, the specific gravity, the reaction, and the total nitrogen (obtained by the Kjeldahl process) of the urine were determined for the different periods. An aliquot of the urine from each period was taken to make a composite sample for the day and a further aliquot of each day's composite sample was combined to make a 3-day composite. In the daily composite samples, determinations of nitrogen and heat of combustion were made, while in the 3-day composite Metabolism Experiment No. 59. 27 the determinations consisted not only of nitrogen and heat of combustion, but also of water, carbon, hydrogen of organic matter, and ash. The results of all determinations by periods are given in table 3. Table 3. — Determinations in urine per period and per day — Metabolism experiment No. 59. Date. Period. (a) Amount. (6) Specific gravity. to) Vol- ume. (a-s-b) (d) Reaction. Nitrogen. •to) Pro- por- tion. (/) Am'nt. (axe) 1903. Dec. 18-19. Dec. 19-20. Dec. 20-31. 7 a.m. to 10 a.m. 10 a.m. 1 p.m. 1p.m. 4 p.m. 4 p.m. 7p.m. 7p.m. 10p.m. 10 p.m. 7 a.m. Total Gh-ams. 190.2 252.2 528.7 129.3 203.7 236.7 1.0230 1.0165 1.0070 1 . 0320 1.0130 1.0240 c.c. 186 248 535 137 301 330 Slightly alkaline. . Slightly alkaline. . Slightly acid Acid Per ct. 1.00 .67 .45 .65 .74 1.49 Grams. 1.90 1.69 2.38 .84 1.51 3.52 do 1540.8 .... 1517 11.84 Total by com- 7 a.m. to 10 a.m. 10 a.m. 1p.m. 1p.m. 4 p.m. 4p.m. 7p.m. 7p.m. 10p.m. 10p.m. 7 a.m. Total 1540.8 1.0160 1517 .77 11.86 Acid 87.2 106.2 114.8 175.5 96.2 401.9 1.0385 1.0370 1.0340 1.0225 1.0250 1.0180 85 103 113 173 94 396 1.78 1.71 1.69 1.38 1.71 1.32 1.55 1.82 1.94 2.35 1.65 4.90 . . . .do . . . .do 981.8 .... 963 14.11 Total by com- 7 a.m. to 10 a.m. 10 a.m. 1p.m. 1p.m. 4p.m. 4p.m. 7p.m. 7p.m. 10p.m. 10 p.m. 7 a.m. Total 981.8 1 .0210 963 1.43 14.04 Acid 111.1 146.8 190.2 114.0 100.4 266.4 1.0270 1.0320 1.0190 1.0345 1.0280 1.0240 108 144 187 111 98 360 1.90 1.54 1.21 1.64 1.91 1.64 2.11 2.36 2.30 1.87 1.92 4.36 . . . .do . . . .do 928.9 .... 908 14.83 Total by corn- Total, 3 days.. Total compos- ite, 3 days. . . 928.9 1.0230 908 1.59 14.77 3451.5 .... 3387 40.77 3451.5 1.0200 3387 1.18 40.73 The accuracy of the nitrogen determinations and the method of taking composite samples is checked by the agreement between the sum of the amounts of nitrogen in the samples for periods and in the composite samples. Thus, on the first day the total amount of nitrogen eliminated, as determined from the total composite sample, was 11.86 grams and the total amount of nitrogen as determined from the 6 periods was 11.84 grams. The nitrogen in the total composite for the 3 days was 40.73 grams, while the nitrogen eliminated for 28 Influence of Inanition on Metabolism. the sum of the 18 periods was 40.77 grams. Similarly, the determinations of the heat of combustion on the 3-day composite showed 314 calories, while the determination of the daily composites showed that the urine contained 308 calories of energy. In addition to the data in table 3. the total 3-day composite yielded 95.4 per cent of water, 0.87 per cent of carbon, 0.26 per cent of hydrogen in organic matter, and 0.69 per cent of ash. The heat of combustion of the urine per grm was 0.059, 0.108, and 0.119 calorie, respectively, and the total energy of the urine for the 3 clays was 91, 106, and 111 calories, respectively. Weight and composition of urine. — The total amounts of nitrogen, carbon, hydrogen in organic matter, oxygen, water, solids, ash, and organic matter excreted per day are shown in table 4. The records given in table 3 show that only the determinations of nitrogen and heat of combustion were made daily, the determinations for water, ash, carbon, and hydrogen being made upon the 3-day composite. In accordance with our previous custom, the amounts of carbon and hydrogen were apportioned between the 3 days according to the amounts of nitrogen excreted, it being assumed that a definite amount of nitrogen will carry with it a proportionate amount of carbon and hydrogen in organic matter. Method of proportioning ash of urine. — Since in this experiment the analy- sis of ash was made only on the 3-day composite of the urine, it became neces- sary, in order to obtain the material katabolized from the body for each indi- vidual day, to apportion this ash among the different days of the experiment. The results of subsequent experiments in which the total solids, ash, and nitrogen were determined daily showed that in fasting experiments the ratio of the nitrogen to the total solids is fairly constant, and that on the average N = total solids X 0.29. This percentage was applied to the nitrogen found in experiment No. 59, and the following results for total solids were obtained : For the first day, 40.9 grams; for the second day, 48.4 grams; and for the third day, 50.9 grams — a total of 140.2 grams. The actual total solids were determined in the 3-day composite sample and found to be 158.8 grams, a result 18.6 grams greater than the amount computed for the 3 days. For lack of a better method of apportionment, this discrepancy was distributed over the 3 days in proportion to the several amounts of solids. The estimated amount of total solids for the first day (40.9 grams) was therefore increased by a certain proportion of the total discrepancy corresponding to the fraction ~| X (158.8 — 140.2). Apportioning this error, 18.6 grams, over the 3 days by the method indicated above, the corrected total solids are 46.3, 54.8, and 57.7 grams. They are recorded in line c of table 4. The total solids consist of ash, urea, and other compounds which may be Metabolism Experiment No. 59. 29 designated " material other than urea." The urea may roughly be computed * by dividing the number of grams of nitrogen eliminated by 46.6, the percentage of nitrogen in urea. For the 3 days of this experiment it was computed to be 25.43, 30.15, and 31.70 grams for the respective days. Given, then, the total solids of urine and the number of grams of urea (obtained by dividing the nitrogen by 46.6), by subtracting from the total solids of urine the number of grams of urea, the " material other than urea plus ash " is obtained. Thus on the first day of the experiment the amount of total solids was computed to be 46.3 grams. Deducting from this the amount of urea for the first day, i. e., 25.43 grams, leaves 20.87 grams as the weight of "material other than urea plus ash." The corresponding figures for the second and third days are 24.65 grams and 26.00 grams. Assuming, then, that the " material other than urea " is constant, the ash is obtained as follows : Total Table 4. — Weight and composition of urine — Metabolism experiment No. 59 (December, 1903). Weight Water Solids, a—b Ash i Organic matter, c—d (/) Nitrogen (g) Carbon (h) Hydrogen in organic matter (i) Oxygen (by difference) in organic mat- ter, e-(f+g + h) (a) (6) (c) (d) ie) Dec. 18-19. Grams. 1540.8 1494.5 46.3 6.94 39.36 11.84 8.71 2.60 16.21 Dec. 19-20. Grams. 981.8 927.0 54.8 8.20 46.60 14.11 10.40 3.11 18.98 Dec. 20-21. Grams. 928.9 871.2 57.7 8.67 49.03 14.82 10.92 3.26 20.03 Total for 3 days. Grams. 3451.50 3292.7 158.8 23.81 134.99 40.77 30.03 8.97 55.22 1 The ash and water for the individual days are calculated as shown on p. 28. Hence the amounts of solids and organic matter for the individual days are not determined but calculated. " material other than urea plus ash " for 3 days is to the " material other than urea plus ash " for any given day as the total ash is to the ash for any given day. Thus, the sum of the " material other than urea plus ash " for the 3 days is 20.87 -f- 24.65 + 26.00 = 71.52 grams. The proportion, then, would be 71.52 : 20.87 : : 23.81 (total ash determined for the three days of the experiment) : 6.94 grains, the amount of ash for the first day. By means of this proportion the values for ash for the two following days were found to be 8.20 and 8.67 grams, respectively. Method of obtaining water in urine in experiment No. 59. — The amount of water of the urine was determined on the 3-day composite in this experiment. In order to compute the amounts excreted each day, the figures for the total amount of urine for each day were decreased by the corrected amount of total 1 It is known that probably at least 1 gram of creatinine is excreted aside from the urea. Hence the calculation is at best but approximate. 30 Influence of Inanition on Metabolism. solids found by the method explained above. Thus 1540.80 (the amount of urine for the first day) — 46.3 = 1494.50, the water for the first day. Corre- sponding amounts for the other two days of the experiment were 927.00 grams and 871.20 grams. Elimination of Water- Vapoe. The amounts of water-vapor in the ventilating air-current are of value to show the loss of water from the body through the lungs and skin, to interpret the hygrometric conditions, and to aid in computing the heat production. The data regarding the water in the ventilating air-current are given in detail in table 5. Column a of the table shows the relative humidity of the air inside the chamber. These data are important, since it is conceivable that there may be noticeable difference? in the gaseous exchange and heat radiation as the result of differences in hygrometric conditions. The method of obtaining the figures is as follows : Knowing the total volume of air in the chamber to be approximately 4900 liters and the temperature about 20° C. and that 1000 liters of air when saturated at 20° contain 17.11S grams of water, it is possible to compute the amount of water-vapor which would be present in the air of the chamber if it were completely saturated at 20°. This amount would be 83.88 grams. From the ratio between this saturation amount and the actual amount of water found, the relative humidity is readily obtained. In pre- senting the results of subsequent experiments in this report the relative humidity will not be given, but the weight of water-vapor in grams at the end of each period, as shown in column b of table 5, will be included in the tables. By following the method outlined above, the relative humidity can by simple computation be readily obtained. In column c is recorded the gain or loss of water-vapor by the air in the chamber from period to period. At the end of each experimental period, the determination of the amount of moisture in the air is made by weighing the water absorbed from a known amount of air in its passage through a U-tube containing pumice stone drenched with sulphuric acid. The total amounts of water-vapor remaining in the chamber as computed from this determination are recorded in column b. The total water of respiration and perspiration, i. e., the amounts retained in the sulphuric acid absorber and recorded in column d corrected for the variations in the amount of water-vapor inside the chamber, are recorded in the last column. In subsequent experiments, only the figures recorded in column b and column e in this table will be presented. In certain experiments other corrections are necessary for the amount of water condensed by the heat- absorbing system and for the differences in weight of the underclothing, chair, bed, bedding, etc. In the experiment here reported, however, no perceptible moisture was observed on the absorbers in the chamber, and the data for fluctuations in the weight of the bed, bedding, and clothing were not obtained. Metabolism Experiment No. 59. 31 A word regarding the relative humidity of the air inside the chamber should be said here. The data of table 5 show that, on the whole, the relative humidity falls off somewhat as the experiment progresses, averaging not far from 45 per cent during the 3 days. During work experiments, the humidity increases Table 5. — Record of water of respiration and perspiration — Metaoolism experiment No. 59. Date. Period. Water-vapor in chamber at end of period. (c) Gain(+)or loss (— ) from preceding period. (d) Amount in outgoing air. (e) Total water of respira- tion and perspira- tion. (c+c?) (a) Relative humidity. (W Amount. 1903. Dec. 18 Dec. 18-19... Dec. 19-20... Dec. 20-21... Per ct. 54.8 48.8 49.1 Grams. 45.88 40.86 41.11 Grams. — 5.02 + 0.25 Grams. 130.63 132.95 Grams. 125.61 133.20 1 a.m. to 4 a.m. . 4 a.m. 7 a.m. . 7 a.m. to 10 a.m. . 10 a.m. 1 p.m. . 1 p.m. 4 p.m. . 4 p.m. 7 p.m. . 7 p.m. 10 p.m. . 10 p.m. 1 a.m. . 1 a.m. 4 a.m. . 4 a.m. 7 a.m. . Total 51.7 49.0 47.2 48.2 45.8 43.2 44.9 46.0 43.25 41.05 39.55 40.38 38.37 36.19 37.55 38.54 + 2.14 — 2.20 — 1.50 + 0.83 -2.01 -2.18 + 1.36 + 0.99 134.27 132.21 121.14 132.91 116.35 119.05 114.05 114.91 136.41 130.01 119.64 133.74 114.34 116.87 115.41 115.90 — 2.57 984 . 89 982.32 7 a.m. to 10 a.m. . 10 a.m. 1 p.m. . 1 p.m. 4 p.m. . 4 p.m. 7 p.m. . 7 p.m. 10 p.m. . 10 p.m. 1 a.m. . 1 a.m. 4 a.m. . 4 a.m. 7 a.m. . Total 48.3 44.2 44.0 40.5 44.5 42.2 42.6 49.2 40.39 37.01 36.84 33.87 37.27 35.32 35.66 41.19 + 1.85 -3.38 -0.17 -2.97 + 3.40 -1.95 + 0.34 + 5.53 122.62 125.35 115.15 123.88 118.28 117.10 111.19 115.73 124.47 121.97 114.98 120.91 121.68 115.15 111.53 121.26 .... .... + 2.65 949.30 951.95 7 a.m. to 10 a.m. . 10 a.m. 1 p.m. . 1 p.m. 4 p.m. . 4 p.m. 7 p.m. . 7 p.m. 10 p.m. . 10 p.m. 1 a.m. . 1 a.m. 4 a.m. . 4 a.m. 7 a.m. . Total 42.6 43.8 42.8 43.7 43.1 40.7 46.5 44.7 35.71 36.70 35.81 36.57 36.04 34.05 38.94 37.45 — 5.48 +0.99 -0.89 + 0.76 -0.53 -1.99 + 4.89 — 1.49 125.08 119.63 120.23 113.35 119.96 113.24 117.09 118.31 119.60 120.62 119.34 114.11 119.43 111.25 121.98 116.82 -3.74 946.89 943.15 considerably. It is important to note that during this experiment the air was relatively dry. In the fasting experiments there was no sensible perspiration observed by any of the subjects. ElJMINATION OF CARBON DIOXIDE. Carbon dioxide is one of the most important products of oxidation in the body. Hence determinations of this factor were made in all the experiments 32 Influence of Inanition on Metabolism. here reported. The details of the determinations for the different periods of this experiment are given in table 6. The ventilation conditions are here considered in detail, since the air is distinctly abnormal so far as its carbon Table 6. — Record of carbon dioxide- —Metabolism experiment No. 59. Date. Period. Carbon dioxide. (ff) Carbon in carbon dioxide exhaled (ex3/ll) In chamber at end of period. (c) Gain (+) orloss(-) from preced- ing period. (d) Am'nt ab- sorbed from out- coming air. (e) Cor- rected weight exhaled by subject (c+d). (/) Volume exhaled by subject (exO.6091) (a) Parts in 10,000. CM Am'nt. 1903. Dec. 18 1 a.m. to 4 a.m. 36 28 Grams. 33.37 25.82 Grams. — 7.55 Grams. 77.23 Grams. 69.68 Liters. 35.47 Grams. 19.00 Dec. 18-19.. 4 a.m. 7 a.m. 7 a.m. 10 a.m. 35 32.39 + 6.57 72.39 78.96 40.20 21.54 44 40.72 + 8.33 93.43 101.76 51.80 27.75 10 a.m. 1 p.m. 56 51.29 + 10.57 78.25 88.82 45.24 24.22 1 p.m. 4 p.m. 49 44.99 - 6.30 94.59 88.29 44.95 24.08 4 p.m. 7 p.m. 88 81.72 + 36.73 61.85 98.58 50.19 26.89 7 p.m. 10 p.m. 63 58.79 — 22.93 104.63 81.70 41.59 22.28 10 p.m. 1 a.m. 41 38.43 —20.36 94.48 74.12 37.74 20.21 1 a.m. 4 a.m. 26 24.60 — 13.83 79.12 65.29 33.24 17.80 Dec. 19-20.. 4 a.m. 7 a.m. Total 7 a.m. to 10 a.m. 45 42.12 + 17.52 54.47 71.99 36.65 19.63 .... + 9.73 660.82 670.55 341.40 182.86 41 38.11 — 4.01 98.97 94.96 48.35 25.90 10 a.m. 1 p.m. 32 29.80 — 8.31 94.24 85.93 43.75 23.43 1p.m. 4 p.m. 38 34.84 + 5.04 78.31 83.35 42.43 22.73 4 p.m. 7 p.m. 37 33.83 - 1.01 89.46 88.45 45.03 24.12 7 p.m. 10 p.m. 42 38.85 + 5.02 85.72 90.74 46.19 24.75 10 p.m. 1 a.m. 31 28.80 -10.05 82.13 72.08 36 69 19.66 1 a.m. 4 a.m. 25 22.77 - 6.03 71.91 65.88 33.54 17.97 Dec. 20-21.. 4 a.m. 7 a.m. Total ... 7 a.m. tolO a.m. 42 38.39 + 15.62 — 3.73 61.93 77.55 39.48 21.15 .... 662.67 658.94 335.46 179.71 34 30.48 — 7.91 103.76 95.85 48.80 26.14 10 a.m. 1 p.m. 46 41.98 + 11.50 72 . 05 83.55 42 . 53 22.79 1 p.m. 4 p.m. 35 31.39 — 10.59 96.34 85.75 43.66 23.38 4 p.m. 7 p.m. 51 45.71 + 14.32 69.88 84.20 42.87 22.96 7 p.m. 10 p.m. 36 31.84 — 13.87 100.24 86.37 43.97 23.55 10 p.m. 1 a.m. 44 38.89 + 7.05 67.07 74.12 37.73 20.21 1 a.m. 4 a.m. 37 32.83 -6.06 79.33 73.27 37.30 19.98 4 a.m. 7 a.m. Total 46 41.73 + 8.90 58.23 67.13 650.24 34.18 18.31 + 3.34 646.90 331.04 177.32 dioxide content is concerned. In the first column the carbon dioxide is expressed in parts per 10,000. With variations in muscular activity there are variations in the amounts of carbon dioxide in the chamber at the end of different periods. These variations, expressed as gains or losses, are given in column c. The major Metabolism Experiment No. 59. 33 portion of the carbon dioxide in the ventilating air-current is absorbed in the soda-lime cans or purifiers. The amount so absorbed is recorded in column d. The amount exhaled by the subject, i. e., the amount actually pro- duced during any period, is the algebraic sum of columns c and d. These amounts are recorded in column e. Since the volumes of the carbon dioxide are used in computing the respiratory quotients, they are calculated by multi- plying the figures in column c by the factor 0.5091 and the results are recorded in column /. Column g records the amounts of carbon in carbon dioxide exhaled. These values are used in subsequent calculations affecting the gain or loss of chemical elements in the body. When it is remembered that normal air contains from 3 to 4 parts of carbon dioxide per 10,000, the data in column a of the table appear of peculiar interest. The results serve to show that the carbon dioxide content of the air in this experiment was iu general from 8 to 10 times that of normal air, and in one instance, at 7 p. m., December 18, it reached more than 20 times the normal. The amount of carbon dioxide remaining in the chamber at the end of each period is determined simultaneously with the amount of water by absorbing the carbon dioxide from the known volume of air passed through soda lime. The ratio of the total volume of the sample to the volume of air in the calorimeter chamber being known, the amount of residual carbon dioxide can be computed. The amount of carbon dioxide, in parts per 10,000, can in all cases be roughly approximated by multiplying the residual amount in grams of carbon dioxide by 1.12, hence in subsequent experiments the ventilation conditions will not be expressly tabulated. Oxygen Consumed. The role played by oxygen in katabolism is a complex one, in that all three components of the body, namely, protein, fat, and glycogen, are oxidized in the process of katabolism. The amount of oxygen absorbed during the oxi- dation is of great importance in interpreting the kinds and amounts of body materials oxidized. While in the case of carbohydrates oxygen is used to oxidize only carbon, since hydrogen and oxygen in the carbohydrate molecule exist in the proportion to form water, with fat and protein, on the other hand there is oxidation of both carbon and hydrogen. Hence the ratio of the oxygen consumed to the carbon dioxide exhaled is markedly different with the three groups of compounds. For the proper interpretation, therefore, of the specific katabolism, a knowl- edge of the amount of oxygen absorbed is essential. In these experiments the amount thus absorbed is determined by noting the loss in weight of a steel cylinder from which oxygen is admitted to the ventilating system from time to time, and also any drafts upon the residual amount of oxygen in the air 3 34 Influence of Inanition on Metabolism. of the chamber. There are about 1000 liters of oxygen in the air of the chamber at the beginning of the experiment. The subject can draw upon this residual for oxygen to support the vital processes and these drafts would of necessity result in a difference in the composition of the air and the residual amounts of the elements in it. Furthermore, the supply of oxygen admitted may at times be larger than that actually consumed by the subject and hence the residual amount of oxygen may increase. A reference to the original description of the methods and appliances for the methods of determining oxygen2 will show that the amount of oxygen supplied varies with the barometric pressure. The admission of oxygen is determined by the height of the rubber diaphragms on the two pans which equalize the pressure of the air. If the barometer falls, the air in the chamber expands, thus causing the rubber diaphragms to rise, consequently decreasing the admission of oxygen during this period. The oxygen is being consumed out of the air and a minimum amount is being supplied. Thus there is a draft upon the residual quantity of oxygen. If the barometer rises the air inside the chamber is compressed and diminishes in volume, resulting in a lowering of the diaphragm on the pans. It becomes necessary, therefore, to so increase the supply of oxygen as to keep the rubber diaphragm from touching the bottom of the pans and thus eliminate any chance for a rarefaction inside the chamber. Under these conditions the oxygen supplied to the chamber may be much greater than that actually required by the subject and the residual amount be thereby considerably increased. The determinations of the actual amounts of oxygen admitted to the system and the variations in the residual amounts are recorded in table 7. Column a records the per cent of oxygen in the air in the calorimeter chamber at the end of each period, and the actual amount expressed in liters is tabulated in column b. These amounts were obtained as a result of the analysis of air at 7 o'clock each morning and the computation of the amount of oxygen, which takes into consideration the amount of water-vapor and carbon dioxide as determined by the residual analysis, the amount of nitrogen added with the oxygen, and the total air of the chamber. The only factor that is not shown is the true volume of air inside the apparatus at the end of each period. While this volume is in general not far from 4890 liters, there may be slight fluctu- ations in volume, due to the size of the subject and number of miscellaneous articles inside the chamber. From these data the amounts of oxygen remaining in the chamber at the end of each period were readily computed.3 During the period from 4 to 7 a. m. on December 18, the preliminary night, the increase in the amount of oxygen residual in the chamber amounted to 2 Carnegie Institution of Washington Publication No. 42. 3 For a detailed explanation of the method of computation, see U. S. Dept. Agr., Office of Exp. Sta. Bui. 175. Metabolism Experiment No. 59. 35 9.80 liters. On the contrary, during the period from 4 to 7 a. m. on the experimental day, December 19-20, there was a decrease of 39.10 liters in the amount of oxygen in the chamber. These and similar fluctuations in the residual amount of oxygen are recorded in column c. Table 7. — Record of oxygen — Metabolism experiment No. 59. Date. Period. Oxygen. (a) Volume of oxygen con- sumed (/X0.7). In chamber at end of period. Gain (+) or loss (— ) from preceding period. («) Am'nt ad- mitted to cham- ber. (/) Cor- rected am'nt con- sumed by subject (e-d). (a) Pro- por- tion. (6) Am'nt. (c) Volume. (d) Weight (c-s-0.7). 1903. Dec. 18 P.ct. Preliminary 20 . 6 1 a.m to 4 a.m. 20.9 Liters. 958.56 979.32 Liters. + 20.76 Grams. + 29^66 Grams. 88^3 Grams. 58^87 Liters. 41.21 Dec. 18-19.. 4 a.m. 7 a.m. 21.1 989.12 + 9.80 + 14.00 84.44 70.44 49.31 7 a.m. to 10 a.m. 21.1 992.971 + 3.85 + 5.50 100.83 95.33 66.73 10 a.m. 1 p.m. 21.0 985.301- 7.67 — 10.96 68.70 79.66 55.76 1 p.m. 4 p.m. 21.3 1002.80 + 17.50 + 25.00 99.64 74.64 52.25 4p.m. 7p.m. 21.1 998.20 — 4.60 - 6.57 90.03 96.60 67.63 7 p.m. 10 p.m. 20.9 991.09 — 7.11 - 10.16 75.48 85.64 59.95 10 p.m. 1 a.m. 21.4 1017.27 + 26.18 + 37.40 106.62 69.22 48.46 1 a.m. 4 a.m. 21.5 1019.72 + 2.45 + 3.50 66.85 63.35 44.35 Dec. 19-20.. 4 a.m. 7 a.m. Total 21.3 1010.26 — 9.46 - 13.52 51.46 64.98 45.49 .... + 21 14 -4- SO 19 659.61 629.42 440.62 7 a.m. to 10 a.m. 21.1 1003.45J— 6.81 — 9.73 92.79 102.52 71.76 10 a.m. 1 p.m. 21.1 1000.071— 3.38 — 4.83 67.69 72 . 52 50.76 1 p.m. 4 p.m. 20.9 988.56- 11.51 - 16.44 61.29 77.73 54.41 4 p.m. 7 p.m. 20.8 982.46 — 6.10 - 8.71 71.46 80.17 56.12 7 p.m. 10 p.m. 20.4 962.491— 19.97 - 2S.53 61.03 89.56 62.69 10 p.m. 1 a.m. 20.3 951.44 — 11.05 - 15.79 52.48 68.27 47.79 1 a.m. 4 a.m. 20.0 936.32'— 15.12 - 21.60 42.49 64.09 44.86 Dec. 20-21.. 4 a.m. 7 a.m. Total 19.3 897.22 — 39.10 — 55.86 18.52 74.38 52.07 113.04 — 161.49 467.75 J629.24 440.46 7 a.m. to 10 a.m. 19.4 899.01 + 1.79 + 2 . 56 100.58 98.02 68.61 10 a.m. 1 p.m. 19.0 874.42 — 24.59 — 35.13 47.60 82.73 57.91 1p.m. 4 p.m. 18.1 826.98 — 47.44 — 67.78 28.23 96.01 67.21 4 p.m. 7 p.m. 17.4 792.36 — 34.62 — 49.46 30.38 79.84 55.89 7p.m. 10p.m. 17.7 803.91 + 11 . 55 + 16.50 100.98 84.48 59.14 10 p.m. 1 a.m. 17.7 806.36 + 2.45 + 3 . 50 74.92 71.42 49.99 1 a.m. 4 a.m. 17.9 815.61 + 9.25 + 13.21 90.76 77.55 54.29 4 a.m. 7 a.m. Total 18.1 830.12 + 14. 5J + 20.73 - 95.87 76.78 56.05 39.24 — 67.10 550.23 646 . 10 452.28 Since weights rather than volumes are employed in many of the computa- tions, the equivalent in grams of the volume of oxygen lost or gained to the residual air is computed by dividing the volume by the factor 0.7, and recorded in column d. The actual weight of oxygen admitted to the chamber from the 3G Influence of Inanition on Metabolism. steel cylinder is obtained by noting its loss in weight, making due allowance for the nitrogen contained in the compressed oxygen. The weights of oxygen admitted are recorded in column e. Since these amounts do not represent the quantities actually consumed by the subject, they have been corrected for the changes in the residual amounts. The correct amounts consumed by the subject are recorded in grams in column f and in liters in column g, since the volume absorbed is of importance in subsequent computations. Table 8. — Elements katabolized in the body — Metabolism experiment No. 59. (a) Total weight. (b) Nitro- gen. (c) Carbon. W) (e) ^et0- Oron. Ash. First day, Dec. 18, 1903. Income: Oxygen from air Outgo: Grams. 629 . 42 Grams. Grams. Grams. Grama. 629.42 Grams. 1494.50 46.30 982.32 670.55 li!84 8!7i 182.86 167.23 2.60 109.92 1327.27 16.21 872 . 40 487.69 e'94 Solids in urine Carbon dioxide Total 3193.67 2564.25 11.84 11.84 191.57 191.57 279.75 12703.57 279.75 2074.15 6.94 6.94 Second day, Dec. 19, 1903. Income: Oxygen from air Outgo: Water in urine 629.24 .... .... .... 629.24 .... 927.00 54.80 951.95 658.94 14^11 io!46 179 .71 103.73 3.11 106.52 823.27 18.98 845.43 479.23 8^20 Solids in urine Water of respiration x Carbon dioxide Total 2592.69 1963.45 14.11 14.11 190.11 190.11 213.36 213.36 2166.91 1537.67 8.20 8.20 Loss Third day, Dec. 20, 1903. Income: Oxygen from air Outgo: Water in urine 646.10 .... .... .... 646.10 .... 871.20 57.70 943.15 650.24 14.82 l6!92 177\32 97.49 3.26 105.54 773.71 20.03 837.61 472.92 8!o7 Solids in urine Water of respiration l Carbon dioxide Total 2522.29 1876.19 14.82 14.82 188.24 188.24 206.29 206.29 2104.27 1458.17 8.67 8.67 Loss 1 Includes also water of perspiration. Material Katabolized in the Body. In fasting experiments, aside from the isolated instances of anabolic trans- formations, the transformations of matter are wholly katabolic, that is, the body is losing substance continually. In the fasting experiments here reported the income consisted solely of drinking-water and oxygen of the air. Of these two, oxygen alone is capable of entering into chemical combination. Using the chemical analyses of the urine and the measurements of carbon dioxide and Metabolism Experiment No. 59. 37 water output and oxygen intake, it is possible to strike a complete balance of intake and outgo. Such a balance may for convenience be tabulated in the form shown in table 8. The income, which on the first day amounted to 629.42 grams of oxygen, is small as compared with the total outgo, which amounted to 3193.67 grams of material. The outgo is subdivided into water of urine, solids of urine, water of respiration and perspiration, and carbon dioxide. In considering this table, it should further be stated that there were no feces passed during the time of this experiment, and moreover it was impossible to separate any feces which could with any certainty be designated " fasting feces." For this reason the question of the status of feces in connection with this table is eliminated. From the chemical composition of water and carbon dioxide and the chemical analysis of the solids in urine, it is possible to compute the amounts of the different elements involved in the katabolic transformations. Thus, the 1494.5 grams of water in the urine of the first day can be subdivided into 167.23 grams of hydrogen and 1327.27 grams of oxygen. In the case of solids in urine we have, from the chemical analyses, 11.84 grams of nitrogen, 8.71 grams of carbon, 2.6 grams of hydrogen of organic matter, 16.21 grams of oxygen, and 6.94 grams of ash, the ash here being treated as an element. The amounts of carbon and oxygen in carbon dioxide are also shown, together with the total outgo in terms of nitrogen, carbon, hydrogen, oxygen, and ash. By deducting the oxygen of the intake, a total loss to the body of 2564.25 grams of material is obtained, of which the larger part is obviously oxygen derived from the water of respiration and carbon dioxide and water in the urine. A similar computation for the second and third days shows losses to the body of 1963.45 grams and 1876.19 grams of material, respectively. From these losses of the chemical elements it is possible to compute the losses of compounds. The computations are based upon the chemical analyses of body protein, body fat, and glycogen. For body protein, the work of Koehler 4 has been taken. For body fat, the analyses of human fat in this laboratory are used,5 and the carbo- hydrates existing in the body are assumed to be chiefly glycogen. The following is a tabular statement of the composition assumed in these computations : Body material. N. C. H. O. Mineral matters (includ- ing S). Proteids Per cent. 16.67 Per cent. 52.80 76.10 44.40 Per cent. 7.00 11.80 6.20 11.19 Per cent. 22.00 12.10 49.40 88.81 Per cent. 1.53 Fat Carbohydrates Water , 4Zeit. f. physiol. Chemie (1901), 31, 498. 5 Benedict & Osterberg, Amer. Jour. Physiol. (1900), 4, p. 74. 38 Influence of Inanition on Metabolism. Disregarding the mineral matters, the following equations may be derived from these data, letting p = protein, t = fat, r = carbohydrates, and w = water : 0.16670 = N 0.4440r + 0.7610* + 0.5280» = C 0.1119m? + 0.0620r + 0.1180* + 0.0700p = H 0.8881W + 0.4940r + 0.1210* + 0.2200p = 0 Solving these equations in terms of 1ST, C, H, and O, the following formulae are obtained: Protein = 6.0 N. Fat = 0.005 C + 9.693 H — 1.221 O — 2.476 N. Carbohydrates = + 2.243 C — 16.613 H + 2.093 O — 2.892 N. Water = — 1.248 C + 7.920 H + 0.128 O + 0.460 N. The quantities of each element lost from the body, expressed in grams in table 8, may be substituted in these equations for the chemical elements N", H, O, and C. For purposes of illustration the computation of the losses of body compounds for the first day of experiment 59 are given in some detail in table 9. Table 9. — Illustrative table showing elements and material katabolized in body — Metabolism experiment No. 59. (a) Total weight. (b) Nitro- gen. (c) Carbon. (d) Hydro- gen. M Oxygen. (f) Ash. First day, Dec. 18, 1903 Outgo: Water in urine Grams. 629.42 Grams. Grams. Grams. Grams. 629.42 Grams. 1494.50 46.30 982.32 670.55 li'84 8!7i 182 ! 86 167.23 2.60 109.92 1327.27 16.21 872.40 487.69 e>'94 Solids in urine Water of respiration ' Carbon dioxide Total Loss 3193.67 2564.25 11.84 11.84 191.57 191.57 279.75 279.75 2703.57 2074.15 6.94 6.94 Katabolized body material: Protein 71.04 150.72 89.16 2247.48 5.85 11.84 37.51 114.70 39.59 4.97 17.78 5.53 251.49 15.63 18.24 44.04 1995.99 1.09 5^85 Fat Glycogen Water Ash Total 2564.25 11.84 191.80 279.77 2073.90 6.94 1 Includes also water of perspiration. As a mathematical verification of these computations the results are expressed in tabular form, and by use of the percentage composition shown above the amounts of nitrogen, carbon, hydrogen, and oxygen in the quantities of each compound lost may be recomputed. The sum of the different elements should Metabolism Experiment No. 59. 39 then equal the original quantities of elements lost. The mathematical verifi- cation of these computations is shown in table 9. Two features of this table demand special discussion. In the first place, while the actual amount of ash determined by chemical analyses is seen in table 4 to have been 6.94 grams/ here it is seen that only 5.85 grams of ash were katabolized. On the other hand, it must be borne in mind that in the 71.04 grams of protein katabolized there were 1.09 grams of ash, since protein con- tains 1.53 per cent of mineral matter. It is necessary, therefore, to deduct from the total amount of ash in the output, 6.94 grams 6 in this particular experiment, the total amount of ash in the protein, 1.09 grams, since the ash of the kata- bolized protein appears in the urine. Therefore the difference, 5.85 grams, is ash other than ash of protein. In the second place, while it has been stated above that the sum of the elements in these compounds, as computed from their percentage composition, should equal the quantities of elements lost, there are slight discrepancies, amounting to but 0.1 or 0.2 of a gram, which are due to the dropping of Table 10. — Elements and materials katabolized in the body — Metabolism experiment No. 59. Date. (a) Nitro- gen. (tt Carbon. (c) Hydro- gen. (d) Oxygen. (e) Water.1 (/) Protein. (a) Fat. (h) Carbo- hydrates (as glyco- gen). (i) Ash.i 1903. Dec. 18-19 Dec. 19-20 Dec. 20-21 Total, 3 days. Grams. 11.84 14.11 14 . 82 Grams. 191.57 190.11 188.24 Grams. 279.75 213.36 206.29 Grams. 2074.15 1537.67 1458.17 Grams. 2247. 4S 1655.87 1592.35 Grams. 71.04 84.66 88.92 Grams. 150.72 156.61 183.39 Grams. 89.16 59.41 4.22 Grams. 6.94 8.20 8.67 40.77 569.92 699.40 5069.99 5495 . 70 244 . 62 490.72 152.79 23.81 1 See pp. 28-29. supernumerary figures in the computations. The agreement is sufficiently close, however, to demonstrate the mathematical accuracy of the computation of the protein, fat, glycogen, water, and ash katabolized. Elements and materials katabolized in the tody. — As a result of the com- putations cited above, we have not only the amounts of the chemical elements lost from the body, but also the chemical compounds expressed in terms of water, protein, fat, carbohydrates, and ash. These are shown in table 10 for each day of the experiment. The figures for the first day are also shown in table 9. By inspection of the figures for the daily amounts it is seen that, save in the case of the carbohydrates, about the same quantities of materials were lost on 6 In this particular experiment the quantity of ash was calculated, not determined. See p. 28. 40 Influence of Inanition on Metabolism. each day, suggesting constancy of katabolism. In comparing the results of the three different days, there is an increasing loss of nitrogen, a nearly constant loss of carbon, a rapidly diminishing loss of hydrogen and oxygen, and consequently a corresponding diminution in the water lost. The striking difference, as has been pointed out above, is in the case of carbohydrates, in which the loss of glycogen becomes markedly less each day. But 4.22 grams of glycogen were oxidized in the body on the last day. Since experiment No. 59 is one of the earliest experiments in the series in which fasting metabolism was studied, it seems undesirable to discuss the peculiar characteristics of the fasting metabolism until opportunity is had to examine all of the available data; consequently, the data for the subsequent experiments will be presented in due course and the main discussion deferred until all the evidence has been considered. It does seem fitting, however, to explain somewhat more fully the use of terms employed in the tables above. In discussing material katabolized in the body, objection can be raised to the use of the expression " water katabolized in the body/' since in the compu- tations of the quantities of material katabolized water is involved only as pre- formed water. Aside from this preformed water, which leaves the body as such and is considered in the table as " water katabolized in the body," there is other water resulting from the oxidation of the protein, fat. and carbohydrates. The total water eliminated from the body, namely, the sum of the water of urine, respiration and perspiration, and feces, if any, includes not only the water katabolized in the body in the sense in which the expression is used above (i. e., preformed water), but also the water resulting from the katabolism of the protein, fats, and carbohydrates. Certain discrepancies appear when the attempt is made to compare the water katabolized in the body in table 9 with the total water output. There is an excess of water eliminated over the so-called " water katabolized in the body," but if the water of oxidation of the organic hydrogen of protein, fat, and carbohydrates katabolized is deducted from the total water elimination, the discrepancy disappears. Since a portion of the organic hydrogen of the protein katabolized, i. e., the organic hydrogen of the urine, is not oxidized, this amount must be deducted from the organic hydrogen of the protein. A mathematical verification of these points may be made in the following way : The total water eliminated equals the weight of the water in the urine, 1494.50 grams (see table 9) plus the water of respiration and perspiration, 982.32 grams, or a total of 2476.82 grams. The water katabolized from the body as preformed water is 2247.48 grams. Thus there was an excess of 229.34 grams excreted as the result of oxidized hydrogen of organic matter. During this day there were katabolized 71.04 grams of protein, 150.72 grams of fat, and 89.16 grams of glycogen. These materials contain 4.97, 17.78, and 5.53 grams of organic hydrogen, respectively — a total of 28.28 grams. Deduct- Metabolism Experiment No. 59. 41 ing the organic hydrogen of the urine, 2.60 grams, there remains 25.68 grams, the amount of organic hydrogen that was oxidized during the day. From the ratio of hydrogen to water it is computed that there are 229.6 grams of water resulting from this oxidation. Thus the discrepancy apparently existing be- tween the total elimination of water and the water katabolized from the body is explained by the water of oxidation of organic hydrogen. This computation assumes that all the organic hydrogen of the urine results from the decomposition products of protein. While the numerous tests for albumen and sugar, which were invariably negative, would preclude the possibility of organic hydrogen from these two sources, nevertheless the marked acidosis observed by Brugsch 7 would lead to the inference that organic matter other than that of the disintegration products of protein might well be present in the solids of urine in fasting experiments. With the data at hand, however, no better method of distinguishing between preformed water and water of oxidation of organic hydrogen has as yet been obtained. Table 11. — Distribution of intake and outgo of water — Metabolism experiment No. 59. Date. Outgo from the body. Balance of preformed water. to) Water of oxidation of organic hydrogen. (c-d) (a) Water of urine. (M Water of respira- tion and perspira- tion. (c) Total. (a + b) (d) Pre- formed (katabol- ized) water in outgo. (6) Intake and drink. (/) Loss of pre- formed water. (d-e) 1903. Dec. 18-19 Dec. 19-20 Dec. 20-21 Total, 3 days. . Av.perday. . . . Grams. 1494.50 927.00 871.20 Grams. 982.32 951.95 943.15 Grams. 2476.82 1878.95 1814.35 Grams. 2247.48 1655.87 1592.35 Grams. 1342.50 1360.20 1188.00 Grams. 904.98 295.67 404.35 Grams. 229.34 223.08 222.00 3292.70 1097.57 2877.42 959.14 6170.12 2056.71 5495.70 1S31.90 3890.70 1296.90 1605.00 535.00 674.42 224.81 Balance of water. — While the figures in table 10 indicate the loss of elements and materials from the body, the income of water has not been taken into consideration in any of the computations thus far, while the oxygen of the income has been duly considered. Hence it is clear that the loss of water from the body as given in column e of table 10 is not the net amount lost, since the body received certain amounts of water daily. Thus, on the first day, there were consumed 1342.5 grams of water; on the second day, 1360.2 grams; on the third day, 1188.0 grams. If the amount of drinking-water is taken into consideration, the actual loss of water from the body is much less than appears in table 10. The true loss of preformed water is therefore the 7Zeit. f. exper. Path. u. Ther. (1905), 1, p. 419. Zeit. f. exper. Path. u. Ther. (1906), 3, p. 675. Cf. also Bonniger and Mohr, 42 Influence of Inanition on Metabolism. difference between the loss represented in table 10 and the intake of water as given in table 2; consequently, on the first day, the loss of water from body tissues would be 2247.48 — 1342.50, or 904.98 grams. A similar computation shows the losses of the second and third days to be 295.67 and 404.35 grams, respectively. The data for obtaining these losses are tabulated in table 11. Aside from the preformed water in' the muscles and tissues of the body which was lost during fasting, there remains to be considered the water resulting from the oxidation of the protein, fats, and carbohydrates katabolized, or the water of oxidation of the organic hydrogen contained in these compounds. Since this water was measured together with the rest of the output, it is included in the total amount of water eliminated from the body, namely, column c. Deducting the preformed water lost (column d), the values for the water of oxidation of the organic hydrogen are obtained. These are recorded in column g. Output of Heat. Measurements with the respiration calorimeter include the determination of the heat output of the body. As these measurements have a very direct and important bearing upon the measurements of body material katabolized, a careful inspection of the heat data is of interest. The larger portion of the heat given off from the body is measured by the current of cold water which passes through the heat absorbers inside the respiration chamber. From the differences in temperature of the water as it enters and leaves the chamber, and the mass of water, the heat absorbed can be computed. Correction for specific heat of water. — The measurement of heat by this method involves the use of a varying standard, i. e., the calorie at the different ranges of temperature through which the water is warmed, and it is accordingly necessary to reduce these observed heat values to the standard calorie, which in all of the work with the respiration calorimeter thus far has been the calorie at 20° C.8 There are not immaterial differences in the specific heat of water at different temperatures. These specific heats have been computed and placed in tabular form elsewhere.9 The method of reducing this heat to terms of C20 is shown in table 12. Thus, in column a is recorded the heat measured as the product of the mass of water and the differences in temperature. In column o, the range of temperature as expressed in degrees centigrade, and in column c the specific heat for these ranges of temperature are shown. By multiplying the heat measured by the specific heat of water at the temperatures used, the heat in terms of C20 recorded in column d can be obtained. 6 The calorie used in all measurements discussed in this report is the quantity of heat required to raise 1 kilo of water from 19.5° to 20.5° C. 0 U. S. Dept. of Agr., Office of Exp. Sta. Bui. 63, p. 56. Metabolism Experiment No. 59. 43 While these changes are small, amounting to less than 4 calories for the whole day, and while it may appear that other discrepancies much greater are neglected, nevertheless it is our custom to include this change in an endeavor to secure the greatest possible accuracy with every factor determined by means of this apparatus. An extended discussion of the errors involved in the different determinations is given elsewhere.10 Correction for temperature of water and dishes. — In addition to the heat brought away by the water-current, there are minor changes in the amount of heat in the chamber resulting from the introduction of material at a tem- perature below or above that of the chamber. In fasting experiments these changes are at a minimum and consist chiefly of the introduction of cold Table 12. — Heat reduced to terms of C-0 — Metabolism experiment No. 59 (December 18-19, 1903). Date and period. (a) Heat measured. (6) Range of temper- ature. (O Specific heat. (d) Heat in terms Of C20. (axe) 7 a.m. to 10 a.m. . . Calories. 257.3 9-14 1.0022 Calories. 257.9 10 a.m. 1 p.m. . . 204.0 9-14 1.0022 204.4 1 p.m. 4 p.m. . . 199.9 8-14 1.0025 200.4 4 p.m. 7 p.m. . . 222.6 9-14 1.0022 223.1 7 p.m. 10 p.m. . . 205 . 7 9-14 1.0022 206.2 10 p.m. 1 a.m. . . 131.2 9-15 1.0021 131.5 1 a.m. 4 a.m 117.1 9-17 1.0018 117.3 4 a.m. 7 a.m. . . Total for day. . . 157.9 8-17 1.0020 158.2 1495.7 1499.0 drinking-water contained in glass bottles. Occasionally the urine or feces may be withdrawn before they reach the temperature of the chamber. In all cases the time at which the material is put in or taken from the chamber, the nature of the material, and the weight and temperature are recorded. The amounts of heat lost or gained by the water-current due to the differences between the temperature of the calorimeter and the water and dishes used on the first day of experiment No. 59 are given in table 13. The specific heat of glass is taken as 0.2. From the specific heat of the material, the weight in grams, and the differences between the temperature of the material and the calorimeter chamber, the quantities of heat absorbed or given off are computed. In only one instance during this experiment was any material introduced into the respiration calorimeter above the temperature of the chamber. At 4.30 p. m., December 18, four glass urine bottles, weighing 1743 grams, were introduced at a temperature 1° above that of the chamber. All other cor- 10 Carnegie Institution of Washington Publication No. 42; U. S. Dept. of Agr., Office of Exp. Sta. Bui. 175. 44 Influence of Inanition on Metabolism. rections were clue to cold drinking-water. The sum of the corrections for this day is 22.7 calories, or, in other words, 22.7 calories of heat were required to warm the objects to the temperature of the calorimeter chamber, and hence were not measured in the water-current. Capacity correction of the calorimeter. — Even with the most skillful physical assistants slight fluctuations in the temperature of the calorimeter are unavoid- able, and since the large mass of metal is capable of storing and yielding considerable quantities of heat, the result of these fluctuations must be taken into consideration to determine correctly the total heat elimination for any given period. From a number of tests, which, however, admittedly partake more Table 13. — Correction applied to the heat measured for temperature of water and dishes — Metabolism experiment No. 59 (December 18-19, 1903). Time. Article. (a) Specific heat. CM Weight. W> Temper- ature. (d) Temper- ature of calo- rimeter. (6) Differ- ence. if) Heat (axbXe). 8h00m a.m. . Water.. Glass . . 1.0 .2 12h 10™ p.m. . Water.. Glass . . 1.0 .2 4h 30m p.m. . Water.. Glass . . Do .. 1.0 .2 .2 10ii5m p. m. . Water.. Glass . . 1.0 . 2 Total.. Grams. 400 297 400 640 400 297 1743 400 297 °c 7.6 7.6 °C 19.9 °C 12.3 8.5 8.5 20.0 11.5 7.8 7.8 21.0 20.0 20.0 12.2 i.o 7.8 7.8 20.0 12.2 Calories. + 4.920 + .731 + 4.600 + 1.472 + 4.880 + .725 - .349 + 4.880 + .725 (0) Correc- tion. Calories. + 5.7 + 6.1 + 5.3 + 5.6 22.7 of the nature of close estimates than of scientific experiments, the hydrothermal equivalent of the calorimeter has been found to be 60 kilos. Consequently, to obtain the capacity correction of the calorimeter, the temperature fluctuations in degrees centigrade are multiplied by 60. Correction for heat in bedding. — The necessity of correcting the heat measured for the heat in the bedding is based upon the fact that during the period from 10 p. m. to 1 a. m., when the subject retires at night, a considerable quantity of heat is produced which is not liberated into the air of the chamber, but is retained by the bed and bedding until some time after the subject has risen in the morning. From the heat eliminated and other factors it has been computed, though by crude methods, that 30 calories of heat are thus absorbed and liberated by the bedding. Consequently, during the first period in the morning, there is a certain increment of heat measured by the water- Metabolism Experiment No. 59. 45 current which was stored in the bedding during the previous night, and hence must be deducted. Table 14. — Summary of calorimetric measurements and total heat eliminated — Metabolism experiment No. 59. Date and period. 1903. Dec. 18: Preliminary: 1 a.m. to 4 a.m. 4 a.m. 7 a.m. Total. Dec. 18-19: 7 a.m. to 10 a.m. 10 a.m. lp.m. 4 p.m. 7 p.m. 10 p.m. 1 a.m. 4 a.m. Total. 1 p.m. 4 p.m. 7 p.m. 10 p.m. 1 a.m. 4 a.m. 7 a.m. Dec. 19-20: 7 a.m. to 10 a.m. 1 p.m. 10 a.m. 1 p.m. 4 p.m. 7 p.m. 10 p.m. 1 a.m. 4 a.m. 4 p.m. 7 p.m. 10 p.m. 1 a.m. 4 a.m. 7 a.m. Total , Dec. 20-21 : 7 a.m. to 10 a.m. 10 a.m. 1 p.m. 4 p.m. 7p.m. 10 p.m. 1 a.m. 4 a.m. Total. 1 p.m. 4 p.m. 7 p.m. 10 p.m. 1 a.m. 4 a.m. 7 a.m. (a) Heat meas- ured in terms of C20. Cals. 146.07 140.57 286.64 (W Change in temper- ature of calo- rimeter. °C. + 0.10 - .10 (O Capaci- ty cor- rection of calo- rimeter. (b X 60) Cals. + 6.00 -6.00 (d) Correc- tion due to temper- ature of water and dishes. Cals. (e) Correc- tion for heat in bedding1 retained (+). liberated (-)• Cals. (/) Water vapor- ized equals total amount in out- going air plus gain or loss from preced- ing period. Grams. 125.61 133.20 258.81 (fif) Heat used in vapori- zation of water. (/X0.593) Cals. 74.36 78.85 153.21 (h) Total heat elimi- nated. (a+c+ d+e+g). Cals. 226.43 213.42 439.85 257.90 + .10 + 6.00 + 5.70 -30.00 136.41 80.75 204.40 + .02 + 1.20 + 6.10 • • • ■ 130.01 76.97 200.40 + .01 + .60 .... .... 119.64 70.83 223 . 10 — .14 -8.40 + 5.30 • • • • 133.74 79.17 206.20 + .07 + 4.20 + 5.60 • • • • 114.34 67.69 131.50 — .04 -2.40 • • • . + 30.00 116.87 69.19 117.30 + .03 + 1.80 • • • • .... 115.41 68.32 158.20 + .05 +3.00 .... .... 115.90 68.61 1499.00 + .10 | + 6.00 + 22.70 .... 982.32 581.53 j 320.35 288.67 271.83 299.17 283.69 228.29 187.42 229.81 !109.23 246.60 203.50 192.80 212.50 222.10 136.20 134.40 172 . 50 1520.60 .09 .01 .11 .08 .07 .03 .13 .06 — .04 — 5.40 — .60 + 6.60 — 4.80 + 4.20 + 1. -7 + 3 80 80 60 -2.40 + 5.60 + 5.90 + 4.80 + 5.40 + 21.70 -30.00 + 30.00 124.47 121.97 114.98 120.91 121.68 115.15 111.53 121.26 73.69 72.21 68.06 71.58 72.03 68.17 66.03 71.79 951.95 563.56 290.49 281.01 267.46 284.08 303.73 236.17 192.63 247.89 2103.46 248.60 _ .04 -2.40 194.00 + .10 + 6.00 199.80 — .07 — 4.20 218.10 + .03 + 1.80 221.70 — .05 -3.00 140.80 — .04 -2.40 161.30 + .09 + 5.40 153.20 — .04 -2.40 1537.50 — .02 -1.20 + 5.60 + 5.30 - 6!50 + 4.50 + 14.90 -30.00 119.60 70.80 ... 120.62 71.41 .... 119.34 70.65 • > * ■ 114.11 67.55 .... 119.43 70.70 + 30.00 111.25 65.86 • • ■ • 121.98 72.21 116.82 69.16 943 . 15 558.34 292.60 276.71 266.25 286.95 293 . 90 234.26 238.91 219 96 2109.54 Consequently, during the first period in the morning, the correction is negative, since the heat is in excess of that normally produced, while during 46 Influence of Inanition on Metabolism. the period from 10 p. m. to 1 a. m. the correction is positive. It is important to note that no matter what error affects this correction, it is a compensating one and hence it does not affect the measurement of the heat elimination for the 2-1 hours. It affects only the two periods to which the corrections are applied. Correction for the heat of vaporization of water. — A very considerable pro- portion of the total heat given off from the body, especially during rest, is used for the vaporization of water from the lungs and skin. The amount of water thus vaporized equals the total amount in the outgoing air plus the gain or loss of water vapor in the residual air from the preceding period. For the want of a more accurate factor for the heat of vaporization of 1 gram of water at the temperature of the calorimeter we have used the factor 0.592, based on Begnault's work. Summary of calorimetric measurements. — The various corrections mentioned above to be applied to the heat measured by the calorimeter are given in table 14, the total corrected heat eliminated being recorded in column h. Total Heat Production. The simple measurement of heat lost by the body is not sufficient for the proper interpretation of the processes of metabolism. Since metabolism and heat production rather than heat elimination go hand in hand, it is of vital importance to distinguish between heat production and heat elimination; and for accurate measurements of heat production, knowledge of the fluctuations in the store of heat in the body is absolutely essential. Changes in the residual heat in the body due to variation in body weight and temperature. — The body of the subject may be said to be a large reservoir for heat and the quantity of heat thus stored in the body varies with every period of the day. Even with constant body temperature there are noticeable fluctuations in body-weight which involve considerable changes in the amount of heat actually existing in the body. Similarly, differences in body temper- ature cause fluctuations in the storage of heat. For example, if a subject weighs 60 kilos and there is a rise of temperature of 1° C, this is practically equivalent to the storage of 50 calories, since the specific heat of the body is not far from 0.83. A corresponding fall of temperature is followed by a corresponding loss from the store of heat. But even though the temperature be constant, if the body loses 500 grams in weight, obviously 500 grams of material have been cooled from the temperature of the body, 37° C, to the temperature of the chamber, 20° C, and thus 7.1 calories of heat have been given up which were not actually produced during the period. If the calorimeter chamber were provided with a weighing arrangement whereby body-weights could be taken at the end of each experimental period, part of the data for these corrections could be readily secured. Unfortunately, Metabolism Experiment No. 59. 47 as yet no simple arrangement seems to be possible for securing the body- weight of the subject at the end of each period without the extraneous muscular effort of adjusting the furniture in the chamber and suspending the weighing chair on the special scale provided for the purpose. Therefore the weighings are usually made but once each day. From the weights so found, the changes in body-weight from period to period were computed. The details of such computations are shown for the first day of this experiment in table 15. The income consists of oxygen and water consumed, and the outgo, of water of respiration and perspiration, carbon dioxide, and urine. During the period Table 15. — Change of body-weight during different periods of the day — Metabolism experiment No. 59 (December 18-19, 1903). Time. Income. (a) Oxygen con- sumed. (b) Water con- sumed. Outgo. (O Water of respira- tion.1 (d) Carbon dioxide elimina- ted. (e) Urine. (/) Balance (o+6) -(c+d+e). (ff) Body- weight. 7 a.m 7 to 10 a.m 10 a.m 10 a.m. to 1 p.m 1 p.m 1 to 4 p.m 4 p.m 4 to 7 p.m 7 p.m 7 to 10 p.m 10 p.m 10 p.m. to 1 a.m 1 a.m 1 to 4 a.m 4 a.m 4 to 7 a.m 7 a.m Ora ms. 95 33 79 66 74 64 96 60 85 64 69 .22 63 .35 64 .98 • Grams. 400.00 400.00 400.00 142.50 Orams. 136 41 130 01 119 64 133 74 114 34 116 87 115 .41 115 .90 • • • • Orams. 10l".76 88 '.82 88/29 98^58 81.70 74.12 65.29 71.99 Orams. Orams. 455 80 -198.64 190 20 + 70.63 252 20 — 385.49 528 70 — 264.42 129 30 -239.70 203 70 -182.97 • -117.35 . . . -122.91 • .... Kilos. 67.796 67.597 67.668 67.282 67.018 66.778 66.595 66.478 66.355 1 Includes also water of perspiration. from 7 to 10 a. m. there was a total loss from the body of 198.64 grams over and above the income of oxygen and water. Thus the body-weight at 10 o'clock was less by 199 grams than it was at 7 a. m. During the next period the income was slightly larger than the output and therefore the body-weight was slightly increased. The computations for the remaining periods of the day, with the final corrected body-weight, appear in the table. Unfortunately, in this particular experiment the special scale for weighing the man had not been installed and the body-weights were very defective, but from later observations on this same subject it is assumed that when naked he weighed 67.796 kilos. That this figure is very nearly correct seems fairly 48 Influence of Inanition on Metabolism. certain from a comparison of these data with weights taken by the subject, the weight of clothes being taken into consideration. The figure above was therefore taken as a basis. Table 16. — Determinations of body temperature and body-weight, with correspond- ing heat corrections — Metabolism experiment No. 59. Date and time. Body temperature. Body-weight. (e) Correction for change of body temperature and weight. (a) By rectal thermome- ter. (ft) Rise (+)or fall (-) from preceding period. (O Calculated. (d) Gain (+) or loss (-) from preceding period. 1903. °C. 36.72 °C. Kilos. 167.796 Grams. Calories. 37.00 + 0.28 67.597 — 199 + 12.92 36.75 - .25 67.668 + 71 + 13.04 36.84 + .09 67.282 - 386 — .34 7 p.m 36.94 + .10 67.018 — 264 + 1.86 lOp.m 36.55 - .39 66.778 — 240 — 24.96 36.35 - .20 66.595 — 183 — 13.58 36.45 + .10 66.478 - 117 + 3.94 Total for day.. . Dec. 19, 7 a.m 36.56 + .11 66.355 - 123 + 4.38 - .16 — 1441 — 28.82 36.56 • * • • 66.355 36.96 + .40 66.401 + 46 + 22.68 36.83 — .13 66.579 + 178 — 4.72 4 p.m 36.85 + .02 66.352 — 227 — 2.06 7 p.m 36.73 - .12 66.499 + 147 — 4.56 lOp.m 36.61 — .12 66.370 - 129 — 8.40 Dec. 20, 1 a.m 36.45 - .16 66.155 — 215 - 11.72 36.61 + .16 66.041 — 114 + 7.20 Total for day... Dec. 20, 7 a.m 36.73 + .12 65.917 — 124 + 4.86 + .17 — 438 + 3.28 36.73 65.917 36.89 + .16 65.798 - 119 + 7.08 36.99 + .10 65.953 + 155 + 7.62 4 p.m 37.00 + .01 65.697 - 256 — 3.04 37.01 + .01 65.388 - 309 - 3.84 lOp.m 36.58 _ .43 65.553 + 165 - 21.06 36.41 - .17 65.339 — 214 — 12.16 36.45 + .04 65.221 — 118 + .58 7 a.m Total for day. . . 36.80 + .35 65.093 — 128 + 17.14 + .07 — 824 — 7.68 1 The weight given for " a. m. arbitrarily assumed. In addition to the fluctuations in body-weight, it is necessary in order to get the true heat produced, to know also the changes in body temperature. These latter were taken with the electrical rectal thermometer, which has been described in detail.11 While reserving the list of thermometric observations for "Benedict & Snell, Archiv f. d. ges. Physiol. (1901), 88, pp. 492-500; Carnegie Institution of Washington Publication No. 42, p. 156. Metabolism Experiment No. 59. 49 discussion elsewhere, it will suffice for the particular purpose in view to give the body temperature determinations for the end of each period. These are recorded in column a of table 16. The differences from period to period are recorded in the second column. The data recording the body-weight are given in columns c and d, while the final column gives the differences, due to changes in body-weight and temperature, in the amount of heat stored in the body. Table 17. — Total heat production — Metabolism experiment No. 59. Date and period. (a) Total heat eliminated. CM Correction due to change of body tem- perature and weight. (c) Total heat production. (a + b) 1903. Dec. 18-19, 7 a.m. to 10 a.m Calories. 320.35 288.67 271.83 299.17 283.69 228.29 187.42 229.81 Calorics. + 12.92 -13.04 — .34 + 1.86 -24.96 — 13.58 + 3.94 + 4.38 Calories. 333.27 275.63 271.49 301.03 258.73 214.71 191.36 234 . 19 Total 2109.23 — 28.82 2080.41 Dec. 19-20, 7 a.m. to 10 a.m 290.49 281.01 267.46 284.08 303.73 236.17 192 . 63 247.89 + 22.68 — 4.72 - 2.06 - 4.56 — 8.40 -11.72 + 7.20 + 4.86 313.17 276.29 265.40 279.52 295.33 224.45 199.83 252.75 10 a.m. 1 p.m 4 p.m. 7 p.m 1 a.m. 4am 4 a.m. 7 a.m Total Dec. 20-21, 7 a.m. to 10 a.m 2103.46 + 3.28 2106.74 292.60 276.71 266.25 286.95 293.90 234.26 238.91 219.96 + 7.08 + 7.62 — 3.04 — 3.84 -21.06 — 12.16 + .58 + 17.14 299.68 284.33 263.21 283.11 272.84 222.10 239.49 237.10 10 a.m. 1 p.m 7 p.m. 10 p.m Total 2109.54 - 7.68 2101.86 In order to show the method by which the chauge in the amount of heat stored in the body of the subject (column e) is found, a concrete example is here taken. At 7 a. m., December 18, the subject weighed 67.796 kilos, the body temperature was 36.72°, and the calorimeter temperature was 20°. If his body had cooled down to the calorimeter temperature immediately, the quantity of heat which would have been liberated would be 67.796 X 0.83 (the specific heat of body material) X 16.72 (difference between the temperature of the 50 Influence of Inanition on Metabolism. body and the calorimeter) = 9-10.88 calories. Similarly, at 10 a. m. the body weight was 67.597 kilos, and the difference between body temperature and calorimeter temperature was 17°. The formula would then be 67.597 X 0.83 X 17.0 = 953.80. The difference between the amounts of heat in the body reduced to 20° at 7 a. m. and 10 a. m. is 953.80 — 940.88 = 12.92 calories. There were, therefore, 12.92 calories of heat stored in the body more than were present at 7 a. m., and consequent^, since this quantity of stored heat was derived from the heat produced during this period, it should be added to the heat eliminated to give the heat production. The results for all the different periods are recorded in column e. Relation of heat production to lieat elimination. — The total heat production is the heat eliminated corrected for changes in the amount of heat residual in the body. The corrections for the heat elimination have previously been Table 18. — Amounts of body protein, fat, and glycogen katabolized and energy of each1 — Metabolism experiment No. 59. Date. Protein. Fat. Glycogen. (a) Total energy. (a) (6) (c) (d) (e) (/) Amount. Energy. Amount. Energy. Amount. Energy. (b+d+f) 1903. Dec. 18-19 Grams. 71.04 Calories. 401 Grams. 150.72 Calories. 1438 Grams. 89.16 Calories. 374 Calories. 2213 Dec. 19-20 84.66 478 156.61 1494 59.41 249 2221 Dec. 20-21 88.92 502 183.39 1750 4.22 18 2270 1 Factors for heat of combustion per gram of protein, 6.65 calories; fat, 9.54 calories; glycogen, 4.19 calories. considered and are tabulated in columns c to g of table 14. The corrections for the changes in the amounts of heat residual in the body as found in table 16 are applied to the corrected heat elimination and thus the total heat production is obtained. These results are shown in table 17. Energy of body material lost. — The quantities of protein, fat, and carbo- hydrates katabolized in the body have been computed from the chemical analyses and the formulae given on page 38. The energy of these compounds may be computed from the weights katabolized and the heats of combustion of body protein, fat, and carbohydrates. The heat of combustion of fat-free muscular tissue from which the nitrogenous extractives have not been removed is not far from 5.65 calories per gram. A large number of determinations of the heat of combustion of human fat made in this laboratory " averaged 9.54 calories per gram. The heat of combustion of glycogen has frequently been determined as 4.19 calories per gram. The computation, therefore, of the energy resulting from the katabolism of any one of these three different compounds in the "Benedict & Osterberg, Amer. Journ. Physiol. (1900), 4, pp. 69-76. Metabolism Experiment No. 59. 51 body may be made by multiplying its amount in grams by its heat of com- bustion. The results are stated in table 18. The total energy which would result from the katabolism of the varying amounts of the protein, fat, and glycogen is given in the last column. It should be here noted that these figures represent the total potential energy of the different materials and not the actual energy liberated in the body, for while in the katabolism of fat and carbohydrates the total heat of oxidation is liberated, in the katabolism of protein a certain moiety is eliminated as the potential energy of urine. Balance of Energy. As the total heat produced is derived from the katabolism of body protein, fat, and carbohydrates, there should be an exact balance between the computed Table 19. — Comparison of energy derived from katabolized body material loith total heat production — Metabolism experiment No. 59. Date. Energy derived from different sources. (ff) Total heat produc- tion. Energy from body material From body protein. id) From body fat. (e) From body glyco- gen. if) Total. (c+cl+e) greater than output. (a) Energ-y of pro- tein katabo- lized. (b) Poten- tial energy of urine. (c) Net energy (a-b). CO Amt. (f-g) (i) Pro- portion. (K+g) 1903. Dec. 18-19 Dec. 19-20 Dec. 20-21 Total, 3 days. Av. per day. . Cals. 401 478 502 Cals. 91 106 111 Cals. 310 372 391 Cals. 1438 1494 1750 4682 1561 Cals. 374 249 18 Cals. 2122 2115 2159 Cals. 2080 2107 2102 Cals. 42 8 57 Per ct. 2.0 0.4 2.7 1381 460 308 103 1073 357 641 214 6396 2132 6289 2096 107 36 1.7 total energy of the materials oxidized in the body and the total heat produced. That such a balance exists, though it is not a perfect one, appears from a consideration of table 19. Since, however, not all the potential energy of the protein is actually liberated in its katabolism, the net energy from the body protein is derived by deducting the potential energy of the urine from the energy of the protein katabolized. This computation is made in the first three columns of the table. The energy of body fat and glycogen katabolized are iden- tical with the amounts shown in table 18. The total energy derived from the different sources, therefore, is the sum of the net energy of the protein, the heat of combustion of the body fat, and the heat of combustion of the body glycogen. This total is recorded in column /. The total heat production shown in table 17 is recorded in column g above. Theoretically at least, we should expect that the total heat production would correspond to the total energy derived from the Influence of Ixaxitiox ox Metabolism. different sources. The difference between these two quantities is shown in column h and the percentage discrepancy is given in the last column. The difficulties of striking an accurate balance of this kind for an experiment of 24 hours' duration are obvious, since there are a number of physiological errors involved which preclude extremely accurate work. On the other hand, if the experimental period be made to cover several days, the physiological errors may and doubtless do in a large measure compensate. From the average of the three days of this experiment, it appears that the average heat production as measured by the calorimeter with corrections is 36 calories less than the estimated energy derived from the different sources or an error of 4- 1.7 per cent. Further discussion of the magnitude of this error and its relation to other experiments is deferred. In computing the percentage discrepancy between the total heat production and the estimated energy of material oxidized in the body the former is used as the standard of measurement and the error assumed to be due to the estimates of the energy of material oxidized in the body. Although the direct measurement of oxygen furnishes data for a more exact estimation of the products of katabolism than has heretofore been obtained, it is believed that at present the errors in the assumption of the energy of material katabolized in the body are greater than the errors in the determination of the heat pro- duction and hence the latter factor is used as the standard for comparison. It is to be noted, however, that not only in this, but in the whole series of experi- ments here reported, the agreement between the total heat production and the estimate of the energy of body material actually oxidized is, on the whole, very satisfactory. Relations between Oxygen Consumption, Carbon Dioxide Elimination, and Heat Production. Since heat is the result of the oxidative processes in the body it is natural to suppose that there would be some simple relations existing between the quantities of oxygen absorbed, the quantities of carbon dioxide eliminated, and the heat produced. The data on this point are given in table 20. In earlier experiments, where the amounts of oxygen and carbon dioxide were determined, the heat was not directly determined, but the ratio of the respiratory gases has been of great value in estimating the heat production. Oxygen thermal quotient. — The ratio between the amount of oxygen con- sumed and the amount of heat produced may be termed the oxygen thermal quotient, and this ratio can be expressed as the number of grams of oxygen that is absorbed to produce 100 calories of heat. The ratios between the oxygen, carbon dioxide, and heat are of interest not only for the whole day, but also for the shorter periods, and hence in column c the oxygen thermal quotient for each 3-hour period has been recorded. Metabolism Experiment No. 59. 53 Carbon dioxide thermal quotient. — The ratio between the carbon dioxide eliminated and the heat produced is also of value in interpreting the nature of the metabolism. This ratio, which is expressed in terms of the number of Table 20. — Oxygen and carbon dioxide thermal quotients, and respiratory quotients — Metabolism experiment No. 59. Date and period. 1903. Dec. 18: Preliminary: 1 a.m. to 4 a.m. 4 a.m. 7 a.m. Dec. 1S-19: 7 a.m. to 10 a.m. 1 p.m. 10 a.m. 1 p.m. 4 p.m. 7 p.m. 10p.m. 1 a.m. 4 a.m. Total. 4 p.m. 7 p.m. 10 p.m. 1 a.m. 4 a.m. 7 a.m. Dec. 19-20: 7 a.m. to 10 a.m. 10 a.m. 1 p.m. 4 p.m. 7 p.m. 10 p.m. 1 a.m. 4 a.m. Total Dec. 20-21: 7 a.m. to 10 a.m. 1 p.m. 4 p.m. 7 p.m. 10 p.m. 1 a.m. 4 a.m. 7 a.m. 10 a.m. 1 p.m. 4 p.m. 7 p.m. 10 p.m. 1 a.m. 4 a.m. Total. 1 p.m. 4 p.m. 7 p.m. 10 p.m. 1 a.m. 4 a.m. 7 a.m. (a) Total heat produc- tion. (6) Oxygen con- sumed. Cals. 333.3 275.6 271.5 301.0 258.7 214.7 191.4 234.2 2080.4 299.7 284.3 263.2 283.1 ! 272.9 222.1 239.5 237.1 2101.9 Grams. 58.9 70.4 (c) (d) Oxygen Carbon thermal dioxide quotient elimina- (100 b+a). ted. Grams. .... 69.7 .... 79.0 (e) Carbon dioxide thermal quotient (100 d+a). 95 79 74.6 96.6 85.6 69.2 63.4 65.0 629.4 (/) (g) Volume of Volume carbon of dioxide oxygen elimi- con- nated sumed (dX 0.5091). (b X 0.7). Liters. Liters. 35.5 41.2 40.2 49.3 (70 Respi- ratory quo- tient 28.6 28.9 27.5 32.1 33.1 32.2 33.1 27.8 30.3 101.8 88.8 88.3 98.6 81.7 74.1 65.3 72.0 670.6 30.5 32.2 32.5 32.8 31.6 34.5 34.1 30.7 51.8 45.2 45.0 50.2 41.6 37.7 33.2 36.7 32.2 341.4 66.7 55.8 52.3 67.6 59.9 48.5 44.3 45.5 440.6 313.2 102.5 32.7 95.0 276.3 72.5 26.3 S5.9 265.4 77.7 29.3 83.4 279.5 80.2 28.7 88.5 295.3 89.6 30.3 90.7 224.5 68.3 30.4 72.1 199.8 64.1 32.1 65.9 252.7 74.4 29.4 77.5 2106.7 629.3 29.9 659.0 30.3 31.1 31.4 31.6 30.7 32.1 33.0 30.7 31.3 48.4 43.8 42.4 45.0 46.2 36.7 33.5 39.5 335.5 71.8 50.8 54.4 56.1 62.7 47.8 44.8 52.1 440.5 98.0 32.7 95.9 82.7 29.1 83.6 96.0 36.5 85.7 79.8 28.2 84.2 84.5 31.0 86.4 71.4 32.2 74.1 77.6 32.4 73.3 56.1 23.6 67.1 646.1 30.7 650.3 32.0 29.4 32.6 29.7 31.7 33.4 30.6 28.3 48.8 42.5 43.7 42.9 44.0 37.7 37.3 34.2 68.6 57.9 67.2 55.9 59.2 50.0 54.3 39.2 30.9 331.1 , 452.3 0.86 .82 .78 .81 .86 .74 .69 .78 .75 .81 .78 .67 .86 .78 .80 .74 .77 .75 .76 .76 .71 .73 .65 .77 .74 .76 .69 .S7 73 grams of carbon dioxide accompanying the production of 100 calories of heat, has been given in column e. A particular discussion of this, as well as other quotients, will be found beyond. Respiratory quotient. — The relationship first recognized and studied was that existing between the carbon dioxide eliminated and the oxygen consumed — 54 Influence of Inanition on Metabolism. the so-called respiratory quotient. This respiratory quotient is based upon relations of the volume of these two gases, and therefore the volumes of carbon dioxide eliminated and oxygen consumed have been recorded in columns f and g. These are readily obtained from the weights of carbon dioxide and oxygen by means of the factors given in the column headings. The respiratory quotient is the ratio between them, as expressed in the last column. Numerous incidental errors affect at times very noticeably the value of the respiratory quotient for the different periods. These errors have been discussed in detail elsewhere." For the 2-i hours, however, the respiratory quotients are probably not far from correct. These values range from 0.78 on the first day of the experiment to 0.73 on the last day. "Carnegie Institution of Washington Publication No. 42; U. S. Dept. Agric, Office of Exp. Sta. Bui. 175. Metabolism Experiment No. 68. 55 Metabolism Experiment No. 68. This experiment, which lasted two days, followed immediately a series of experiments covering 11 consecutive days, made with the same subject inside the respiration calorimeter. The previous series has been reported elsewhere." While in the majority of the fasting experiments here reported, accurate knowledge of the metabolic activity before the fast was not available, in this instance the metabolism for several days before the fasting period was accurately measured. Three series of experiments preceded the fasting period. The first was a 3-day experiment, during which the subject was engaged in hard muscular labor riding a bicycle ergometer15 for 6 hours each day and received a diet furnishing about 100 grams of protein and 4630 calories of energy per day. The diet during this period consisted in large part of carbohydrates. The second was also a work experiment of 3 days' duration, during which a diet containing approximately the same amount of nitrogen as the preceding, but with slightly increased energy, averaging not far from 4750 calories per day, was given. During this experiment the diet consisted in large part of fat, chiefly in the form of cream. The third series consisted of 4 experiments, beginning with a 1-day experiment, during which a diet containing 19.11 grams of nitrogen and 5393 calories of energy, chiefly in the form of cream, was provided. On this day the subject did an unusual amount of muscular work on the bicycle ergometer, riding until far into the night. The severe work day was followed by a rest experiment of 1 day, during which the subject spent the most of his time in bed, recuperating from the excessive work of the preceding day. The diet contained 15.4 grams of nitrogen and 2369 calories of energy. During the next experiment, which was also of 1 day's duration, the subject prepared the ergometer and removed clothing for riding, but immediately dressed himself and did not ride, the purpose being to determine the energy required for the extraneous muscular work other than that involved in riding the ergometer. The diet for this day furnished 14.11 grams of nitrogen and 2062 calories of energy. On each of the two following days the subject rode the bicycle ergometer for 6 hours. The riding corresponded to the free leg- motion of coasting, since no resistance was applied to cause the subject to do work. During these two days, which are of more especial interest in considering the subsequent fasting period, the diet supplied about 11 grams of nitrogen 14 U. S. Dept. Agriculture, Office of Exp. Sta. Bui. 175. 15 The bicycle ergometer is an apparatus for measuring accurately the external muscular work of riding. It is practically a stationary bicycle. For a description, see Carnegie Institution of Washington Publication No. 42, p. 164. 56 Influence of Inanition on Metabolism. and 2100 calories of energy. The diet was extremely simple, consisting of crackers, sugar, and milk. The details of the preceding experiments are given in another publication.18 The subject had undoubtedly become thoroughly accustomed to life inside the respiration calorimeter and to the daily routine during the previous 11 days, consequently the fasting experiment was made under conditions ideal for obtaining minimum muscular activity. No prescribed program was insisted upon. He was provided with reading and writing materials and every care was taken to make him as comfortable as possible. The clothing worn by the subject was essentially the same as that worn by all others, namely, union suit, stockings, trousers, sweater or coat as desired, and slippers or rubber-soled shoes. A. L. L. was a student in the university. In temperament he was rather phlegmatic and slow in speech and movement. His physical strength was above the ordinary. The records of his body measurements are given below. From the date of the anthropometric records, it will be seen that these measurements were taken six months prior to the date of the experiment. Measurements of A. L. L. — Date, Oct. 2Jf, 1903. Age 22 years, 8 months. Weight kilograms. Height centimeters . Length of — Sternum do . . . Navel do . . . Pubis do . . . Sitting do. . . Knees do. . . Shoulders, elbow do . . . Elbow to tip do . . . Arm reach do . . . Right foot do . . . Left foot do . . . Girth of— Head do. . . Neck do. . . Chest- Depressed do . . . Inflated do. . . Normal do . . . At 9th rib, full do... At 9th rib, depressed. .do. . . Waist do. . . Hips do... Right biceps do. . . Left biceps do . . . Right arm do . . . 73.3 166.3 138.9 101 83.4 90.2 44.1 37.5 45.2 175.9 26.8 26.7 57.1 37 93.5 102 99.8 94.5 93.4 81.3 97.7 33.6 33.5 30.2 Girth of— Left arm centimeters Right elbow do . . Left elbow do . . Right forearm do. . Left forearm do . . Right wrist do. . Left wrist do . . Right thigh do . . Left thigh do . . Right knee do . . Left knee do . . Right calf do.. Left calf do.. Right instep do . . Left instep do . . Breadth of — Head do. . Neck do. . Shoulders do . . Chest do. . Waist do. . Hips do. . Depth of— Chest do. . Abdomen do . . 30.5 24.8 24.9 30 29.5 17.3 17.5 56.5 56 37.8 38.2 36.9 36.7 25.9 25.6 15.1 11.4 43.2 28.8 27.3 33.9 21 20.4 10 U. S. Dept. Agriculture, Office of Exp. Sta. Bui. 175. Metabolism Experiment No. 68. 57 The following notes from the diary of the subject are of especial interest as an index of his physical state : Notes from diary. April 27, 1904. Took a lampblack capsule this morning and drank a little water. The day passed quite quickly because of interesting reading matter. Felt a little craving for food in the evening and some weakness. Retired and slept well, waking a little before 7 a. m. April 28, 1904. Felt a definite craving for food about breakfast time, but it soon passed away. Felt weak and lifeless; belched considerable wind. About bedtime I felt unusually weak. This was particularly noticeable because I had not been moving around for some time. After lying down I didn't feel very hungry and soon went to sleep. Was wakeful and restless during the latter part of the night. Routine. — Since it interferes materially with fasting subjects to insist upon a prescribed routine, the subject of this experiment was only cautioned to minimize so far as possible the muscular movements. His experience of the 11 preceding days aided materially in securing uniform experimental conditions during the fasting period. The experimental day began as usual at 7 o'clock in the morning and, since the last meal was taken at 6 o'clock the night before, there had been a fast of 13 hours before the experiment proper began. Body movements. — The usual movements incidental to drinking water, taking pulse, caring for excreta, opening and closing the food aperture, etc., were unavoidable. Although the subject attempted to secure uniform bodily activity on the two days of the experiment, an inspection of the record of body move- ments will show that this was extremely difficult. The major body movements obtained by the physical observer and reported in the diary of the subject are given herewith. Movements of subject. — Duration, tico days, from Apr. 27, 7 a. m., to Apr. 29, 7 a. m., 1904. April 21. A. M. p. M. A. M. 8h 50m telephone. 3" 32m telephone. 7h00m rise. 9 16 telephone. 3 56 telephone. 7 02 weigh self. 9 54 telephone. 4 00 telephone, food ap- 7 04 weigh clothes. 10 02 food aperture. erture, undress, 7 06 weigh absorbers. 10 16 telephone. take temperature. 7 10 urinate. 10 20 food aperture. 4 04 prepare ergometer, 7 12 take temperature. 10 24 undress, prepare mount and dis- 7 20 dress. ergometer. mount, dress, sit. 7 24 sit. 10 28 mount ergometer. 4 20 read. 7 28 food aperture. 10 30 dismount, dress, 5 12 telephone. 7 32 sit. sit. 5 14 read. 7 40 food aperture. 11 48 telephone. 5 36 undress. 8 16 move about, pre- P. M. 5 42 dress. pare ergometer,1 12h 02m food aperture. 5 46 sit, read. undress.2 12 06 telephone. 6 06 telephone. 8 18 mount ergometer. 1 00 urinate. 6 08 food aperture. 8 20 dismount, dress. 3 00 read. 6 10 sit, read. 8 30 sit. 3 30 food aperture. 7 02 food aperture. 1 Places the bicycle ergometer in the position in which it is usually ridden. 2 Record of "undress " during the day indicate that trousers and sweater were removed. 58 Influence of Inanition on Metabolism. Movements of subject. — Continued. April 27 (cont.) A. M. p. M. P. M. i7h 44m telephone. 1' 52m mount. 7h03n" urinate. 7 54 food aperture. 1 54 dismount, dress. 7 04 telephone. 8 20 move about, un- 1 56 sit, read. 7 06 food aperture. dress, prepare er- 3 32 food aperture. 7 08 take temperature. gometer. 4 00 rise, undress, take 7 10 food aperture. 8 24 mount. temperature. 7 12 read. 8 30 dismount, dress. 4 04 food aperture. 7 32 telephone. 8 36 sit, read. 4 06 prepare ergometer, 7 44 read. 10 06 take temperature, mount. 10 00 take temperature. telephone. 4 08 dismount, dress, 10 52 open bed, undress. 10 08 food aperture. sit. 10 56 urinate. 10 10 sit, read. 4 10 read. 11 00 retire. 10 16 food aperture. 4 32 telephone. 10 24 prepare ergometer, 5 28 rise, undress. April 28. mount. 5 30 dress, read. A.M. 10 28 dismount, dress. 6 02 rise, food aperture. 7h 00m rise. 10 30 sit, read. 6 06 telephone. 7 04 weigh self. 10 36 telephone. 6 28 food aperture. 7 06 weigh clothes. 11 10 telephone. 7 00 take temperature. 7 08 weigh absorbers. P. M. 7 02 urinate. 7 10 urinate. lh02m urinate. 7 48 rise. 7 14 take temperature. 1 04 food aperture, take 7 50 sit, read. 7 16 sit. temperature. 8 02 read. 7 20 rise, dress. 1 10 sit, read. 10 04 take temperature. 7 22 read. 1 44 collect drip.1 10 08 read. 7 26 food aperture. 1 50 undress, prepare 10 52 collect drip. 7 32 sit, read. ergometer. 11 00 retire. 1 The water which has accumulated in the drip cans is put into a bottle and passed out through the food aperture. Table 21. — Record of water consumed — Metabolism experiment No. 68. Date. Period during which water was consumed. Total for day. 7 to 10 a. m. 10 a. m. to 1 p. m. 1 to 4 p. m. 1904. Apr. 27-28 Apr. 28-29 Grams. 149.2 330.2 Grams. 119.2 Grams. 286.6 Grams. 268.4 616.2 Drinking-water. — Water for drinking purposes was furnished ad libitum. In order to compute the heat production, the quantity of water consumed during each period of the day (when the record of body movements indicated that water was drunk) was estimated by methods similar to those followed in experiment No. 59. Data for making this apportionment were rather more complete in this experiment than in the previous one. All water was cooled and its temperature taken before it was received by the subject. This is true also of all the experiments following. The amounts obtained by means of these estimates are recorded in table 21. From table 21 it appears that the quantity of water drunk on the second day of the experiment was much larger than that consumed on the first day; in Metabolism Experiment No. 68. 59 fact more than twice as large. Furthermore, all the water was consumed before 4 p. m., which was unusual in these experiments. The quantity of water con- sumed by A. L. L. was considerably less than that drunk by the subject of experiment No. 59. Urine. In this experiment the urine was collected in four periods, from 7 a. m. to 1 p. m., 1 to 7 p. m., 7 to 11 p. m., and 11 p. m. to 7 a. m. This was done in the hope that the analysis of the urine for different periods would throw some light upon the rate of elimination of different ingredients in the urine; for while Table 22. — Determinations in urine per period and per day — Metabolism experiment No. 68. Date. Period. (a) Amount. (b) Specific gravity. (O Volume (a + b). (d) Reaction. (e) Nitro- gen. 19M. Apr. 27-28. Apr. 28-29. 7 a.m. to 1 p.m. 1 p.m. 7 p.m. 7 p.m. 11 p.m. 11 p.m. 7 a.m. Total Grams. 267.5 264.6 172.1 297.5 1.0190 1 . 0220 1.0215 1.0230 c.c. 262 259 168 290 Faintly acid. . Slightly acid. . do Acid Grams. 2.89 3.31 2.08 3.98 1001.7 1001.7 1.0230 979 979 12.26 12.32 Total by com- 7 a.m. to 1 p.m. 1p.m. 7 p.m. 7 p.m. 11 p.m. 11 p.m. 7 a.m. Total 221.9 223.1 I 398.8 1.0225 1.0260 1.0270 217 217 388 3.08 3.47 6.48 do do 843.8 843.8 1.0265 822 822 13.03 13.05 Total by com- posite Total for 2 days. Total by compo- site for 2 days. Acid 1845.5 1845 . 5 1.0255 1801 1801 25.29 25.04 the collections were made more frequently in experiment ISTo. 59, with the increased demands upon the time of the assistants, the analysis of urine for short periods had been impossible. The amounts of urine and the specific gravity, volume, reaction, and nitrogen, determined not only by periods but also in the composite sample, are given in table 22. In addition to the data given in table 22, there were determined on the daily composite samples the heat of combustion and on the composite for the two days the heat of combustion, the water-free material, the carbon, the hydrogen of organic matter, and the ash. The heat of combustion per gram for the urine for April 27-28 was 0.092 calorie. For April 28-29 it was 0.117 calorie. The 2-day composite sample gave 94.94 per cent of water, 1.09 per cent of carbon, 0.25 per cent of organic hydrogen, and 1.22 per cent of ash. GO Influence of Inanition on Metabolism. Weight and composition of urine. — The total weights of material excreted in the urine are given in table 23. These include the total weight of urine, the weight of water, solids, ash, organic matter, nitrogen, carbon, hydrogen in organic matter, and oxygen in organic matter. The heat of combustion is also given. As in experiment Xo. 59, the determinations of carbon and organic hydrogen were made only on the 2-day composite, and hence it was necessary to apportion the total amounts of carbon and organic hydrogen eliminated according to the amount of nitrogen eliminated on the two days. In apportioning the amounts of water and ash, which were also determined only in the 2-day composite, the Table 23. — Weight, composition, and heat of combustion of urine — Metabolism experiment No. 68 (April, WO-'t). Constituents. (a) Weight grams. . (6) Water do . . . (c) Solids, a — b do... (d) Ash i do . . . (e) Organic matter, c — d do . . . (/) Nitrogen do . . . (g) Carbon do . . . {h) Hydrogen in organic matter do. . . (i) Oxygen (by difference) in organic mat- ter, e — {f+g + h) do . . . (j) Heat of combustion calories. . . Apr. 27-28. 1001.7 956.4 45.3 10.91 34.39 12.26 9.75 2.23 10.15 92 Apr. 28-29. 843.8 795.7 48.1 11.6 36.50 13.03 10.37 2.38 10.72 99 Total for 2 days. 1845.5 1752.1 93.4 22.51 70.89 25.29 20.12 4.61 20.87 191 1 The ash and water for the individual days are calculated as shown on page 28. Hence the amounts of solids and organic matter for the individual days are not determined but calculated. same processes were followed as those described in experiment ISTo. 59. The amounts of ash and water for the individual days are affected by the errors in the apportionment of the total solids. Elimination of Water- Vapor. The amount of water withdrawn from the respiration chamber by the venti- lating air-current varies from period to period and depends in large measure upon the absolute amount of water remaining in the chamber at the end of each period and upon the rate of ventilation. Since as was explained in discussing experiment No. 59 (see page 30) the relative humidity for each period may be readily computed, in table 24 are recorded only the total amount of water- vapor in the atmosphere of the calorimeter at the end of each period and the total water of respiration and perspiration. On the two days the amount of water lost in respiration and perspiration was essentially the same, there being 745.3 grams on the first day and 760.7 grams on the second. Metabolism Experiment No. 68. 61 While in experiment No. 59 there was no recorded change due to loss or gain of water in the weight of the heat-absorbers, chair, bedding, and other articles inside the chamber, in this experiment there was a loss of 47 grams of water on the first day and a gain of 21 grams on the second day. The correct apportion- ment of this gain or loss of water by individual periods is impracticable, since the weights are taken only once each day. It has therefore been our custom to inspect the hygrometric conditions in the chamber at the end of each period, and thus ascertain whether there was a continuous drying out process or a condensation and absorption of moisture from the air. An inspection of the figures in column a in table 24 shows that the water in the respiration chamber Table 24. — Record of water of respiration and perspiration — Metabolism experiment No. 68. (a) ■ • • Apr. 27-28: 7 a.m. to 10 a.m. . 313.9 101.1 32.2 110.1 35.1 56.0 70.7 0.79 10 a.m. 1 p.m. . 284.9 76.5 26.9 89.3 31.4 45.5 53.6 .85 1 p.m. 4 p.m. . 275.8 82.3 29.9 90.3 32.8 46.0 57.6 .80 4 p.m. 7 p.m. . 303.2 89.8 29.6 93.7 30.9 47.7 62.9 .76 7 p.m. 10 p.m. . 287.9 84.8 29.5 90.2 31.3 45.9 59.4 .77 10 p.m. 1 a.m. . 268.9 74.9 27.9 81.4 30.3 41.4 52.4 .79 1 a.m. 4 a.m. . 219.0 65.8 30.0 67.7 30.9 34.5 46.0 .75 4 a.m. 7 a.m. . Total 213.8 04 . S 30.3 71.6 33.5 36.5 45.4 .80 2167.4 640.0 29.5 694.3 32.0 353.5 448.0 .79 Apr. 28-29: 7 a.m. to 10 a.m . . 290.0 99.4 34.3 98.9 34.1 50.4 69.6 .72 10 a.m. 1 p.m. . 300.6 83.8 27.9 90.2 30.0 45.9 58.7 .78 1 p.m. 4 p.m. . 300.3 83.7 27.9 88.7 29.6 45.2 58.6 .77 4 p.m. 7 p.m. . 292.3 86.2 29.5 92.4 31.6 47.0 60.3 .78 7 p.m. 10 p.m. . 281.0 81.6 29.1 87.7 31.2 44.6 57.1 .78 10 p.m. 1 a.m. . 263.8 71.8 27.2 80.4 30.5 40.9 50.2 .82 1 a.m. 4 a.m. . 238.3 69.0 28.9 70.0 29.4 35.7 48.3 .74 4 a.m. 7 a.m. . Total 251 . 1 67.1 26.7 71.0 28.3 36.1 47.0 .77 2217.4 642.6 29.0 679.3 30.6 345.8 449.8 .77 The same discrepancies which have been pointed out before as obtaining for the determinations of oxygen, heat, and carbon dioxide during short periods vitiate the accuracy of the ratios for individual periods, yet the results for 24 hours are probably accurate. The oxygen thermal quotient remains nearly constant for the two days, falling from 29.5 on the first day to 29.0 on the sec- ond day. The carbon dioxide thermal quotient indicates a somewhat greater fall, varying from 32.0 on the first day to 30.6 on the second day. The respir- atory quotient, although indicating marked fluctuations in the different periods, on the average for the 24 hours is 0.79 for the first clay and 0.77 for the second. Metabolism Experiment No. 69. 71 Metabolism Experiment No. 69. This experiment, which continued for 4 days, was made with the same subject as experiment No. 68, but 7 months later. The subject entered the respiration chamber on the evening of December 15, 1901, the experiment proper beginning at 7 a. m., December 16. Preliminary practice in adjustment of furniture and apparatus inside the calorimeter chamber was not necessary, as this subject had previously participated in a large number of experiments. He was also thoroughly accustomed to the environment. The usual clothing was worn and the general plan of the whole experiment was similar to that of experiment No. 68. It has already been suggested that A. L. L., the subject of this experiment, was of phlegmatic temperament. The following notes from his diary contain practically everything he wrote, and so brief are they that their value for deduction as to the physical or psychical condition of the subject a** practically worthless. They simply serve to indicate that the stay in the calorimeter was not distasteful to the subject and that he felt little ill effect from his fast. IS Notes from diary. Dec. 16, 1904: Lay down this morning at 9 and re- mained on the bed until 1 p. m. Slept the greater part of the time, but with disturbed dreams during the last of it. Dec. 17, 1904: While adjusting the heat absorbers this morning, I felt a little faint. Have had no desire for water, but have drunk as ordered.20 Dec. 18, 1904: Nothing worthy of note regarding this day. Dec. 19, 1904: Day passed quite as the rest. Pulse. — In this experiment the subject counted his pulse twice each day. The count was made for two minutes and the average number of beats per minute is recorded below. Time. Pulse rate. Time. Pulse rate. Dec. 16, 1904, lh 00" P- m. . 55 Dec. 18, 1904, 7h45ma. m. . 62 8 15 P- m. . 52 11 00 P- m. . 48 Dec. 17, 1904, 9 50 a. m. . 57 Dec. 19, 1904, 8 30 a. m. . 62 3 00 P. m. . 62 | 10 20 P- m. . 56 Routine. — Aside from conforming to the general experimental period so far as hours of sleep, rising, collection of urine, etc., was concerned, the subject was allowed to do as he pleased except that he was cautioned to secure, in so far as possible, uniform muscular activity on the several days of the experiment. 20 Owing to the unusually high specific gravity of the urine of this subject in previous experiments he was requested to drink liberal amounts of water. 72 Influence of Inanition on Metabolism. Body movements. — A careful record was kept of the bodily activity, and for the better interpretation of the results it is given herewith. Movements of subject. -Duration, k days, from Dec. 16, 7 a. m., to Dec. 20, 7 a. m., 1904. December 16. A. M. P. M. A. M. gh Q4m telephone. lh02m read. 7h00m rise. 8 06 read. 1 20 food aperture. 7 02 weigh self. 8 36 stop reading. 2 24 stop reading. 7 04 weigh absorbers. 9 06 lie, read. 2 26 write. 7 06 dress. 9 50 count pulse. 6 26 telephone. 7 10 urinate. 10 36 telephone, food ap- 6 32 food aperture. 7 18 fold bed. erture. 7 02 urinate. 7 20 adjust table, sit. 10 42 lie. 7 06 food aperture. 7 24 read, sit. 11 30 stop reading. 7 08 read. 8 06 telephone. 11 36 asleep. 9 02 read. 8 08 read. P. M. 9 38 asleep. 9 08 recline, read. lh 02m rise, urinate, food 11 00 count pulse. 9 16 stop reading. aperture. 11 02 urinate, retire. 9 36 asleep. 1 04 sit, read. December 19. P. M. 3 00 count pulse. A. M. 12h 52m awake, read. 3 54 telephone. 7h 00m rise. 1 00 telephone. 3 56 drink. 7 04 weigh self. 1 02 rise, urinate, sit. 4 04 telephone. 7 06 weigh absorbers. 1 10 food aperture. 5 06 close curtain. 7 10 urinate. 1 24 count pulse. 5 10 open curtain, sit. 7 12 dress. 1 30 write. 5 14 telephone. 7 15 food aperture. 2 26 telephone. 5 16 read. 7 16 sit. 2 28 food aperture. 6 24 food aperture. 7 25 food aperture. 2 30 read. 6 28 read. 8 10 telephone. 4 22 telephone. 7 02 urinate. 8 14 food aperture. 4 28 telephone. 7 06 read. 8 20 write. 4 30 read. 11 08 urinate. 8 30 count pulse. 7 00 urinate. 11 10 retire. 8 37 food aperture. 7 02 7 04 read, telephone. December IS. 8 44 9 08 read, lie. 7 05 food aperture. A. M. 7h02n' 7 04 rise. 9 44 asleep. 7 06 read. weigh self, weigh absorbers. 11 02 telephone. 8 15 count pulse. 7 06 11 04 sit, food aperture. 11 00 11 06 urinate, retire. 7 12 urinate, food aper- ture. 11 06 P. M. lie, read. December 11. 7 14 dress. lh02" rise, urinate, food A. M. 7 24 read. aperture. 7h00" ' rise. 7 45 count pulse. 1 04 sit, read. 7 02 weigh self. 9 10 lie, read. 2 23 food aperture. 7 04 weigh absorbers. 10 30 telephone. 3 54 telephone. 7 08 urinate. 10 32 sit, food aperture. 7 04 telephone, urinate, 7 10 dress. 10 40 lie, read. food aperture. 7 16 sit. P. M. 7 06 read. 7 20 read. 12h 58m rise, sit. 10 20 count pulse. 7 30 food aperture. 1 00 urinate, food aper- 11 00 urinate, undress, 7 32 read. ture. retire. Drinking-water. — During this experiment the subject did not appear to care for water and would have drunk very little of his own accord. In previous experiments he had shown a similar tendency, and in some instances the quantity of urine voided had been very small and of a high specific gravity. Metabolism Experiment No. 69. 73 Accordingly the subject was requested to drink considerable amounts of water and no doubt consumed more than he desired, though it gave him no discomfort. The daily amounts consumed and the estimates for the water consumed by periods during the experiment are given in table 33. In estimating the quantity of water consumed during each of the periods for which the amounts were not definitely shown, it was assumed that after a weighed amount of water was placed in the calorimeter chamber equal amounts were drunk during each 2-hour period until all this water was consumed. For example, it was known that between the hours of 1 and 7 p. m. on December 16 the subject drank 4S9.10 grams of water. These 6 hours constituted three experimental periods and it was assumed that one-third of this amount (163.03) grams of water was taken during each period. While the apportion- ment of the quantities for the different periods is at best but roughly approxi- Table 33. — Record of water consumed — Metabolism, experiment No. 69. Date. Period during which water was consumed.1 Total for day. 7 to 9 a. m. 9 to 11 a. m. 11 a. m. to 1p.m. 1 to 3 p. m. 3 to 5 p. m. 5 to 7 p. m. 7 to 9 p. m. 9 to 11 p.m. 1904. Dec. 16-17... Dec. 17-18. . . Dec. 18-19... Dec. 19-20... Grams. 38.33 166.04 148.00 136.37 Grams. 148.00 136.37 Grams. 166.03 148.00 136.36 Grams. 163.04 163.00 131.80 Grams. 163.03 166.03 163.00 Grams. 163.03 55.50 162.90 131.80 Grams. 119.60 76.20 77.90 Grams. 76^20 77.90 Grams. 647.03 553.60 1085.30 828.50 1 Assumed in some instances. (See above.) mate, it must be borne in mind that the absolute amount of water taken during the day was determined with accuracy. A similar apportionment has been made of the water consumed on the subsequent days of the experiment. Urine. As in experiment No. 68, the urine was collected 4 times daily. In the samples of urine thus collected a number of determinations were made which are recorded in table 34. As a check on the nitrogen determinations the total nitrogen was determined in the daily composite as well as by periods. Weight and composition of urine. — In addition to the determinations of nitrogen, specific gravity and reaction made upon urine collected by periods, there were determined in the daily composite the water, total solids, ash, nitrogen, carbon, hydrogen in organic matter, phosphorus, sulphur, and heat of combustion. From the weight of urine and the percentages obtained in the above determinations, the total amounts of each ingredient analyzed were computed. These amounts are recorded in table 35. While in experiments Nos. 59 and 68 determinations of ash, carbon, and hydrogen were made only on the composite urine for the whole experiment, in 74 Influence of Inanition on Metabolism. this experiment the determinations were made on each daily sample, thus avoiding the complicated and somewhat erroneous calculations for the distri- bution or apportionment of the water, ash, solids, carbon, and hydrogen over the individual days of the experiment. Table 34. — Determinations in urine per period and per day — Metabolism experiment No. 69. Date. Period. (a) Amount. Specific gravity. (c) Volume (a-t-b). (d) Reaction. (e) Nitro- gen. 1904. Dec. 16-17. Dec. 17-18. Dec. 18-19. Dec. 19-20. 7 a.m. to 1 p.m 1 p.m. 7 p.m 7 p.m. 11 p.m Total Total by composite. . . 1 p.m. 7 p.m 7 p.m. 11 p.m Total Total by composite. . . 7 a.m. to 1 p.m 7 p.m. 11 p.m 11 p.m. 7 a.m Total Grams. 115.4 123.3 63.1 150.0 1.0320 1.0330 1 . 0360 1.0335 c. c. 112 119 61 145 Acid. ..do.. ..do.. ..do.. . . . . Grams. 2.45 2.51 1.52 3.61 451.8 451.8 1.0330 437 437 10.09 9.95 174.4 167.9 92.2 155.6 1.0280 1.0280 1.0325 1.0325 169 163 89 151 Acid. . . do. . 3.94 3.80 2.39 4.13 . . do. . ..do.. . . . . 590.1 590.1 1.0310 572 572 14.26 14.34 255.9 208.2 107.9 190.3 1.0205 1.0255 1.0265 1.0270 250 203 105 185 Acid. ..do.. 4.32 4.03 2.34 4.35 ..do.. ..do.. 762.3 762 . 3 1.0250 743 743 15.04 15.03 Total by composite. . . 1 p.m. 7 p.m 7 p.m. 11 p.m Total , 250.2 227.9 100.9 163.1 1.0195 1.0215 1.0265 1.0280 245 223 98 159 Acid. ..do.. ..do.. ..do.. • • • 4.12 3.47 2.23 3.15 742.1 742.1 1.0230 725 725 12.97 13.18 Total by composite. . . Total, 4 days 2546.3 2477 52.36 The determinations of phosphorus were made by fusion with sodium peroxide and not by titration with uranium salts. For purposes of further comparison the quantity of phosphorus is expressed in line ;' of table 35 as the element and in line h as phosphoric acid, i. e., phosphorus pentoxide. The amounts of sulphur are likewise expressed both as sulphur and as sulphur trioxide. Elimination of Watee-Vapob. The amounts of water-vapor remaining in the chamber at the end of each 2-hour period are recorded in table 36, together with the amounts of water of respiration and perspiration. On the preliminary night, the residual amounts Metabolism Experiment No. 69. 75 were determined in two periods of 3 hours each immediately preceding the experiment. In this experiment as in experiment No. 68, there were certain changes in the weights of the chair, bed, bedding, etc. The amounts of water involved in these changes are noted at the end of the table. While inspection of the humidity conditions in this experiment showed that the losses must have been fairly regular for the first, third, and fourth days, the gain on the second day apparently occurred during the last 4 periods of the day and it was so apportioned. Table 35. — Weight, composition, and heat of combustion of urine — Metabolism experiment No. 69. Dec. 16-17. Dec. 17-18. Dec. 18-19. Dec. 19-20. Total for 4 days. {a) Weight grams . . (&) Water do (c) Solids, a — b do (d) Ash do (e) Organic matter, c—d do. . . . (/) Nitrogen do {* i . 7 04 sit, write. 3 16 write. 12" 08m telephone. 7 20 food aperture. 3 40 sit. 12 46 rise, sit. 7 8 32 00 lie. count pulse. 3 4 42 18 write. food aperture. 1 02 telephone, rise, uri- nate, move about. 9 15 asleep. 4 20 drink, sit. 1 04 sit. 11 00 rise. 4 26 read. 1 30 count pulse. 11 02 open bed. 5 04 rise, move about. 1 32 lean on table, 11 04 urinate, undress, 5 08 lean on table. write. retire. 5 10 telephone. 2 30 write. January 9. 5 12 write. 3 20 food aperture. A. M. 5 30 count pulse, write. 3 22 lean on table, lh 30m rise, count pulse, 5 32 telephone. write. write. 6 00 food aperture. 3 40 telephone. 7 00 rise, urinate. 6 16 sit. 4 02 sit. 7 04 begin weighing. 7 00 rise, urinate. 4 04 stand. 7 11 finish weighing. 7 04 sit, read. 4 14 move about, drink. 7 22 close curtain. 7 30 write. 4 22 sit, write. 7 24 defecate. 8 00 food aperture, 4 24 read. 7 25 food aperture. count pulse. 5 04 rise, stretch arms. 11 -J Influence oe Inanition on Metabolism. Movements of subject. — Continued. January 10 (cont). P. M. p. M. P. M. 7" 00m rise. 9h 00m lean on table, read 5" 06m drink. 7 02 take temperature. 9 12 stand. 5 16 lean on table, 7 04 urinate. 9 14 arrange bed. write. 7 08 food aperture. 9 16 lie. 5 26 telephone. 7 10 lean on table, read. 11 00 rise. 5 30 count pulse, write. 7 12 telephone. 11 02 close curtain, un 5 54 move about. 7 15 write. dress, urinate, re 5 58 sit, read. 8 08 sit, read. tire. Drinking-water. — Drinking-water was furnished from the city supply and the subject drank much larger amounts than any previous subject. The quantities for each day, apportioned as nearly as possible among the different experimental periods, are given in table 65. Great differences in the amount of water consumed daily may be observed. While about 1200 grams were con- sumed on the first day of the fast, on the second and third days the amount consumed per day averaged more than 2 liters. On these 2 days a considerable Table 65. — Record of water consumed'1 — Metabolism experiment No. 71. Date. 7 to 9 a. m. 9 to 11 a. m. 11 a. m. to 1p.m. 1 to 3 p. m. 3 to 5 p.m. 5 to 7 p. m. 7 to 9 p. m. 9 to 11 p. m. Total for day. 1905. Jan. 7-8 .... Jan. 9-10 Jan. 10-11 Grams. 202.05 397.20 404.50 384.60 Grams. 202.05 393.10 3S9.10 395 . 60 Grams. 198.60 397.20 197.20 194.00 Grams. 198.60 197.40 197.20 194.00 Grams. 201.55 197.40 391.60 158.55 Grams. 201.55 194.80 158.55 Grams. 195.45 194.80 Grams. 195.45 343.70 Grams. 1204.40 1973.20 2312.90 1485.30 1 Period during which water was consumed was assumed in some instances. amount of water was taken between 7 and 11 p. m. On the first and last days no water was consumed after 7 p. m. Urine. The urine was collected as usual at the end of each of the 4 periods, but on the first day the subject accidentally mixed the urine of the first and second periods. Determinations were made of the specific gravity, reaction, and nitrogen of the urine for each period. These are recorded in table 66. The quantity of urine voided is nearly proportional to the amount of water drunk, there being on the third day over 2.5 liters passed. Throughout the experiment the specific gravity was low and the reaction acid. Weight, composition, and heat of combustion of urine. — Aside from the determinations given in table 66 on the samples of urine for the 4 usual periods, each daily composite sample was analyzed. Determinations were made of the water, total solids, ash, nitrogen, carbon, organic hydrogen, phosphorus, sulphur, and heat of combustion. From the percentages thus obtained, the heat of bustion per gram and the weight of urine, the quantities of the various elements and the total heat of combustion are computed. These are recorded Metabolism Experiment No. 71. 113 in table 67. As in experiments Xos. 69 and 70 the results were obtained from actual determinations made in duplicate in all cases, and in many cases in triplicate. The phosphorus and sulphur determinations are expressed not only in terms of elements but also as oxides. Both elements were determined by the fusion method. Table 66. — Determinations in urine per period and per day — Metabolism experiment No. 71. Date. Period. (a) Amount. (&) Specific gravity. (c) Volume (a + b). (d) Reaction. (e) Nitro- gen. 1905. Jan. 7-8 Jan. 8-9 Jan. 9-10.. . Jan. 10-11.. 7 a.m. to 1 p.m. 1 p.m. 7 p.m. 7 p.m. 11 p.m. 11 p.m. 7 a.m. Total Grams. 1 9S5 . 6 65.5 108.4 1.0072 1.0153 1.0215 c.c. 979 64 106 Slightly acid. Grams. 3.29 0.77 1.78 . . . .do 1159.5 1159.5 1.00S9 1149 1149 5.84 5.82 Total by com- 7 a.m. to 1 p.m. 1 p.m. 7 p.m. 7 p.m. 11 p.m. 11 p.m. 7 a.m. Total Total by com- 7 a.m. to 1 p.m. 1 p.m. 7 p.m. 7p.m. 11p.m. 11 p.m. 7 a.m. Total . 830.4 550.6 307.5 332.6 1 . 0042 1.0061 1.0058 1.0101 827 547 306 329 3.04 2.74 1.94 3.32 . . . .do ... .do . . . .do 2021.1 2021.1 1.0058 2009 2009 11.04 11.02 Acid Acid 753 . 2 608.7 443.8 735.9 1.0046 1.0073 1.0048 1 . 0054 750 605 442 732 3.46 3.47 2.10 4.07 do do ... .do 2541.6 2541.6 1.0048 2529 2529 13.10 13.16 Total by com- 7 a.m. to 1 p.m. 1 p.m. 7 p.m. 7 p.m. 11 p.m. 11 p.m. 7 a.m. Total Acid do do do 644.0 522 . 5 105.9 213.4 1.0052 1.0067 1.0190 1.0173 640 519 104 210 3.36 2.85 1.47 3.06 1485.8 1485.8 1.0087 1473 1473 10.74 10.67 Total by corn- Total, 4 days... 7208.0 7160 40.72 Elimination of Water- Vapor. The water of respiration and perspiration, together with the amount of water-vapor residual in the chamber at the end of each 2-hour period, is recorded in table 68 (see page 115). The daily amounts of water gained or lost by articles in the chamber are given in the foot-note to the table. This water was apportioned in equal amounts among all the 2-hour periods of the day. 8 114 Influence of Inanition on Metabolism. An inspection of the figures in column a shows that the relative humidity in the chamber steadily decreased during the 4 days, thus explaining the continual loss of moisture from the bedding and furniture. The total water of respiration and perspiration diminished from day to day, amounting to 744.84 grams on the first day and 517.66 grams on the fourth day. Cutaneous excretion of nitrogenous material. — During this experiment and experiment No. 72 immediately following there was an average elimination of nitrogen of 0.029 gram per day found in the perspiration. Table 67. — "Weight, composition, and heat of combustion of urine — Metabolism experiment No. 71. (a) Weight grams . . (6) Water do (c) Solids (a— b) do (d) Ash do (e) Organic matter (c — d) do (/) Nitrogen do (g) Carbon do (A) Hydrogen in organic matter, grams (i) Oxygen (by difference) in organic matter, e— {f+g + h) . . .grams.. ( j) Phosphorus do ... . (k) Phosphoric acid by fusion (P2 05), grams (I) Sulphur grams. . (m) Sulphur trioxide (S03) . . . .do. . . . (n) Heat of combustion. . . .calories. . Jan. 7-8. Jan. 3-9. Jan. 9-10. Jan. 10-11. Total for 4 days. 1159.5 2021.1 2541.6 1485.8 7208.0 1133.99 1984.11 2500.68 1449.25 7068.03 25.51 36.99 40.92 36.55 139.97 7.65 6.06 5.85 5.94 25.50 17.86 30.93 35.07 30.61 114.47 5.84 11.04 13.10 10.74 40.72 5.22 8.29 8.64 7.73 29.88 1.16 2.02 2.29 2.08 7.55 5.64 9.58 11.04 10.06 36.32 .322 .579 .873 .889 2.663 .736 1.326 2.000 2.038 6.100 .519 .711 .758 .670 2 . 658 1.298 1.774 1.893 1.671 6.636 58 91 99 86 334 Elimination of Cakbon Dioxide and Absorption of Oxygen. From the analyses of the respiratory gases, the quantities of carbon dioxide and oxygen residual in the chamber at the end of each period are obtained. These are recorded in columns a and c of table 69. The weights of carbon dioxide exhaled and the amounts of oxygen consumed by the subject are shown in the same table in columns b and d, respectively. There are especially noticeable differences in the absolute amounts of carbon dioxide present in the chamber, both from one period to another and from day to day. The residual amounts tend to diminish as the experiment progresses, while on the other hand the amounts of oxygen tend to increase. For instance, the residual oxygen varies from 831.9 liters at 11 a. m., January 7, to 1090.2 liters at the end of the experiment. Corresponding to these wide differences in the amounts of oxygen, there are of course variations in the absolute percentages of oxygen in the air, but as has been shown by Zuntz, Durig, and Metabolism Experiment No. 71. 115 others,28 the respiratory exchange is unaffected by a diminished percentage of oxygen unless that percentage falls below 11. In none of the experiments thus Table 68. — Record of water of respiration and perspiration- No. 11. -Metabolism experiment (a) (ft) (a) (&) Total Total Total Total amount water of amount water of Date and period. of vapor respira- Date and period. of vapor respira- in tion in tion chamber and chamber and at end perspira- at end perspira- of period. tion.1 of period. tion.1 1905. 1905. Jan. 7: Grams. Grams. Jan. 9-10: Grams. Grams. Preliminary : 1 a.m. . . . 43.0 ... 34.5 52.0 40.6 64.0 31.4 51.0 3 a.m. 5 a.m. 37.4 59.7 30.1 45.0 38.3 62.3 28.9 48.7 Total .... 186.0 30.2 29.8 30.9 48.3 52.1 46.9 Jan. 7-8: 7 a.m. to 9 a.m 45.6 81.6 28.7 48.1 49.5 81.2 28.6 44.2 43.8 69.9 27.7 44.7 43.1 64.3 3 a.m. 5 a.m 28.0 43.5 3 p.m. 5 p.m 5 p.m. 7 p.m 7p.m. 9p.m 9 p.m. 11 p.m 41.3 42.3 39.5 33.9 70.1 66.7 70.5 51.1 Total 26.8 43.1 567.6 Jan. 10-11: 33.1 55.3 30.6 51.6 33.7 50.7 27.5 42.6 37.8 33.4 27.8 42.5 33.9 50.0 29.9 43.8 28.4 28.1 28.1 43.3 48.4 42.8 Total .... 744.8 Jan. 8-9: 7 a.m. to 9 a.m 42.4 82.5 23.5 39.7 9 a.m. 11 a.m 39.6 67.9 26.3 41.6 11 a.m. 1 p.m 35.6 54.5 25.6 43.3 1p.m. 3 p.m 34.0 58.2 27.4 40.6 3 p.m. 5 p.m 5 p.m. 7p.m 7 p.m. 9 p.m 31.5 30.8 31.1 50.5 55.7 52.7 Total 23.7 37.5 517.7 9 p.m. 11 p.m 27.7 49.2 11 p.m. 1 a.m 29.9 49.1 1 a.m. 3 a.m , 27.6 48.9 3 a.m. 5 a.m 28.9 44.6 5 a.m. 7 a.m Total 28.7 51.4 665.2 1 Allowance has been made for water lost by the absorber, chair, bedding, and miscellaneous articles as follows : Jan. 7-8, 24.45 grams ; Jan. 8-9, 53.45 grams ; Jan. 9-10, 31.78 grams ; Jan. 10-11, 37.01 grams. far made has the percentage of oxygen remaining in the chamber approached this latter point, and hence the atmosphere breathed by the subjects of these experiments has not influenced the character or amount of the respiratory exchange. S9U. S. Dept. Agriculture, Office of Exp. Sta. Bui. 175. 116 Influence of Inanition on Metabolism. Table 69 — -Record of carbon dioxide and oxygen — Metabolism experiment 2s0. 71. Carbon dioxide. Oxygen. (a) (6) (c) (d) Date. Period. Amount in Total Amount in Total chamber weight chamber amount at end of exhaled at end of consumed period. by subject. period. by subject. 1905. Preliminary : Grams. Grams. Liters. Grams. Jim. 7. . . 1 a. m 32.9 918.2 1 a. m. to 3 a. m. . 28.8 42^7 907.3 35 A 3 a. m. 5 a. m. . 30.6 44.7 887.1 28.9 Jan. 7-8 5 a. m. 7 a. m. . Total 34.4 51.8 863.9 48.7 139.2 113.0 7 a. m. to 9 a. m. . 41.5 86.1 845.9 77.3 9 a. m. 11a. m. . 56.9 105.2 831.9 71.5 11 a. m. 1 p. m. . 41.0 61.6 843.3 51.6 1 p. m. 3 p. m. . 39.3 53.4 846.9 50.6 3 p. m. 5 p. m. . 33.1 51.3 858.4 45.6 5 p.m. 7 p. m. . 34.5 51.7 861.9 47.9 7 p. m. 9 p.m.. 35.3 60.6 871.0 56.0 9 p. m. 11 p. m. . 24.9 39.1 888.1 36.5 lip. m. 1 a. m. . 23.2 39.9 892.2 35.0 1 a. m. 3 a. m. . 24.6 41.1 899.6 37.1 3 a. m. 5 a. m. . 35.1 38.5 927.3 40.2 Jan. 8-9 5 a. m. 7 a. m. . Total 27.7 40.5 948.4 39.8 669.0 .... 1 589 . 1 7 a. m. to 9 a. m . . 35.1 74.7 942.8 74.1 9 a. m. 11 a. m. . 30.2 54.7 951.4 67.7 11 a. m. 1p.m.. 30.5 53.3 963.0 46.5 1 p. m. 3 p. m. . 29.0 52.7 971.5 54.7 3 p. m. 5 p. m. . 23.6 39.1 993.0 31.9 5 p. m. 7 p. m. . 24.4 48.5 1003.5 48.4 7 p. m. 9 p. m . . 26.1 50.3 1015.9 52.6 9 p. m. lip. m. . 19.9 36.8 1035.7 33.0 11 p. m. 1 a. m. . 21.2 41.2 1054.3 38.3 1 a. m. 3 a. m. . 20.4 39.4 1053.5 39.7 3 a. m. 5 a. m . . 22.7 40.1 1060.4 33.6 Jan. 9-10 5 a. m. 7 a. m. . Total 21.9 39.4 1070.2 33.6 570.2 554.1 7 a. m. to 9 a. m. . 34.6 62.7 1070.4 66.5 9 a. m. 11 a. m. . 26.8 44.5 1078.3 44.5 11 a. m. 1 p. m. . 28.2 47.2 1074.8 42.6 1 p. m. 3 p. m. . 28.3 53.2 1078.2 49.7 3 p. m. 5 p. m. . 30.1 51.4 1070.7 50.6 5 p. m. 7 p. m. . 28.7 51.1 1074.6 48.5 7 p. m. 9 p. m. . 29.3 46.2 1069.6 47.5 9 p. m. 11 p. m. . 26.3 57.4 1059.8 56.2 lip. m. 1 a. m. . 22.0 36.3 1051.9 32.3 1 a. in. 3 a. m . . 19.7 33.5 1049.9 32.1 3 a. m. 5 a. m. . 23.3 37.2 1041.7 36.7 5 a. m. 7 a. m . . Total 19.5 33.3 1039.6 30.8 .... 554.0 538.0 Metabolism Experiment No. 71. 117 Table 69. — Record of carbon dioxide and oxygen — Continued. Date. Period. Carbon dioxide. Oxygen. (a) Amount in (6) Total (c) Amount in Total chamber weight chamber amount at end of exhaled at end of consumed period. by subject. period. by subject. 1905. Grams. Grams. Liters. Grams. Jan. 10-11. .. 7 a. m. to 9 a. m. . 27.7 56.8 1034.6 55.1 9 a. m. 11 a. m. . 26.9 45.4 1038.0 41.8 11 a. m. 1 p. m. . 29.0 46.5 1037.0 41.4 1 p. m. 3 p. m. . 29.9 48.1 1038.8 53.5 3 p. m. 5 p. m. . 28.2 47.1 1048.7 46.5 5 p. m. 7 p. m. . 23.0 44.3 1062.4 42.9 7 p. m. 9 p. m. . 26.9 46.6 1066.0 45.1 9 p. m. 11 p. m. . 20.0 37.2 1074.4 36.0 11 p. m. 1 a. m . . 20.3 35.4 1074.2 32.2 1 a. m. 3 a. m . . 18.9 33.2 1080.4 36.9 3 a. in. 5 a. m. . 19.5 31.8 1083 . 1 34.1 5 a. m. Total . 7 a. m. . 19.0 35.7 1090.2 27.2 .... 508.1 492.7 That the amount of carbon dioxide eliminated is greatest on the first day is largely due to the extra muscular exercise taken during the second period. During this period it is seen that there were 105.2 grams of carbon dioxide eliminated, while for the same period on the subsequent days from 45 to 55 grams were exhaled. The minimum carbon dioxide elimination was reached on the last day. The largest consumption of oxygen occurred on the first day and the smallest on the last day. The marked increase in the absorption of oxygen that would naturally be expected as a result of extra muscular work is not apparent during the second period of the first day. The difficulty of determining the respiratory quotient and especially the oxygen consumption for short periods during severe muscular work has previously been pointed out.26* In general, the falling off in the consumption of oxygen from day to day is roughly proportional to the decreasing quantities of carbon dioxide eliminated. Elements Katabolized in the Body. The amounts of elements katabolized, as computed from the result of the various elementary analyses, are given in table 70. Elements and materials katabolized in the body. — The elements katabolized from the body and the amounts of katabolized materials (computed by means of the formulas) are given for each day in table 71. Of especial significance 'U. S. Dept. Agriculture, Office of Exp. Sta. Bui. 175. 118 Influence of Inanition on Metabolism. is the marked fluctuation of the protein katabolized, the quantity for the first day being very small, i. e., 35.04 grams. The usual large katabolism of carbo- hydrates takes place on the first day, while on the 3 succeeding days the loss is much less and very constant. Table 70. — Elements katabolized in body — Metabolism experiment No. 71. (a) Total weight. (6) Nitro- gen. (c) Carbon. (d) Hydro- gen. (e) Oxygen. Ash. First day, Jan. 7, 1905. Income: Oxygen from air Solids in urine Grams. 589 . 12 Grams. Grams. Grams. Grams. 589.12 Grams. 1133.99 25.51 744.84 669 . 04 5^84 5.22 182 ! 45 126.89 1.16 83.35 1007.10 5.64 661.49 486.59 7.65 Total Loss Second day, Jan. 8, 1905. Income: Oxygen from air Outgo: Water in urine 2573.38 1984.26 5.84 5.84 187.67 187.67 211.40 211.40 2160.82 1571.70 7.65 7.65 554.11 .... 554.11 .... 1984.11 36.99 665 . 19 570.17 ii!o4 8~29 155.51 222.02 2.02 74.43 1762.09 9.58 590 . 76 414.66 6^06 Solids in urine Water of respiration 1 Carbon dioxide Total. . 3256 . 46 2702.35 11.04 11.04 163.80 163.80 298.47 298.47 2777.09 2222.98 6.06 6.06 Loss Third day, Jan. 9, 1905. Income: Oxygen from air Outgo: Water in urine 537.97 .... 537.97 .... 2500.68 40.92 567.66 553.97 13' 10 8'64 15i:69 279.83 2.29 63.52 2220 . 85 11.04 504 . 14 402 . 88 5^85 Solids in urine Water of respiration l Carbon dioxide Total 3663.23 3125.26 13.10 13.10 159.73 159.73 345 . 64 345.64 3138.91 2600.94 5.85 5.85 Fourth day, Jan. 10, 1905. Outgo: Water in urine 492.74 492.74 .... 1449.25 36.55 517.66 508.13 10 '74 7^73 138! 58 162.17 2.08 57.93 1287.08 10.06 459 . 73 369 . 55 5^94 Solids in urine Water of respiration 1 Carbon dioxide Total 2511.59 2018.85 10.74 10.74 146.31 146.31 222 . 18 222 . 18 2126.42 1633.68 5.94 5.94 Loss 1 Includes also water of perspiration. Balance of water. — The usual method of obtaining the actual loss of water to the body was followed in this experiment. While the quantity of urine voided results in a large elimination of preformed water, this output is nearly com- pensated by the unusually large amounts of water consumed. The amounts lost Metabolism Experiment No. 71. 119 (column f, table 72) were fairly constant save on the last day of the experiment. The data are given in table 72. Changes in Body-Weight Compared with Balance of Income and Outgo. As in preceding experiments, actual weighings were used to check the computed losses of body material. From the records of the weights of drinking- Table 71. — Elements and materials katabolized in body — Metabolism experiment No. 71. Date. (a) Nitro- gen. (6) Carbon. (c) Hydro- gen. (d) Oxy- gen. (e) Water. Protein. <0) Fat. (70 Carbo- hydrates las gly- cogen). («) Ash. 1005. Jan. 7-8 Jan. 8-9 Jan. 9-10 Jan. 10-11 Total, 4 days. Grams. 5.84 11.04 13.10 10.74 Grams. 187.67 163.80 159.73 146.31 Grams. 211.40 298.47 345.64 222.18 Grams. 1571.70 2222.98 2600.94 1633.68 Grams. 1643.94 2449.08 2877.07 1791.12 Grams. 35.04 66.24 78.60 64.44 Grams. 116.53 152.29 142.90 133.01 Grams. 181.64 29.69 22.04 25.33 Grams 7.65 6.06 5.85 5.94 40.72 657.51 1077.69 8029.30 8761.21 244.32 544.73 258.70 25.50 Table 72. — Distribution of intake and outgo of water- No. 71. -Metabolism experiment Date. 1905. Jan. 7-S Jan. 8-9 Jan. 9-10 . . . Jan. 10-11 . . . Total for 4 days. Average per day. Outgo from the body. (a) Water of urine. Grams. 1134.0 1984.1 2500.7 1449.2 7068.0 1767.0 (b) Water of respira- tion and perspira- tion. (c) Total (a+b). Grams. 744.8 665.2 567.7 517.7 2495 . 4 623.8 Grams. 1878.8 2649.3 3068.4 1906.9 9563.4 2390.8 balance of preformed water. (d) Pre- formed (katabo- lized) water in outgo. Grams. U643.9 12449.1 12877.1 1791.1 8761.2 2190.3 (e) Intake in drink. (f) Loss of pre- formed water (cl-e). Water of oxida- tion of organic hydro- gen (c-d). Grams. Grams. 1204.4 1439.5 1973.2 1 475.9 2312.9 !564.2 1485.3 305.8 6975.8 1785.4 1744.0 446.3 Grains. 234.9 200.2 191.3 175.8 802.2 200.5 1 Does not include water of feces. (See p. 122.) water and oxygen consumed, the total income is computed and recorded in line c of table 73. The outgo requires special consideration. Contrary to the ex- perience of the subjects of experiments Nos. 59, 68, and 69, the subject of this experiment defecated on each of the first 3 days. The large amount of vege- table material in his diet affected peculiarly the passage of feces,27 as is espe- 27 For discussion of this point, see Section on Feces, Part 3, of this report. 120 Influence of Inanition on Metabolism. cially noticeable on the first day. The weights of fresh feces were as follows : At 9h 36m a. m., January 7, 241.5 grams; at 7h 24m a. m., January 8, 41.3 grams; and at 7h 24m a. m., January 9, 48 grams. As was customary in all the fasting experiments, the subject had taken a gelatin capsule filled with lampblack with the last meal immediately preceding the fast. Since none of tliis fecal matter was colored with charcoal, it was obvious that it belonged distinctly to food taken previously. It was therefore not a product result- ing from fasting metabolism. Since, however, feces were actually passed they should be taken into consideration in the balance made in table 73. Table 73. — Comparison of changes in body-weigM xoitli balance of income and outgo — Metabolism experiment No. 71. Jan. 7-8. Jan. 8-9. Jan. 9-10. Jan. 10-11. Total for 4 days. Average per day. Income: (a) Water consumed. . . Grams. 1204.40 589.12 Grams. 1973.20 554.11 Grams. 2312.90 537.97 Grams. 1485.30 492.74 Grams. 6975.80 2173.94 Grams. 1743.95 543.49 (c) Total (a + b) Outgo: (d) Urine1 , 1793.52 2527.31 2850.87 1978.04 9149.74 | 2287.44 1459.50 241.50 669.04 744.84 1796.90 41.30 570.17 665.19 2138.30 48.00 553.97 567.66 2008.30 508.13 517.66 7403.00 330.80 2301.31 2495.85 1850.75 82.70 575.33 623.84 (e) Feces2 (/) Carbon dioxide. . . . (g) Water of respira- tion andperspira- (h) Total (d+f+g) (i) Gain ( + ) or loss (— ) of body material (c—Ji). . (j) Gain ( + ) or loss (— ) of 2873.38 3032.26 3259.93 3034.09 12199.66 3049.92 1 — 1079.86 -504.95 -1091.00 —507.00 -409.06 -418.00 -1056.05 -1114.00 — 3049.92 -3130.00 — 762.48 — 782.50 1 The data In this line should not be confounded with urine data in other tables, explanation, p. 66.) -Not included in the total outgo. (See p. 119.) (See Comparison of lines i and / of table 73 shows very satisfactory agreement on the second and third days between the losses of body material and losses in weight. On the last day, however, there is a considerable error. The difficulties of securing the complete accuracy of this balance have already been dwelt upon. In the previous fasting experiments here reported no feces had been passed during the experimental period. In this experiment, however, feces were passed on the first 3 days which evidently were the resultant of food taken previous to the fast. But as it was found impracticable to separate fasting feces from those of food preceding fasting, which were in this and some sub- sequent experiments excreted during the fasting period, the best method of using the weights in the computations was not clear. The uncertainty as to the nature of the feces seemed to make it desirable to eliminate them from the Metabolism Experiment No. 71. 121 Table 74. — Summary of calorimetric measurements and total heat production — Metabolism experiment No. 11. (a) (6) (c) (d) Date. Period. Heat meas- ured in terms Heat used in vaporiza- tion of water. Sum of heat correc- tions.1 Total heat produc- tion (a+b+c). 1905. Jan. 7 Preliminary : 1 a.m. to 3 a.m 3 a.m. 5 a.m Calories. 77.6 87.6 Calories. 37.9 35.4 Calories. 2 +13.2 s — 10.2 Calories. 2 128.7 2112.8 Jan. 7-8 5 a.m. 7 a.m Total 92.8 36.8 1 + 3.0 2132.6 258 . 0 110.1 2 + 6.0 2374.1 7 a.m. to 9 a.m 173.2 235.2 49.5 49.3 — 15.1 - 7.6 207.6 3276.9 11 a.m. 1 p.m 150.2 42.6 + 11.7 204 . 5 1 p.m. 3 p.m 135.6 39.3 — 2.8 172.1 119.8 42.6 -21.4 141.0 134.4 40.7 + 21.3 196.4 7 p.m. 9 p.m 138.3 43.0 -11 .7 169.6 9 p.m. 11 p.m 98.9 31.4 — 43.0 87.3 76.3 34.0 + 47.6 157.9 73.8 31.2 + 5.1 110.1 80.4 21.0 + 30.6 132.0 Jan. 8-9 5 a.m. 7 a.m .... Total 7 a.m. to 9 a.m 9 a.m. 11 a.m 93.8 30.8 — 9.8 114.8 1509.9 455.4 + 4.9 1970.2 201.3 126.8 51.5 42.9 — 12.4 + 12.6 240.4 182.3 11 a.m. 1 p.m 119.7 34.9 + 13.6 168.2 1 p.m. 3 p.m 132.6 37.1 — 1.7 168.0 3 p.m. 5 p.m 90.5 32.6 - 7.6 115.5 132.9 35.6 + 4.5 173.0 7 p.m. 9 p.m 115.7 33.8 -11.2 138.3 9 p.m. 11 p.m 101.9 31.7 -36.1 97.5 98.3 31.7 + 41.0 171.0 88.3 31.6 - 1.5 118.4 3 a.m. 5 a.m 81.6 29.0 + 20.1 130.7 Jan. 9-10 5 a.m. 7 a.m Total 82.1 33.1 + 25.4 140.6 1371.7 425.5 + 46.7 1843.9 7am to 9 a m 180 0 32 3 —27 6 184 7 9 a.m. 11 a.m 111.6 31.8 + 10.7 154.1 11 a.m. 1 p.m 129.3 28.3 + 2.8 160.4 1 p.m. 3 p.m 123.4 30.4 - 5.1 148.7 3 p.m. 5 p.m 125.8 30.1 — 3.5 152.4 131.4 32.4 + 20.7 184 . 5 130.7 29.4 — 22.0 138.1 9 p.m. 11 p.m 125.6 30.1 — 26.1 129.6 52.1 27.7 + 51.8 131.6 95.9 28.0 -11.1 112.8 82.6 27.3 + 18.3 128.2 5 a.m. 7 a.m Total 76.6 27.0 + 17.6 121.2 1365.0 354.8 + 26.5 1746.3 1 See pp. 42-49. 2 Not corrected for change in body temperature and weight. 3 Not corrected for 2.74 calories introduced electrically in magnetizing the fields of the bicycle ergometer. 122 Influence of Inanition on Metabolism. Table 74. — Summary of calorimetric measurements and total heat production — Continued. (o) (B) (c) (d) Date. Period. Heat meas- ured in terms C20 Heat used in vaporiza- tion of water. Sum of heat correc- tions.1 Total heat produc- tion (a+ft+c). Jan. 10- 1905. -11 Calories. 161.0 119.3 Calories. 32.3 27.0 Calories. — 23.0 + 8.4 Calories. 170.3 154.7 11 a.m. 1 p.m 118.4 27.0 + 3.2 148.6 119.1 27.7 - 4.7 142.1 112.6 27.5 — 0.0 140.1 112.0 30.5 + 4.0 146.5 125.8 27.2 — 19.5 133.5 9 p.m. 11 p.m 96.5 25.3 — 35.1 86.7 62.5 26.4 + 43.7 132.6 88.2 27.5 — 0.9 114.8 3 a.m. 5 a.m 79.7 25.9 + 11.9 117.5 Total 76.4 24.0 + 17.8 118.2 1271.5 328.3 + 5.8 1605.6 *See pp. 42-49. energy and heat computations. As shown above, the feces passed during the experiment were not an excretory product as a result of fasting metabolism and had they been retained in the colon until the end of the experiment would have affected in no wise the heat or energy balance.28 Indeed, in some of the fasting experiments the feces were artificially removed by means of an enema before the subject entered the chamber, and none were passed during the time of the experiment. Accordingly in the fasting experiments it was assumed that the feces were retained in the body, and hence the weight of material as well as the heat lost by cooling from body temperature to calorimeter temperature were computed and corrected for, in column c of table 74. Allowance has been made in line ; of table 73 for feces passed, the amounts being deducted from the total loss of weight in order to make the data comparable with the balance of income and outgo. The feces passed during the fasting experiments were not analyzed. Output of Heat. The calorimetric measurements are summarized as usual and correction made for changes in body temperature and body-weight. The total heat production is recorded in column d of table 74. Although the data for the preliminary night are also included, it is to be borne in mind that the value of the results for these three periods is somewhat uncertain because of the necessary incompleteness of preliminary heat corrections. 28 The reabsorption of small amounts of material in the large colon is probable, but hardly a quantitative factor. Metabolism Experiment No. "71. 123 Table 74 shows that the total heat production was greatest on the first day and smallest on the last day. The extra heat eliminated as a result of the muscular work during the second period of the first day was markedly evident, for on the first day there were 276.83 calories produced as against an average of less than 165 calories for the corresponding period of the remaining days. Balance of Energy. Table 75 shows the method of deriving the computed heat production and the comparison of these results with the heat production as measured. It is true of this experiment, as of No. 69, that the total energy derived from different sources was on each day larger than the heat production as measured. The smallest discrepancy occurred on the second and the largest Table 75. — Comparison of energy derived from katabolized body material with total heat production — Metabolism experiment No. 11. Date. Energy derived from different sources. (0) Total heat produc- tion. Energy from body material greater (+) or less (— ) than output. From body protein. (d) From body fat. (e) From body glyco- gen. (f) Total (c+d+e). (a) Energy of protein katabo- lized. (b) Poten- tial energy of urine. (c) Net energy (a-b). (h) Amount (f-Q). 1001.89 F5.91 543.34 482.00 9.98 7.89 131.47 889.78 10.05 482.54 350 . 53 6!02 Total 2063.14 1577.60 9.98 9.98 139.36 139.36 174.88 174.88 1732.90 1247.36 6.02 6.02 1 Includes also water of perspiration. Metabolism Experiment No. 73. 147 Elimination of Carbon Dioxide and Absorption of Oxygen. The quantities of carbon dioxide and oxygen residual in the chamber at the end of each experimental period as well as the total amounts of carbon dioxide exhaled and oxygen consumed by the subject are given in table 97, page 144. Fluctuations are noted in the residual amounts of carbon dioxide and oxygen as in the earlier experiments. The amount of carbon dioxide excreted was greatest on the first day and diminished with considerable regularity to the end of the experiment. On the other hand, the daily amounts of oxygen consumed remained nearly constant for the first 2 days, after which there was a gradual diminution from day to day. Table 99. — Elements and materials katabolized in body — Metabolism experiment No. 73. Date. (a) Nitro- gen. (6) Carbon. (c) Hydro- gen. (d) Oxy gen (c) Water. (f) Protein. (a) Fat. (h) Carbo- hydrates (as gly- cogen). Ash. Jan. 1905. 28-29 Oms. 10.29 Oms. 173.54 Oms. 327.44 Oms. 2,491 92 Oms 2,700 44 Oms. 61.74 Oms. 106.63 Oms. 135.32 Oms. 9.96 Jan. 29-30 11.97 161.03 400.92 3,034 86 3,368 29 71.82 151.72 18.05 6.23 Jan. 30-31.. .. 11.54 155.67 371 . 53 2,801 66 3,112 16 69.24 152.62 7.44 5.74 Jan. 31-Feb.l. 10.39 148.17 281.43 2,099. 73 2,317 55 62.34 139.15 21.63 6.09 Feb. 1-2 Total, 5 days. 9.98 139.36 174.88 1,247. 36 1,375 38 59.88 148.07 110.84 6.02 54 . 17 777.77 1556.20 11,675. 53 12,873 82 325.02 698.19 171.60 34.04 1 Glycogen gained. Elements Katabolized in the Body. The elements katabolized in the body, obtained by the methods previously described, are given in table 98. The apparently excessive losses of total weight, hydrogen, and oxygen are partly compensated by the large amounts of water consumed. Elements and materials katabolized in the body. — A summary of the losses of the elements and the compounds computed therefrom by means of formulae previously recorded is given in table 99. As in the preceding table, there is a very large apparent loss of hydrogen, oxygen, and water. The loss in protein averages 65 grams per day. In common with the other fasting experiments, there is a very large loss of glycogen during the first 24 hours of the fast and lesser losses on the 3 days following, but on the fifth day there is apparently a gain of 10.84 grams. The probability of a gain of glycogen after 4 consecutive days of fasting demands special consideration, but discussion of the point is deferred until later (Part 3, Materials Katabolized, Glycogen) in the report. 1 is Influence of Inanition on Metabolism. Balance of water. — In table 100, showing the distribution of intake and output of water, the results are obtained as explained earlier in tins publica- tion. Allowance has not been made in columns d and /, however, for water of feces passed on the first 3 days of the experiment for the reason explained in the discussion of table 73, page 120. Tbe data below show that the actual loss of water to the body was very much less than the apparent loss shown in table 99. The water of oxidation of organic hydrogen is not materially different in amount from that found in the previous fasting experiments. Table 100. — Distribution of intake and outgo of water — Metabolism experiment No. 78. Date. Outgo from the body. (a) Water of urine. (6) Water of respira- tion and perspira- tion. (c) Total (a+b). Balance of preformed water. (d) Pre- formed (katabo- lized) water in outgo.1 (e) Intake in drink. (f) Loss of pre- formed water l (d-e). (P) Water of oxida- tion of organic hydro- gen (c-d). 1905. Jan. 28-29 Jan. 29-30 Jan. 30-31 Jan. 31-Feb. 1 Feb. 1-2 Grams. 2,225.8 2,928.2 2,698.3 1,928.3 1,001.9 Grams. 684.2 636.1 602.3 569.2 543.3 Grams. 2,910.0 3,564.3 3,300.6 2,497.5 1,545.2 Total, 5 days. . . , Average per day, 10,782.5 2,156.5 3035.1 607.0 13,817.6 2,763.5 Grams. Grams. Grams. 2,700.4 2,082.3 618.1 3,368.3 2,746.9 621.4 3,112.2 2,763.2 349.0 2,317.5 1,955.4 362.1 1,375.4 1,075.9 299.5 12,873.8 10,623.7 2250.1 2,574.7 2,124.7 450.0 Grams. 209.6 196.0 188.4 180.0 169.8 943.8 188.8 1 Does not include water of feces, periment only. Feces were passed on the first three days of the ex- Changes in Body-Weight Compared with Balance of Income and Outgo. The subject was weighed as usual each morning throughout the experiment and in table 101 a comparison of the changes in body-weight with the balance of income and outgo is given. The income consists of water and oxygen consumed while the outgo includes the urine, feces, carbon dioxide, and water of respiration and perspiration. Before the fasting period began the subject took a gelatin capsule filled with lampblack. Feces were passed the first 3 days of the experiment in the follow- ing amounts: First day, 71.60 grams; second day, 87 grams; and the third day, 51.10 grams, but as they contained no lampblack they undoubtedly resulted from the food consumed before the fast. In making the balance of income and outgo, however, the feces have in effect been restored to the body of the subject, and the losses shown by the scale have been correspondingly reduced. Metabolism Experiment No. 73. 149 In comparing the losses in body material with the losses in body-weight, it is seen that aside from the discrepancy of over 100 grams on the first day, the agreement is quite satisfactory. It is to be observed that on each of the last 4 days the loss of body-weight is slightly greater than the loss of body material computed from the income and outgo. Output of Heat. The summarized data for the total heat production per 2-hour period are given in table 102. The total heat production is greatest on the first day and gradually diminishes throughout the experiment. Table 101. — Comparison of changes in body weight with balance of income and outgo — Metabolism experiment No. 73. Jan. 28-29. Jan. 29-30. Jan. 30-31. Jan. 31- Feb. 1. Feb. 1-2. Total for 6 days. Average per day. Income : (a) Water consumed (&) Oxygen (c) Total (a + b) Outgo : (d) Urine1 (e) Feces2 (/) Carbon dioxide (g) Water of respiration and perspiration (h) Total (d+f + g) (i) Loss of body material (h-c) (j) Loss of body weight. . Grams. Grams. 2082.30 2746.90 544.24 547.86 2626.54 3294.76 Grams. 2763.20 533.04 3296.24 Grams. Grams. 1955.40 1075.90 502.68 2458.08 485.54 Grams. 10,623.70 2,613.36 Grams. 2124.74 522.67 1561.44 13,237.06 2647.41 2615.50 2770.80 71.60 608.87 684.22 3908 . 59 87.00 560.02 636.05 J.966.87 2966.40 2143.90 51.10 541.67 602.31 4110.38 515.22 569.17 3228.29 1192.10 482.00 543.34 2217.44 11,688.70 209 . 70 2,707.78 3,035.09 17,431.57 2337.74 41.94 541.56 607.02 3486.31 1282.05 1168.00 672.11 686.00 814.14 836.00 770.21 779.00 656.00 668.00 4,194.51 4,137.00 838.90 827.40 1 The data in this column should not he confounded with urine data In other tables, explanation, p. 66. 2 Not included in the total outgo. See p. 148. See Balance of Energy. Allowing for the energy of the incompletely oxidized products of protein katabolism, the total energy derived from all sources may be computed and com- pared with the total heat production. This comparison is shown in table 103. On comparing the total energy derived from different sources with the total heat production it is seen that on the first day the computed energy is 19 calories, or 1.0 per cent, smaller than the total heat production. The largest discrepancy appears on the last day when the total as derived from all sources is 66 calories, i. e., 4.3 per cent greater than the total heat production. In these computations it should be borne in mind that the energy from the body glycogen stored on the last day has been deducted from and not added to the energy from the body protein and fat inasmuch as it represents energy stored and not liberated. The largest discrepancy appears on the day on which the smallest total heat production is measured. 150 Influence of Inanition on Metabolism. Table 102. — Summary of calorimetric measurements and total heat production- Metabolism experiment No. 73. (a) (o) (c) (d) Date. Period. Heat meas- ured in terms 0>o* Heat used in vaporiza- tion of water. Sum of heat correc- tions.1 Total heat produc- tion {a+b+c). 1905. Jan. 28 Preliminary : 1 a.m. to 3 a.m. . . . 3 a.m. 5 a.m. . . . Calories. 96.1 88.7 Calories. 42.6 44.9 Calories. 2- 3.6 2 Calories. 2 135.1 2 133.6 Jan. 28-29 5 a.m. 7 a.m. . . . Total 88.6 39.8 2 2 128.4 273.4 127.3 2 — 3.6 2 397.1 7 a.m. to 9 a.m. . . . 178.4 47.7 — 26.5 199.6 9 a.m. 11 a. m. . . . 138.3 44.6 + 1.2 184.1 11 a.m. 1 p.m. . . . 128.1 39.5 + 11.7 179.:; 1 p.m. 3 p.m. . . . 118.2 38.0 - 1.7 154.5 3 p.m. 5p.m ... 130.4 36.9 -10.6 156.7 5p.m. 7 p.m. . . . 128.4 39.2 + 15.8 183.4 7 p.m. 9 p.m. . . . 126.7 35.9 - 5.9 156.7 9 p.m. 11 p.m. . . . 129.6 35.0 — 22.6 142.0 11 p.m. 1 a.m. . . . 59.6 36.7 + 14.8 111.1 1 a.m. 3 a.m. . . . 70.0 36.3 + 9.3 115.6 3 a.m. 5 a.m. . . . 94.9 34.9 + 14.4 144.2 Jan. 29-30 5 a.m. 7 a. m. . . . Total 87.5 35.1 + 15.8 138.4 1390.1 459.8 + 15.7 1865.6 7 a.m. to 9 a. m. . . . 9 a.m. 11 a.m. . . . 165.0 132.8 37.5 42.4 — 19.6 + 15.5 182.9 190.7 11 a.m. 1 p.m. . . . 128.3 35.0 — 0.7 162.6 1 p.m. 3 p.m. . . . 119.5 33.5 + 5.1 158.1 3 p.m. 5 p.m. . . . 112.6 33.7 + 3.4 149.7 5 p.m. 7 p.m. . . . 137.6 35.2 + 5.8 178.6 7 p.m. 9 p.m. . . . 129.3 34.3 -12.4 151.2 9 p.m. 11p.m.... 123.6 33.0 —15.2 141.4 11 p.m. 1 a. m. . . . 51.5 31.1 + 18.5 101.1 1 a.m. 3 a.m. . . . 77.5 32.3 + 4.4 114.1 3 a.m. 5 a.m. . . . 81.1 30.8 + 24.9 136.8 5 a.m. 7 a.m. . . . Total 7 a.m. to 9 am 89.0 28.6 + 5.9 123.5 1347.8 407.4 + 35.6 | 1790.8 Jan. 30-31 165 8 36 8 — 25.2 + 6.5 177.4 174.3 9 a.m. 11 a.m. . . . 133.5 34.3 11 a.m. 1 p. m. . . . 111.3 32.8 + 14.3 158.4 1 p.m. 3 p.m. . . . 121.3 32.7 + 7.3 161.3 3 p.m. 5 p.m. . . . 134.5 32.9 - 0.9 166.5 5 p.m. 7 p.m. . . . 128.9 32.3 + 8.3 169.5 7 p.m. 9 p.m. . . . 129.4 33.0 -17.5 144.9 9 p.m. 11 p. m. . . . 113.6 29.4 + 0.5 143.5 11 p.m. 1 a.m. . . . 63.2 29.4 - 0.0 92.6 1 a.m. 3 a.m. . . . 76.6 27.1 + 3.1 106.8 3 a.m. 5 a. m. . . . 68.5 29. 9 + 22.9 121.3 5 a.m. 7 a.m. . . . Total 80.5 27.9 + 14.2 122.6 1327.1 378.5 + 33.5 1739.1 1 Spe pp. 42-49. 2 Not corrected for change in body temperature and weight. Metabolism Experiment No. 73. 151 Table 102. — Summary of calorimetric measurements and total heat production- Continued. (a) (6) (c) (d) Date. Period. Heat meas- ured in terms Heat used in vaporiza- tion of water. Sum of heat correc- tions.1 Total heat produc- tion (a+b+c). 1905. Jan. 31-Feb. 1 7 a.m. to 9 a.m. . . . Calories. 149.1 Calories. 32.6 Calories. -23.3 Calories. 158.4 9 a.m. 11 a.m. . . . 123.1 30.9 + 7.5 161.5 11 a.m. 1 p.m. . . . 128.1 32.4 + 7.6 168.1 1 p.m. 3 p.m. . . . 122.1 30.0 + 4.5 156.6 3 p.m. 5 p.m. . . . 115.6 28.6 - 3.4 140.8 5 p.m. 7 p.m. . . . 126.9 29.1 - 0.8 155.2 7 p.m. 9 p.m. . . . 117.3 31.3 — 5.5 143.1 9 p.m. 11 p.m. . . . 112.3 28.9 — 22.2 119.0 11 p.m. 1 a.m. . . . 59.1 29.1 + 18.0 106.2 1 a.m. 3 a.m. . . . 59.0 27.7 + 7.9 94.6 3 a.m. 5 a.m. . . . 80.8 27.5 + 21.5 129.8 Feb. 1-2 5 a. m. 7 a. m . . . . Total 85.6 27.1 + 16.9 129.6 1279.0 355.2 + 28.7 1662.9 7 a.m. to 9 a.m. . . . 9 a. m. 1 1 a. m ... . 155.5 118.6 31.2 28.9 — 32.5 + 6.8 154.2 154.3 11 a.m. 1 p.m. . . . 102.1 27.1 + 1.9 131.1 1 p.m. 3 p.m. . . . 114.1 27.8 — 0.8 141.1 3 p.m. 5p.m. .. . 93.4 24.8 — 1.7 116.5 5 p.m. 7 p.m. . . . 114.8 28.4 - 2.0 141.2 7p.m. 9 p.m. . . . 120.9 29.5 — 6.4 144.0 9 p.m. 11 p.m. . . . 117.7 30.6 — 18.3 130.0 11 p.m. 1 a.m. . . . 54.6 25.7 + 19.2 99.5 1 a.m. 3 a. m. . . . 74.7 26.5 — 6.5 94.7 3 a.m. 5 a.m. . . . 75.0 25.5 + 26.7 127.2 5 a.m. 7 a.m. . . . Total 71.4 27.5 + 14.9 113.8 1212.8 333.5 + 1.3 1547.6 Table 103. — Comparison of energy derived from katabolized body material with total heat production — Metabolism experiment No. 73. Date. Energy derived from different sources. (P) Total heat produc- tion. Energy from body material greater (+) or less ( — ) than output. From body protein. (d) From body fat. (e) From body glyco- gen. Total (c+d-t-e). (a) Energy of protein katabo- lized. (6) Poten- tial energy of urine. (c) Net energy (a-b). (.h) Amount (i) Pro- portion (7i + g). 1905. Jan. 28-29 Jan. 29-30 Jan. 30-31 Jan. 31-Feb. 1. Feb. 1-2 Total, 5 days. Av. per day.. Cals. 349 406 391 352 33S CaU. 86 98 96 92 92 Cals. 263 308 295 260 246 Cals. 1017 1447 1456 1327 1413 Cals. 567 76 31 91 '45 Cals. 1847 1831 1782 1678 1614 Cals. 1866 17'.)! 1739 1663 1548 Cals. — 19 + 40 + 43 + 15 + 66 Per ct. -1.0 + 2.2 + 2.5 + 0.9 + 4.3 1836 367 464 93 1372 274 6660 1332 720 144 8752 1750 8607 1721 + 145 I + 29 +1.7 1 Glycogen gained. 152 I \ [[.[ i:nci: of Inanition on Metabolism. Table 104. — Oxygen and carbon dioxide thermal quotients and respiratory quotients — Metabolism experiment No. 73. Date and period. (a) Total heat produc- tion. (6) Oxygen con- sumed. (c) Oxygen thermal quotient (100b -=- a). (d) Carbon dioxide elimi- nated. (e) Carbon dioxide thermal quotient (100d-=-a). (/) (a) Volume Volume of carbon of dioxide \ oxygen elimi- i con- nated | sumed (dXO.5091) (bxO.T). (fc) Re- spira- tory quo- tient l/H-y). Preliminary : 1905. Jan. 28: 1 a.m. to 3 a.m. . Cals. '135.1 Grams. 35.8 26.5 Orams. 45.0 33.3 Liters. 22.9 Liters. 25.1 0.91 3 a.m. 5 a.m. . '133.6 37.0 27.6 56.2 42.1 28.6 25.9 1.11 5 a.m. 7 a.m. . Total U28.4 38.3 29.9 52.5 40.9 26.8 26.8 1.00 '397.1 111.1 28.0 153.7 38.7 78.3 77.8 1.01 Jan. 28-29: 7 a.m. to 9 a.m. . 199.6 61.6 30.8 68.2 34.2 34.7 43.1 .81 9 a.m. 11 a.m. . 184.1 50.4 27.4 62.7 34.1 31.9 35.3 .90 11 a.m. 1 p.m. . 179.3 48.7 27.2 57.7 32.2 29.4 34.1 .86 1 p.m. 3 p.m. . 154.5 43.7 28.3 52.6 34.1 26.8 30.6 .88 3 p.m. 5 p.m. . 156.7 46.8 29.9 52.2 33.3 26.6 32.8 .81 5 p.m. 7 p.m. . 183.4 50.8 27.7 55.7 30.3 28.3 35.6 .80 7 p.m. 9 p.m. . 156.7 42.6 27.2 45.1 28.8 23.0 29.8 .77 9 p.m. 11 p.m. . 142.0 48.0 33.8 50.2 35.3 25.5 33.6 .76 11 p.m. 1 a.m. . 111.1 36.1 32.5 42.7 38.4 21.7 25.3 .86 1 a.m. 3 a.m. . 115.7 40.1 34.7 37.5 32.5 19.1 28.1 .68 3 a.m. 5 a.m. . 144.2 36.2 25.1 41.8 29.0 21.3 25.3 .84 5 a.m. 7 a.m. . Total 138.4 39.2 28.3 42.5 30.7 21.7 27.4 .79 1865.7 544.2 29.2 608.9 32.6 310.0 381.0 .81 Jan. 29-30: 7 a. m. to 9 a. m . . 182.9 63.7 34.8 62.6 34.2 31.8 44.6 .71 9 a.m. 11 a.m. . 190.7 55.5 29.1 56.8 29.8 28.9 38.8 .75 11 a.m. 1 p.m. . 162.6 47.6 29.3 48.8 30.0 24.8 33.3 .75 1 p.m. 3 p.m. . 158.1 47.2 29.9 49.6 31.4 25.3 33.1 .76 3 p.m. 5 p.m. . 149.2 51.8 34.6 47.4 31.7 24.1 36.3 .67 5 p.m. 7 p.m. . 178.6 49.1 27.5 53.3 29.9 27.2 34.4 .79 7 p.m. 9 p.m. . 151.2 45.6 30.2 46.1 30.5 23.5 31.9 .74 9 p.m. 11 p.m. . 141.4 48.7 34.5 47.2 33.4 24.0 34.1 .71 11 p.m. 1 a.m. . 101. 1 28.4 28.1 36.8 36.4 18.7 19.9 .94 1 a.m. 3 a.m. . 114.1 40.7 35.7 37.3 32.7 19.0 28.5 .67 3 a.m. 5 a. m. . 136.8 37.4 27.3 37.3 27.3 19.0 26.1 .73 5 a.m. 7 a.m. . Total 123.6 32.2 26.1 36.8 29.8 18.8 22.5 .83 1790.8 547.9 30 . 6 560.0 31.3 285 . 1 383.5 .74 Jan. 30-31: 7 a.m. to 9 a.m. . 177.4 68.2 38.4 64.2 36.2 32.7 47.7 .68 9 a.m. 11 a.m. . 174.3 49.5 28.4 52.2 30.0 26.6 34.7 .77 11 a.m. 1 p.m. . 158.4 49.0 31.0 50.5 31.8 25.7 34.3 .75 1 p.m. 3 p.m. . 161.3 50.1 31.0 50.7 31.5 25.8 35.0 .74 3 p.m. 5 p.m. . 166.5 48.7 29.3 50.6 30.4 25.7 34.1 .76 5 p.m. 7 p.m. . 169.5 46.2 27.3 49.9 29.4 25.4 32.3 .79 7 p.m. 9 p.m. . 144.9 43.4 29.9 44.9 31.0 22.9 30.4 .75 9 p.m. 11 p.m. . 143.5 37.3 26.0 40.9 28.5 20.8 26.1 .80 11 p.m. 1 a.m. . 92.6 33.7 36.4 36.5 39.4 18.6 23.6 .79 1 a.m. 3 a.m. . 106.8 33.0 30.9 33.2 31.1 16.9 23.1 .73 3 a.m. 5 a.m. . 121.3 40.2 33.2 40.2 33.1 20.5 28.2 .73 5 a.m. 7 a.m. . Total 122.6 33.7 27.5 27.9 22.8 14.2 23.6 .60 1739.1 533.0 30.7 541.7 31.2 275.8 373.1 .74 See p. 150. Metabolism Experiment No. 73. 153 Table 104. — Oxygen and carbon dioxide thermal quotients and respiratory quotients — Continued. Date and period. (a) Total heat produc- tion. (b) Oxygen con- sumed. (c) Oxygen thermal quotient (lOOb+a). (d) Carbon dioxide elimi- nated. («) Carbon dioxide thermal quotient (lOOdH-a). (/) Volume of carbon dioxide elimi- nated (dxO.5091) (0) Volume of oxygen con- sumed (/*X0.7). (h) Re- spira- tory quo- tient (/-0>. 1905 Jan. 31-Fe 7 a.m. to i>. 1 : 9 a.m. . Cals. 158.4 Orams. 55.1 34.8 Orams. 56.0 35.4 Liters. 28.5 Liters. 38.6 .74 9 a.m. 11 a.m. . 161.5 53.6 33.2 52.7 32.6 26.8 37.5 .72 11 a.m. 1 p.m. . 168.1 49.5 29.4 50.2 29.9 25.6 34.6 .74 1 p.m. 3 p.m. . 156.6 41.6 26.5 45.4 29.0 23.1 29.1 .80 3 p.m. 5 p.m. . 140.8 43.3 30.7 43.1 30.6 21.9 30.3 .72 5 p.m. 7 p.m. . 155.2 43.3 27.9 47.0 30.3 24.0 30.3 .79 7 p.m. 9 p.m. . 143.1 42.9 30.0 46.2 32.3 23.5 30.1 .78 9 p.m. 11 p.m. . 119.0 38.1 32.1 38.9 32.7 19.8 26.7 .74 11 p.m. 1 a. m . . 106.2 30.3 28.5 34.6 32.6 17.6 21.2 .83 1 a.m. 3 a. m . . 94.6 36.3 38.4 33.1 35.0 16.9 25.4 .66 3 a.m. 5 a.m. . 129.8 32.7 25.2 32.7 25.2 16.6 22.9 .73 5 a.m. Total. . Feb. 1-2: 7 a.m. . 129.6 36.0 27.8 35.3 27.2 18.0 25.2 .71 1662.9 502.7 30.2 515.2 31.0 262.3 351.9 .75 7 a.m. to 9 a. m . . 154.2 52.3 33.9 53.5 34.7 27.2 36.6 .74 9 a.m. 11 a.m. . 154.3 44.6 28.9 43.0 27.9 21.9 31.2 .70 11 a.m. 1 p.m. . 131.1 42.0 32.0 42.1 32.1 21.5 29.4 .73 1 p.m. 3 p.m. . 141.1 41.3 29.3 42.9 30.4 21.8 28.9 .75 3 p.m. 5 p.m. . 116.5 38.5 33.0 40.7 34.9 20.7 26.9 .77 5 p.m. 7 p.m. . 141.2 46.8 33.2 43.6 30.9 22.2 32.8 .68 7 p.m. 9 p.m. . 144.0 44.2 30.7 45.2 31.4 23.0 31.0 .74 9 p.m. 11 p.m. . 130.0 41.6 32.0 39.8 30.6 20.3 29.1 .70 11 p.m. 1 a. m . . 99.5 28.1 28.2 31.8 32.0 16.2 19.7 .82 1 a.m. 3 a.m. . 94.7 33.7 35.6 80.0 31.7 15.3 23.6 .65 3 a.m. 5 a.m. . 127.2 36.1 28.4 34.6 27.2 17.6 25.3 .70 5 a.m. Total.. 7 a.m. . 113.8 36.3 31.9 34.8 30.5 17.7 25.4 .70 1547.6 485.5 31.4 482.0 31.2 245.4 339.9 .72 Relations between Oxygen Consumption, Carbon Dioxide Elimination, and Heat Production. The relationships expressed by the oxygen thermal, carbon dioxide thermal, and respiratory quotients are given in table 104. It appears that on the first day the oxygen thermal quotient is the lowest, the carbon dioxide and respir- atory quotient the highest, while on the succeeding days all three ratios are approximately constant. This statement does not take into account the fluctu- ations in these quotients during the 2-hour periods. In this experiment, however, the fluctuations seem less marked than in previous studies here reported. 154 Influence of Inanition on Metabolism. Metabolism Experiment No. 74. Immediately following the 5-day fast (experiment No. 73), an experiment was made with the same subject in order to obtain additional information regarding the effect on metabolism of the ingestion of food. The experiment began February 2, 1905, at 7 a. m., and continued without interruption for 72 hours. Thus, the series (experiments Nos. 73 and 74) represents a contin- uous 8-day experiment inside the respiration chamber, of which 5 days were without and 3 days with food. As previously pointed out the data obtained in the experiment with food, especially on the first and second days, indicate the transformations during transition from fasting to food. Notes from diary, pulse records, and records of body movements. — Such notes from the diary as are deemed essential to an understanding of the condition of the subject during his stay in the calorimeter, the records of his pulse rate, and the usual records of body movements are given below : Notes from diary. Feb. 2, 1905: 7h30,n a. m. Passed a fairly good night; awoke first time at 3h25m a. m., fell asleep again at 3M5m a. m. and awoke at 6h25m a. m. My tongue has a white coat and is swollen, showing my teeth marks on it; have a sweet taste in my mouth. Have just taken a little orange juice. 4 p. m. Feel very comfortable. The swelling of the tongue is gradually diminishing. I feel so good I could undertake another 5 days' fast to- morrow. 10 p. m. Have passed a very comfortable day, but it is becoming a little mo- notonous in here and I will be ex- tremely glad to come out. 10h30m p. m. Pain in heart. Feb. 3, 1905: 7h30m a. m. Have passed a fairly good night; was still sleepy when called. 8h15m a. m. Have just defecated, but with difficulty. 12 m. Finished my second allotment of milk. It is very rich and has a dif- ferent taste from that I usually drink. Tongue still swollen and coated; taste in the mouth only moderately bad. 4 p. m. Feel very comfortable, but a little sleepy. Feb. 4, 1905: 7h30m a. m. Did not sleep very well and was still very sleepy when I re- ceived the rising signal. My tongue is heavily coated with a brownish fur; taste in mouth is bad. I think the milk is making me bilious, as it is very heavy and rich. 8 a. m. My tongue is still swollen a little. 8h15m a. m. Defecated with much diffi- culty. 9 p. m. Have a slight dull (bilious) headache. 10 p. m. My headache is increasing and I have a nauseated feeling; did not take sufficient acids for the amount of milk I drank. Pulse-rate — Experiment No. 711 Time 7.30 8 10 12 2 i 6 8 10 a.m. a.m. a.m. m. 64 p.m. p.m. p.m. p.m. p.m. Feb. 2 62 57 64 2 75 2 75 51 264 59 Feb. 3 55 56 55 54 53 57 62 54 54 Feb. 4 65 66 66 63 2 7o 2 66 264 55 68 l. 1'ulse taken while sitting. 2 Strong. Metabolism Experiment No. 74. 155 Movements of subject, duration 3 days, from Feb. 2, 7 a. m., to Feb. 5, 7 a. m., 1905. A. M. 7h00r 7 7 7 7 7 06 I 10/ 15 18 20 7 30 7 38 8 00 9 02 9 04 9 07 9 08 9 12 9 36 10 00 10 15 11 02 11 04 11 06 12 00 P. M. 12h 04m 12 06 12 32 1 02 1 04 1 08 1 10 1 26 2 00 2 04 2 3 3 3 3 4 4 4 4 4 5 5 5 5 10 02 04 44 50 00 04 08 12 22 02 08 09 12 5 14 5 20 5 25 5 28 February 2. rise, urinate. weigh self, etc. food aperture. sit. Comb hair, food aperture, drink, count pulse, write, sit. count pulse, telephone, rise, food aperture, sit. rise, food aperture, sit. write, telephone, count pulse, drink. move about, stand, read, food aperture, count pulse. move about. sit, write. telephone. rise, move about. food aperture, lean on table, urinate. move about. food aperture. sit, read. count pulse. move about, food aperture. sit, write. move about. sit, write. move about. food aperture. count pulse, write. food aperture. food aperture. lean on table. sit. rise, lean on table. telephone. food aperture. vigorous move- ment. stand, doctor count pulse. stand. take temperature. read. p. M. 6h00n 6 04 6 08 7 7 7 7 7 7 00 04 16 24 42 48 8 00 8 02 8 12 9 00 9 24 9 38 10 00 10 02 10 04 10 30 11 00 11 02 A. M. 7h 00„ 7 7 7 7 7 04 n} 17 18 30 8 00 8 02 8 10 8 15 8 18 8 25 8 26 9 02 9 08 10 00 10 20 10 22 10 24 11 02 11 06 11 08 11 10 11 16 11 18 11 26 11 28 11 34 11 40 11 42 12 00 count pulse. food aperture. sit, read. rise, urinate. move about. food aperture. sit. lean on table, read. sit. count pulse. rise, food aperture. sit. rise. move. sit. count pulse. open bed. lie. write. rise. close curtain, un- dress, urinate, re- tire. February 8. rise, urinate, weigh self, etc. food aperture. sit. count pulse. count pulse. move about. lean on table. defecate. sit, read. telephone, food ap- erture. sit. rise, food aperture. rise, food aperture. sit, count pulse. rise, move about. sit. read. move about, food aperture. lean on table. food aperture. lean on table. sit. read. rise, food aperture. stand. sit, read. stand. sit, read. count pulse, write. P. M. 1 104" 1 move about, uri- nate. 1 08 food aperture, lean on table, read. 1 16 move about, lean on table, read. 1 28 sit, read. 2 00 count pulse. 2 02 food aperture. 2 06 stand, eat. 2 16 stop eating. 2 18 sit, read. 2 20 food aperture. 3 46 asleep, head on table. 3 54 awake. 3 56 count pulse, write. 4 04 rise, food aperture. 4 08 move about. 4 12 sit, head on table, asleep. 4 14 rise, food aperture. 4 16 stand, read. 4 24 sit, asleep. 5 02 awake. 5 06 read. 5 38 telephone. 5 40 stand, doctor read diary. 5 42 move about. 5 46 doctor count pulse. 5 54 food aperture. 5 56 take temperature. 6 00 count pulse. 6 04 rise, move about, sit. 6 05 food aperture. 6 10 read. 7 02 rise, urinate. 7 32 rise. 7 36 lean on table. 8 00 food aperture, count pulse. 8 06 stand, lean on table. 8 20 sit, read. 8 48 sit, head on table. 9 12 sit. 9 14 stand. 9 16 open bed. 9 20 lie. 10 00 count pulse. 10 16 lie. 11 00 rise, urinate. 11 02 close curtain, un- dress, retire. 156 Influence of Inanition on Metabolism. Movements of subject. — Continued. February 4- A. M. p. M. A. M. 11" 06m move about, food 5b 02m rise, move about. 7h 00"' rise, urinate. aperture. 5 08 lean on table, read. 7 04 t weigh self, etc. food aperture, sit. 11 36 sit. 5 38 walk about. 7 09 11 50 food aperture. 5 40 lean on table, read. 7 16 12 00 count pulse. 5 44 telephone. 7 18 food aperture. p. M. 5 52 move about. 7 22 move about. 12h 02m sit, read. 6 00 count pulse, take 7 30 count pulse, write. 1 02 urinate, move temperature. 7 36 sit. about, lean on 6 02 telephone. 7 46 read. table, read. 6 04 food aperture. 8 00 count pulse. 1 06 food aperture. 6 12 rise, lean on table, 8 08 move about. 1 08 lean on table, read. read. 8 10 food aperture. 1 32 sit. 6 14 telephone. 8 15 defecate. 2 00 count pulse. 6 30 sit, read. 8 20 sit read. 2 08 rise, food aperture. 7 00 rise, urinate. 9 02 move about. 2 10 sit. 7 06 lean on table, read. 9 9 04 10 food aperture, sit, read. 2 3 3 3 3 40 12 20 22 30 move about, rise, move about, sit. eat. 7 7 20 24 walk. sit. 9 32 drink. 8 00 food aperture, 9 34 sit. move about. count pulse. 10 00 count pulse. 3 36 sit. 10 00 count pulse. 10 18 move about, lean 3 44 move about. 10 08 open bed. on table. 3 46 pick up papers. 10 12 lie. 10 26 sit. 3 48 walk about. 11 00 rise, undress, uri- 10 32 read. 3 52 sit, read. nate. 11 00 move about. 4 00 count pulse. 11 02 close curtain. Wateb and Oxygen Consumed and Urine, Carbon Dioxide, and Water-Vapor Eliminated. The data showing the amounts of water consumed, the determinations in the urine per period and per day, the oxygen intake and output of carbon dioxide, and the elimination of water-vapor are shown in tables 105 to 109 inclusive. The methods of obtaining the data have previously been described. Table 105. — Record of water consumed ' — Metabolism experiment No. 74- Date. 7 to 9 a. m. 9 to 11 a. m. 11 a. in. to 1 p. m. 1 to 3 p.m. 3 to 5 p. m. Total for day. Feb. 2-3 Grams. 272.1 370.3 335.7 Grams. 148.8 155.1 98.0 Grams. 91.1 73.3 91.1 Grams. 45.6 99.5 205.4 Grains. 45.7 324.3 Grams. 603.3 1022.5 730.2 Feb. 3-4 Feb. 4-5 * Period during which water was consumed, was assumed in some instances. (See page 73.) Metabolism Experiment No. 74. 157 Table 106. — Determinations in urine per period and per day- experiment No. IJf. -Metabolism Date and period. (a) Amount. (6) Specific gravity. (c) Volume (a-s-6). (d) Reaction. (e) • Nitro- gen. '/) Phos- phoric acid by titra- tion (P-A). (a) Total cre- atinine. 1905. Feb. 2-3 : 7 a.m. to 1 p.m. . . . 1 p.m. 7 p.m. . . . 7 p.m. 11 p.m. . . . 11 p.m. 7 a.m. . . . Total Grams. 477.4 619.7 105.1 316.7 1 . 0070 1.00S9 1.0150 1.00S9 c. c. 474 617 104 314 Acid Neutral. . Neutral. . Neutral. . Neutral. . Acid ... Grams. 3.30 3.38 1.49 2.57 Grams. 0.461 .392 .196 .341 Grams. 0.265 .223 .175 .357 1518.9 1518.9 1.0066 1509 1509 10.74 10.76 1.390 1.381 1.020 1.060 Total by composite. Feb. 3-4 : 7 a.m. to 1 p.m. . . . 1 p.m. 7 p.m. . . . 7 p.m. 11 p.m. . . . 11 p.m. 7 a.m. . . . Total Total by composite. Feb. 4-5 : 7 a.m. to 1 p.m. . . . 1 p.m. 7 p.m. . . . 7 p.m. 11 p.m. . . . 11 p.m. 7 a.m. . . . Total 724.1 460.5 215.1 494.7 1.0036 1.0044 1.0061 1.0056 722 458 214 492 2.71 2.39 1.15 2.00 .315 .384 .299 .241 .247 .148 .343 1894.4 1894.4 1.0043 1886 1866 8.25 8.25 1.148 .979 1.010 494.6 484.6 129.1 496.7 1.0034 1.0038 1.0107 1.0056 493 483 127 494 1.79 1.96 0.96 2.07 .238 .308 .403 .220 .226 .170 .224 1605.0 1605.0 1.0048 1597 1597 6.78 6.77 1.141 .840 .860 Total by composite. Total, 3 days 5018.3 .... 4992 25.77 2.839 Table 107. — Weight, composition, and heat of combustion of urine- experiment No. IJf. -Metabolism Feb. 2-3. Feb. 3-4. Feb. 4-5. Total for 3 days. (a) Weight grams (6) Water do. . (c) SolidB, a— b do.. (d) Ash do . . (e) Organic matter, c—d do.. (/) Nitrogen do. . (g) Carbon do . . (h) Hydrogen in organic matter. . . .do. . (i) Oxygen (by difference) in organic mat ter, e— (f + g + h) grams (j) Phosphorus do.. Phosphoric acid (P205): (k) By fusion grams (I) By titration do. . (m) Sulphur do. . (ra) Sulphur trioxide (SOs) do. . (o) Total creatinine do.. (p) Uric acid do. . (g) Chlorine do. . (r) Sodium chloride do. . (s) Heat of combustion calories 1518.9 1485.94 32 . 96 3.49 29.47 10.74 7.29 1.97 9.47 .578 1.323 1.381 .614 1.533 1.06 .329 .72 1.19 88 1894.4 1866.17 28.23 4.17 24.06 8.25 6.06 1.52 8.23 .430 .986 1.148 .511 1.278 1.01 .537 1.31 2.17 68 1605.0 1579.48 25.52 4.82 20.70 6.78 5.46 1.28 7.18 .498 1.141 1.141 .439 1.097 .86 .407 1.34 2.21 61 5018.3 4931.59 86.71 12.48 74.23 25.77 18.81 4.77 24.88 1.506 3.450 3.670 1.564 3.908 2.93 1.273 3.37 5.57 217 158 Influence of Inanition on Metabolism. Table 108. — Record of water of respiration and perspiration — Metabolism experiment No. 7%. Date and period. (a) Total amount of vapor in chamber at end of period. (6) Total water of respira- tion and perspira- tion.1 Date and period. (a) Total amount of vapor in chamber at end of period. (6) Total water of respira- tion and perspira- tion.1 1905. Preliminary : Feb. 2: 5 a.m. Feb. 2-3: 7 a.m. 9 a.m. 11 a.m. 1 p.m. 3 p.m. 5 p.m. 7 p.m. 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. Total Feb. 3-4: 7 a.m. 9 a.m. 11 a.m. 1 p.m. 3 p.m. 5 p.m. to 7 a.m. to 9 B m. . . . 11 a. m. . . . 1 P m. . . . 3 P m. . . . 5 P m. . . . 7 P m. . . . 9 P- m. . . . 11 P m. . . . 1 a m. . . . 3 a m. . . . 5 a m. . . . 7 a m. . . . to 9 a.m. 11 a.m. 1 p.m. 3 p.m. 5 p.m. 7p.m. Grams. 23.4 28.3 26.5 29.0 29.8 29.3 25.8 27.2 25.2 23.8 23.3 24.1 22.3 28.0 25.7 25.5 26.6 26.7 27.1 Grams. 49.9 52.2 50.6 59.0 51.2 52.7 49.6 51.2 39.7 45.8 39.8 43.2 584.9 40.5 48.4 42.3 49.8 45.4 49.9 Feb. 3-4 7 p.m. 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. Total Feb. 4-5 : 7 a.m. 9 a.m. 11 a.m. 1 p.m. 3 p.m. 5 p.m. 7 p.m. 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. Total 1905. to 9 1' m. . . . 11 P m. . . . 1 a m. . . . 3 a m. . . . 5 a m. . . . 7 a m. . . . Grams. Grams. 25.3 41.7 23.3 42.8 24.2 41.7 22.7 41.7 23.6 40.2 23.1 43.2 to 9 a. m. . . . 11 a m. . . . 1 P m. . . . 3 P m. . . . 5 l1 m. . . . 7 P m. . . . 9 P m. . . . 11 P m 1 a m. . . . 3 a m. . . . 5 a m. . . . 7 a m. . . . 28.4 25.8 26.3 25.7 27.4 27.3 26.8 24.5 26.3 24.2 25.4 22.6 527.6 48.7 45.3 42.2 44.9 44.7 49.4 44.1 43.2 40.8 41.6 40.6 39.4 524.9 1 Allowance has been made for water lost by absorbers, chair, bedding, and miscellaneous articles as follows : Feb. 2-3, 14.85 grams ; Feb. 3-4, 12.49 grams ; Feb. 4-5, 23.38 grams. Elements and Materials Katabolized in Body. The usual methods were followed in obtaining the elements and materials katabolized in the body. The data are given in tables 110 and 111. As in the other food experiments reported in this publication the food and water consumption are treated separately from the katabolism. The importance of this conception of the relation of food to katabolism warrants its repetition. Metabolism Experiment No. 74. 159 Table 109.- —Record of carbon dioxide and oxygen — Metabolism experiment No. 7^. Date. Period. Carbon dioxide. Oxygen. (a) Amount in chamber at end of period. (b) Total weight exhaled by subject. (c) . Amount in chamber at end of period. (d) Total amount consumed by subject. 1905. Feb. 2.... Feb.2-3 . . Preliminary: 5 a. m. to 7 a. m. . 7 a. m. to 9 a. m. . Grams. 20.9 Grams. Liters. 924.4 Grams. 28.4 52.6 917.4 53.9 9 a. m. 11 a. m. . 30.2 49.8 919.8 51.0 11 a. m. 1 p. m. . 31.3 50.2 906.6 51.8 1 p.m. 3 p. m. . 35.2 55.0 901.2 55.8 3 p. m. 5 p. m. . 34.7 51.6 907.4 44.4 5 p. m. 7 p. m. . 31.1 45.6 920.8 44.4 7 p. m. 9 p. m. . 32.0 49.3 919.5 51.2 9 p. m. 11 p. m. . 29.1 42.4 924.1 41.2 11 p. m. 1 a. m. . 24.1 32.3 933.0 24.1 1 a. m. 3 a. m. . 23.9 33.0 934.8 33.0 3 a. m. 5 a. m. . 24.2 32.4 939.1 30.7 Feb. 3-4 . 5 a. m. 7 a. m. . Total 24.9 34.4 944.9 30.7 .... 528.6 512.2 7 a. m. to 9 a. m. . 35.7 65.1 936.0 55.2 9 a. m. 11 a. m. . 25.6 44.8 948.5 44.5 11 a. m. 1 p. m. . 32.9 49.0 937.5 43.1 1 p. m. 3 p. m. . 31.3 51.0 943.1 49.6 3 p. m. 5 p. m. . 36.4 48.9 944.3 48.1 5 p. m. 7 p. m. . 31.7 47.5 950.8 42.6 7 p. m. 9 p.m.. 25.0 41.7 964.1 37.6 9 p. m. 11 p. m. . 26.0 40.2 968.1 37.7 11 p. m. 1 a. m. . 21.4 36.9 968.9 32.2 1 a. m. 3 a. m. . 23.4 33.4 972.6 35.0 3 a. m. 5 a. m. . 22.0 36.4 975.7 29.3 Feb. 4-5. 5 a. m. 7 a. m. . Total 23.8 34.8 980.1 34.3 .... 529.7 489.2 7 a. m. to 9 a. m. . 31.1 56.9 975.4 54.4 9 a. m. 11 a. m. . 32.0 44.5 975.6 40.7 11 a. m. 1 p. m. . 30.5 47.0 968.5 35.0 1 p. m. 3 p. m. . 36.5 49.3 961.9 52.3 3 p. m. 5 p. m. . 30.2 45.6 962.4 47.4 5 p. m. 7 p. m. . 36.3 51.7 963.5 44.4 7 p.m. 9 p. m. . 30.0 42.6 973.4 41.4 9 p. m. 11 p. m. . 34.1 44.0 971.5 43.6 11 p. m. 1 a. m. . 28.3 39.7 984.4 29.7 1 a. m. 3 a. m. . 27.2 32.7 986.3 37.6 3 a. m. 5 a. m. . 27.9 38.9 982.6 37.0 5 a. m. 7 a. m. . Total 28.9 34.3 985.3 31.4 527.2 494.9 1G0 Influence of Inanition on Metabolism. Table 110. — Elements katabolized in body — Metabolism experiment No. 74. (a) Total weight. (6) Nitro- gen. (c) Carbon. W) Hydro- gen. («) Oxygen. (/) Ash. First day, Feb. 2, 1905. Outgo: Water in urine Grams. 512.15 Grams. Grams. Grams. Grams. 512.15 I Grams. 1485.94 32.96 584.88 528.55 10^74 7\29 144 .'i4 166.28 1.97 65.45 1319.66 9.47 519.43 384.41 SA9 Solids in urine Water of respiration * Carbon dioxide Total 2632.33 2120.18 10.74 10.74 151.43 151.43 233 . 70 233.70 2232.97 1720.82 3.49 3.49 Loss Second day, Feb. 3, 1905. Income: Oxygen from air Outgo: Water in urine 489.17 .... .... .... 489.17 .... 1866.17 28.23 527.61 529.74 8^25 e!o6 144! 46 208.82 1.52 59.04 1657.35 8.23 468.57 385.28 4!i7 Solids in urine Water of respiration ' Carbon dioxide Total 2951.75 2462.58 8.25 8.25 150.52 150.52 269.38 269.38 2519.43 2030.26 4.17 4.17 Third day, Feb. Jh 1905. Income: Oxygen from air Outgo: Water in urine 494.92 .... .... .... 494.92 .... 1579.48 25.52 524.88 527.19 6\78 5!46 143.79 176.74 1.28 58.73 1402.74 7.18 466.15 383.40 4^82 Solids in urine Water of respiration 1 Carbon dioxide Total 2657.07 2162.15 6.78 6.78 149.25 149.25 236.75 236.75 2259.47 1764.55 4.82 4.82 Loss 1 Includes also water of perspiration. Table 111. — Elements and materials katabolized in body — Metabolism experiment No. 74. Date. (a) Nitro- gen. (6) Carbon. (c) Hydro- gen. (d) Oxy- gen. (e) Water. Protein. iff) Fat. (h) Carbo- hydrates (as gly- cogen). (i) Ash. 1905. Feb. 2-3 Feb. 3-4 Feb. 4-5 Total, 3 days. Grams. 10.74 8.25 6.78 Grams. 151.43 150.52 149.25 Grams. 233.70 269.38 236.75 Grams. 1720.82 2030.26 1764.55 Grams. 1887.12 2209.31 1917.78 Grams. 64.44 49.50 40.68 Grams. 138.30 112.48 124.26 Grams. 27.82 87.88 75.23 Grams. 3.49 4.17 4.82 25.77 451.20 739 . 83 5515.63 6014.21 154.62 375.04 190.93 12.48 Metabolism Experiment No. 74. 161 Output of Heat. The total heat production is recorded in the following table, together with the data from which it is derived. Table 112. — Summary of calorimetric measurements and total heat production — Metabolism experiment No. 74- Date. Period. (a) Heat meas- ured in terms Coo- Heat used in vaporiza- tion of water. (c) Sum of heat correc- tions.1 (d) Total heat produc- tion (a+b+c). 1905. Feb. 2-3 Feb. 3-4 Feb. 4-5 7 a.m. to 9 a.m Cals. 139.2 116.5 114.1 126.4 137.3 128.0 123.2 116.8 66.3 78.6 81.2 66.4 Cals. 30.2 31.6 30.7 35.7 31.0 31.9 30.1 31.1 24.3 27.9 24.3 26.3 Cals. + 8.4 + 5.6 + 14.9 + 20.1 — 10.4 + 0.6 — 4.4 — 7.9 — 2.0 + 10.9 — 2.0 + 8.5 Cals. 177.8 153.7 159.7 182.2 157.9 160.5 148.9 140.0 88.6 117.4 103.5 101.2 9 a.m. 11 a.m 1 p.m. 3 p.m 3 p.m. 5 p.m 5 p.m. 7 p.m 9 p.m. 11 p.m 1 a.m. 3 a.m 5 a.m. 7 a.m. ....... . Total 1294.1 355.0 + 42.3 1691.4 7 a.m. to 9 a.m 9 a.m. 11 a.m 148.1 100.5 109.7 120.0 112.3 116.3 123.0 84.9 71.3 66.7 81.2 71.2 24.6 29.2 25.7 30.1 27.5 30.2 25.3 25.9 25.3 25.3 24.4 26.2 + 13.5 - 0.5 + 19.2 + 22.5 - 0.3 + 6.2 - 8.3 - 10.2 + 2.4 + 8.1 - 0.3 + 8.0 186.2 129.2 154.6 172.6 139.5 152.7 140.0 100.6 99.0 100.1 105.3 105.4 11 a.m. 1 p.m 1 p.m. 3 p.m 3 p.m. 5 p.m 5 p.m. 7 p.m 7 p.m. 9 p.m 9 p.m. 1 1 p. m 11 p.m. 1 a.m 1 a.m. 3 a.m 3 a.m. 5 a.m 5 a.m. 7 a.m Total 1205.1 319.7 + 60.4 1585.2 7 a.m. to 9 a.m 153.3 102 . 4 115.2 103.5 101.7 128.3 115.8 115.6 52.2 69.8 92.4 73.6 30.0 28.0 26.2 27.8 27.6 30.4 27.3 26.7 25.3 25.7 25.1 24.5 + 6.1 + 3.1 + 17.5 + 31.3 - 0.8 — 1.0 - 5.0 — 13.8 + 11.8 + 5.8 + 1.0 + 2.1 189.4 133.5 158.9 162.6 128.5 157.7 138.1 128.5 89.3 101.3 118.5 100.2 9 a.m. 11 a.m 1 p.m. 3 p.m 3 p.m. 5 p.m 7 p.m. 9 p.m 9 a.m. 11 p.m 11 p.m. 1 a.m 1 a.m. 3 am 5 a.m. 7 a.m Total 1223.9 324.5 + 58.1 1606.5 1 See pp. 42-49. 11 162 Influence of Inanition on Metabolism. Balance of Energy. In table 113, the energy derived from the oxidation of body protein, fat, and carbohydrates during the time of this experiment is shown. On the last 2 days the comparison with the heat production shows large discrepancies, approximating 4 per cent. No satisfactory explanation for such discrepancies has as yet appeared. Relations between Oxygen Consumption, Carbon Dioxide Elimination, and Heat Production. The oxygen and carbon dioxide thermal quotients and the respiratory quotients obtained in this experiment are shown in table 114. Table 113. — Comparison of energy derived from Tcataoolized tody material with total heat production — Metabolism experiment No. 74. Date. Energy derived from different sources. From body protein. (a) Energy of protein katabo- lized. (b) Poten- tial energy of urine. (c) Net energy (a-b). (d) From body fat. (e) From body glyco- gen. (/) Total (d+e+/). Total heat produc- tion. Energy from body material greater ( + ) or less (— ) than output. (h) Amount cr-0). (i) Pro- portion (h + g). 1905. Feb. 2-3. Feb. 3-4. Feb. 4-5. Cals. 364 280 2-30 Cals. 88 68 61 Total, 3 days. 874 Av. per day.. 291 217 72 Cals. 276 212 169 Cals. 1319 1073 1185 657 219 3577 1192 Cals. Cals. 116 1711 368 1653 315 1669 799 5033 266 1678 Cals. 1691 1585 1607 Cals. + 20 + 68 + 62 Per ct. + 1.2 + 4.3 + 3.9 4883 1628 + 150 + 50 + 3.1 EFFECT OF INGESTION OF FOOD. Diet. — While the diet of the subject was very simple, it contained in addition to the two articles used in experiment No. 72, a small quantity of apple and a few graham crackers. The same amount of each article of food was consumed each day. The use of modified milk was continued in this experiment. Feces. — At 8.15 a. m., February 3, there were passed 61.4 grams of fresh feces, of which 23 grams were separated as belonging to food eaten before the fast began. The remainder, 38.4 grams, appeared more like " fasting feces " than the feces found in any other experiment following a period of fast but were not sufficiently colored with lampblack to warrant their acceptance as the result of fasting metabolism.31 When partially dried these feces gave 10.1 grams of air-dry material. Subsequently two other portions, on February 4 and 5, were passed which obviously belonged to the food period, and were put together to form one sample, amounting to 67.9 grams. It was, however, considered 31 For further discussion of fasting feces, see section on Feces, Part 3. Metabolism Experiment No. 74. 163 Table 114. — Oxygen and carbon dioxide thermal quotients and respiratory quotients — Metabolism experiment No. 74. Date and period. (a) Total heat produc- tion. Oxygen con- sumed. (c) Oxygen thermal quotient (100 b+a). (d) Carbon dioxide elimina- ted. (e) Carbon dioxide thermal quotient (lOOtl-5-a). (/) 1 (!/) Volume of Volume carbon of dioxide oxygen elimi- con- nated sumed (d X 0.5091). (bx 0.7). (ft) Respi- ratory quo- tient (/-s-ff). 1905 Feb. 2-3: 7 a.m. to 9 a.m. Cals. 177.8 Grams. 53 . 9 30.3 Grams. 52.6 29.6 Liters. 26.8 Liters. 37.7 .71 9 a.m. 11 a.m. 153.7 51.0 33.2 49.8 32.4 25.4 35.7 .71 11 a.m. 1 p.m. 159.7 51.8 32.4 50.2 31.5 25.6 36.2 .71 1 p.m. 3 p.m. 182.2 55.8 30.6 55.0 30.2 28.0 39.1 .72 3 p.m. 5 p.m. 157.9 44.4 28.1 51.6 32.7 26.3 31.1 .85 5 p.m. 7 p.m. 160.5 44.4 27.7 45.6 28.4 23.2 31.1 .75 7 p.m. 9 p.m. 148.9 51.2 34.4 49.3 33.1 25.1 35.8 .70 9 p.m. 11 p.m. 140.0 41.2 29.4 42.4 30.2 21.5 28.9 .75 11 p.m. 1 a.m. 88.6 24.1 27.2 32.3 36.4 16.4 16.8 .98 1 a.m. 3 a.m. 117.4 33.0 28.1 33.0 28.1 16.8 23.1 .73 o a.m. 5 a.m. 103 . 5 30.7 29.6 32.4 31.3 16.5 21.5 .77 5 a.m. Feb. 3-4: 7 a.m. 101.2 30.7 30.3 34.4 34.0 17.5 21.5 .82 1691.4 512.2 30.3 528 . 6 31.3 269.1 358.5 .75 7 a.m. to 9 a. m . 186.2 55.2 29.7 65.1 35.0 33.1 38.6 .86 9 a.m. 11 a.m. 129.2 44.5 34.4 44.8 34.6 22.8 31.1 .73 11 a.m. 1 p.m. 154.6 43.1 27.9 49.0 31.7 24.9 30.2 .83 1 p.m. 3 p.m. 172.6 49.6 28.7 51.0 29.6 26.0 34.7 .75 3 p.m. 5 p.m. 139.5 48.1 34.5 48.9 35.0 24.9 33.7 .74 5 p.m. 7 p.m. 152.7 42.6 27.9 47.5 31.1 24.2 29.9 .81 7 p.m. 9 p.m. 140.0 37.6 26.8 41.7 29.8 21.3 26.3 .81 9 p.m. 11 p.m. 100.6 37.7 37.5 40.2 40.0 20.5 26.4 .78 11 p.m. 1 a.m. 99.0 32.2 32.5 36.9 37.3 18.8 22.5 .84 1 a.m. 3 a.m. 100.1 35.0 35.0 33.4 33.4 17.0 24.5 .69 3 am. 5 a.m. 105.3 29.3 27.8 36.4 34.5 18.5 20.5 .90 5 a.m. Total. Feb. 4-5: 7 a.m. 105 . 4 34.3 489.2 32.5 34.8 529.7 33.0 33.4 17.7 24.0 .74 1585.2 30.9 269.7 342.4 .79 7 a.m. to 9 a.m. 189.4 54.4 28.7 56.9 30.0 29.0 38.1 .76 9 a.m. 11 a.m. 133.6 40.7 30.5 44.5 33.3 22.6 28.5 .79 11 a.m. 1 p.m. 158.9 35.0 22.1 47.0 29.6 24.0 24.5 .98 1 p.m. 3 p.m. 162.6 52.3 32.1 49.3 30.3 25.1 36.6 .69 3 p.m. 5 p.m. 128.5 47.4 36.9 45.6 35.5 23.2 33.2 .70 5 p.m. 7 p.m. 157.6 44.4 28.2 51 .7 32.8 26.3 31.1 .85 7 p.m. 9 p.m . 138.1 41.4 30.0 42.6 30.8 21.7 28.9 .75 9 p.m. 11 p.m. 128 . 5 43.6 33.9 44.0 34.2 22.4 30.5 .74 11 p.m. 1 a.m. 89.3 29.7 33.3 39.7 44.4 20.2 20.8 .97 1 a.m. 3 a.m. 101.3 37.6 37.2 32.7 32.3 16.7 26.3 .63 3 a.m. 5 am. 118.5 37.0 31.2 38.9 32.8 19.8 25.9 .77 5 a.m. Total. 7 a.m. 100.2 31.4 31.3 34.3 34.2 17.4 22.0 .79 1606.5 494.9 30. S 527.2 32.8 268.4 346.4 .78 164 Influence of Inanition on Metabolism. advisable to make complete analyses of both samples of feces in so far as practicable, and the more essential determinations were fortunately secured on both samples. It was impossible to determine fat in feces owing to the small amount of material. After it had been discovered that the method ordinarily Table 115. — Percentage composition of food and feces — Metabolism experiment No. 74. Labor- atory num- ber. Kind of material. fa) Water. (&) Pro- tein. (0) (d) (e) Fat. Carbo- hy- drates. Ash. P. ct. 12.03 P. ct. 4.86 P. ct. 0.66 > ■ ■ • 13.52 .22 .... 11.27 .37 10.32 77.77 1.63 !8.82 3.52 3.52 !6.56 12.62 6.77 (f) Nitro- gen. (9) Car- bon. (ft) Hydro- gen. W Heat of com- bus- tion per gram. 3826 3827 3828 3829 3838 3839 P. ct. Milk 79!47' Apple 86.11 Orange juice 87 . 87 Graham crackers. 5.10 Feces 75.08 Feces 67.51 P. ct. 2.98 .15 .49 5.18 9.06 6.54 P. ct. 0.48 .03 .08 .91 1.45 1.04 P. ct. P.ct. 12.28 1.92 5.70 .90 4.64 .76 44.46 7.08 13.32 1.91 17.02 2.57 Cals. 1.478 540 439 462 533 016 1 Assumed. See Analyses of Feces, Part 3, of this report. Table 116. — Weight, composition, and heat of combustion of food and feces- Metabolism experiment No. 74 (Quantities of food per day). (a) Weight grams. . (&) Water do (c) Protein do. . . . (d)Fat do (e) Carbohydrates, .do. .. . (/) Ash do (g) Nitrogen do. . . . (h) Carbon ... do. . . . (i) Hydrogen do. . . . (j) Oxygen (by difference) grams (k) Heat of combustion calories Food. 1 3826. Milk. 1201.00 953 . 64 35.76 144.36 58.32 7.92 5.76 147.36 23.04 62.28 1774 1 3827. Apple. 122.70 105.66 .18 16.59 .27 .04 6.99 1.10 8.64 66 1 3828. Orange juice. 312.90 274.95 1.53 35.26 1.16 .25 14.52 2.38 19.64 1 3829. Gra- ham crack- ers 35 . 00 1.79 1.81 3.61 27.22 .57 .32 15.56 2.48 14.28 156 Total for day. 1670.60 1336.04 39.28 147.97 137.39 9.92 6.37 184.43 29.00 104.84 2133 Feces. !3838. 38.40 28.83 3.48 3.39 1.35 1.35 .56 5.11 1.82 59 13839. 67.90 45.84 4.44 4.45 8.57 4.60 .71 1 1 . 56 1.75 3.44 137 Total for 3 days. 106.30 74.67 7.92 7.84 9.92 5.95 1.27 16.67 2.48 5.26 196 Aver- age per day. 35.43 24.89 2.64 2.61 3.31 1.98 .42 5.56 .83 1.75 65 1 Laboratory number. used for determining the fat in feces gives results that are too low (see below), it was found that unfortunately the samples for this experiment had been used up in making other analyses. The amount of fat in the feces was therefore estimated by the method explained under Analyses of Feces. Metabolism Experiment No. 74. 165 Percentage composition of food and feces. — The percentage composition and heat of combustion per gram of food and feces are given in table 115. The amounts of food and feces and the elements and nutritive ingredients, of which they consisted, together with the heat of combustion, are shown in table 116. Elements and materials absorbed from food. — From the analyses of the food and feces, the amounts of nitrogen, carbon, hydrogen, oxygen, and ash of the absorbed food were obtained. Since the food was identical on all 3 days, and since the total amount of feces belonging to the experiment was apportioned equally among the different days of the experiment, the only variations in the quantities of elements absorbed on the different days were such as occurred Table 117. — Elements absorbed from food — Metabolism experiment No. 14. (a) Total weight. (6) Nitrogen. (c) Carbon. (d) Hydrogen. (e) Oxygen. (f) Ash. First day, Feb. 2, 1905. Food and drink : Solids in food Orams. 334.56 1336.04 603.30 Orams. 6.37 Orams. 184.43 Orams. 29.00 149.50 67.51 Orams. 104.84 1186.54 535.79 Orams. 9.92 Water in food Water in drink Total 2273.90 6.37 184.43 246.01 1827.17 9.92 Feces : Solids 10.54 24.89 .42 5.56 .83 2.79 1.75 22.10 1.98 Water Total.. 35.43 .42 5.56 3.62 23 . 85 1.98 2238.47 5.95 178.87 242.39 1803.32 7.94 JFor February 3, the amounts of hydrogen and oxygen absorbed are 289.30 and 2175.61 grams respectively ; for February 4, the corresponding amounts are 256.59 and 1916.02 grams. as a result of different amounts of water drunk. The elements absorbed on February 2 are shown in table 117, and to avoid repetition, the amounts of hydrogen and oxygen absorbed on the second and third days of the experiment are given in a note following the table. As has been explained in considering previous experiments, the quantities of absorbed material are computed in terms of body protein, body fat, and glycogen, by means of the formulas given on page 38. The results show that on the first day there were absorbed 35.70 grams of protein, 133.80 grams of fat, and 131.52 grams of carbohydrate. Although there were fluctuations in the quantity of water consumed on the two subsequent days, the quantities of nitrogen, carbon, hydrogen of organic matter, and oxygen of organic matter were constant on all days, and the same amounts of materials were absorbed from the food daily. In carrying out the calculations by means of the formulas, however, slight differences are obtained in the amounts of fat and carbohy- 166 Influence of Inanition on Metabolism. drates for each of the 3 days, but the differences are mostly in the second decimal place. These variations may be noted in lines d and g of table 121 beyond. Amounts of ingredients of food absorbed and corresponding amounts of body materials. — A comparison of the amounts of food protein, fat, and carbo- hydrates determined by proximate analyses with the computed amounts of body protein, fat, and carbohydrates derived from the elementary analyses is shown in table 118. It has previously been stated that the quantities of food were the same each day. Since the feces for the experiment were divided equally among the 3 days, the data for all 3 days are identical. Energy of material absorbed from food. — The nutrients of the absorbed food expressed in terms of body materials, yielded for each day the following Table 118. — Amounts of ingredients of food absorbed and body materials derived from them — Metabolism experiment No. 74 (quantities per day). (a) Food. (6) Feces. (o) Absorbed (a-b). (d) Body material.1 Protein grams . Fat .do. . . Carbohydrates do . . . Ash do. . . Energy calories. 3y.28 147.97 137.39 9.92 2133 2.64 2.61 3.31 1.98 65 36.64 145.36 134.08 7.94 2068 35.70 133.85 131.48 7.94 2030 1 The amounts of water absorbed as calculated bv the formulae were as follows : Feb. 2, 1930.06 grams ; Feb. 3, 2349.24 grams, and Feb. 4, 2056.95 grams. amounts of energy: From protein, 202 calories; from fat, 1277 calories; and from carbohydrates, 551 calories, a total of 2030 calories. Changes in body-weight compared with balance of income and outgo. — The comparison made in table 119 has been previously explained for similar tables. When the multiplicity of weighings is taken into consideration, the balance obtained in this experiment is extremely satisfactory. Balance of Intake and Output. In order to determine to what extent the food supplied the necessary material for the metabolic activity during this experiment, the balance tables 120 and 121 were prepared. There was an increasing gain of water throughout the experiment. The daily loss of protein decreased rapidly as the experiment progressed. The body was nearly in equilibrium as regards fat, while there was a marked storage of carbohydrates and ash on all days. The diet furnished an average excess of 368 calories of energy. Metabolism Experiment No. 74. 167 Table 119. — Comparison of changes in body weight with balance of income and outgo — Metabolism experiment No. 74- Feb. 2-3. Feb. 3-4. Feb. 4-5. Total for 3 days. Average per day. Income: (a) Food Grams. 1(570.60 603.30 512.15 Grams. 1670.60 1022.50 489.17 Grams. 1670.60 730.20 494 . 92 Grams. 5011.80 2356.00 1496.24 Grams. 1670.60 785.33 498.75 (b) Water consumed (d) Total (a+b + c) 2786.05 3182.27 2895.72 8864.04 2954.68 Outgo : 1433.10 528.55 584.88 1716.40 61.40 529 . 74 527.61 1603.00 30.00 527.19 524.88 4752.50 91.40 1585.48 1637.37 1584.17 30.47 528.49 545.79 (/) Feces , {h) Water of respiration and per- spiration (i) Total (e +f+ g + h) 2546.53 2835.15 2685.07 8066.75 2688.92 (J) Gain of body material (d—i).. (k) Gain of body weight 239 . 52 231.00 347.12 342 . 00 210.65 200.00 797.29 773.00 265.76 257.67 1 The data in this line should not be confounded with urine data in other tables, explanation, p. 66. See Table 120. — Distribution of intake and outgo of water — Metabolism experiment No. 74. Date. Outgo from the body. Balance of preformed water. to) Water of oxidation of organic hydrogen (C-d). (a) Water of urine and feces.1 (b) Water of respira- tion and perspira- tion. (c) Total (a + b). (d) Pre- formed (katabol- ized) water in outgo.2 (e) Intake in food and drink. (/) Gain of pre- formed water (d-e). 1905. Feb. 2-3 Feb. 3-5.. Grams. 1485.9 1909.3 1600.6 Grams. 584.9 527.6 524.9 Grams. 2070.8 2436.9 2125.5 Grams. 1887.1 2252.4 1938.9 Grams. 1939.3 2358.5 2066.3 Grams. 52.2 106.1 127 .4 Grams. 183.7 184.5 186.6 Feb. 5-6 Total, 3 days. . Av. per day. . . . 4995.8 1665.3 1637.4 545.8 6633.2 2211.1 6078.4 2026.1 6364.1 2121.4 285.7 95.3 554.8 184.9 1 Water In feces passed on the second and third days assumed as 43.13 and 21.07 grams re- spectively. In obtaining these amounts it is assumed that water existed in the feces for these two days in the same proportion that it did in the total feces of the experiment. 2 Figures for second and third days include assumed water in feces. 168 Influence of Inanition on Metabolism. Table 121. — Balance of intake and output of nutrients, ash, and energy — Metabolism, experiment No. 74- Feb. Feb. Feb. Total for Average 2-3. 3-4. 4-5. 3 days. per day. Body protein : (a) Computed from elements absorbed from food. . . .gms. . 35.70 35.70 35.70 107.10 35.70 64.44 49.50 40.68 154.62 51.54 28.74 13.80 4.98 47.52 15.84 Body fat: (d) Computed from elements ab- 133.80 133.93 133.83 401.56 133.85 138.30 112.48 124.26 375.04 125.01 (/) Gain (+) or loss ( — ) to body -4.50 + 21.45 + 9.57 + 26.52 + 8.84 Body carbohydrates : (g) Computed from elements ab- sorbed from food gms. . 131.52 131.41 131.50 394.43 131.48 27.82 103.70 87.88 43.53 75.23 56.27 190.93 203.50 63.64 67.83 Ash: 7.94 7.94 7.94 23.82 7.94 3.49 4.17 4.82 12.48 4.16 4.45 3.77 3.12 11.34 3.78 Energy: (m)Of food absorbed (determined) 2068 2068 2068 6204 2068 M Heat production plus potential 1779 1653 1668 5100 1700 289 415 400 1104 368 Metabolism Experiment jSTo. 75. 169 Metabolism Experiment No. 75. Immediately following the 5-day fast, experiment No. 73, the subject had remained in the respiration chamber for 3 days with food. From the morning of February 5, until the evening of March 3, he had been engaged in light occupation, assisting about the laboratory. During this preliminary period his diet had consisted in large part of vegetable materials, milk, and eggs. On the evening of March 3, 1905, he entered the respiration chamber and went to bed at 11 p. m. At 1 a. m., March 4, the preliminary analyses of the respiratory products were begun. The experiment proper commenced at 7 a. m., March 4, and continued 7 days, during which time no food was eaten. The general routine was similar to that followed in previous experiments with this subject. This experiment was the most successful of the series conducted with S. A. B. Though he made a large number of notes concerning various symptoms of illness, nevertheless his verdict at the end of each day (except the sixth), was that he had " passed a comfortable day." Notes from diary, pulse records, and records of body movements. — The notes from the diary of the subject which embrace practically all his observations concerning his physical and mental condition, are shown below, together with the pulse records and the record of body movements. Notes from diary. March 4, 1905: 7h15m a. m. Passed a fairly good night, though I awoke a few times. 9h10m a. m. Defecated. 9h15m a. m. Commenced wearing rectal thermometer. 2h30m p. m. Feel sleepy. 3h40™ p. m. Awoke from a nap lasting about 1 hour. 4 p. m. Am commencing to have a sour taste in the mouth; tongue coated thinly with a white fur. 10 p.m. Have a slight headache; eyes feel tired and I am very sleepy. Passed a very comfortable day ex- cept for slight pain in the heart. March 5, 1905. 7h15m a. m. Slept fairly well but did not feel like rising. Am nervous and very weak. 7h30m a. m. Have an acrid taste in my mouth; tongue is coated with a whitish fur, and is swollen. 8h10m a. m. Had an inclination to defe- cate. 8''20m a. m. Feel very weak. 9h10m a. m. Made another ineffectual at- tempt to defecate. 10 p. m. Passed a comfortable day. March 6, 1905: 7h15m a. m. Slept fairly well, though not continuously. Feel weak and ex- tremely nervous. Tongue is swollen and coated with a white fur; sweet- ish taste in mouth. Eyes are clear and bright. 8h10m a. m. Attempted to defecate, but could not. Colon is filled with feces and if they could be removed I would feel considerably better. 9"50m a. m. Made another ineffectual at- tempt to defecate. 10 p. m. Have passed a very comfortable day. The weakness I felt in the morning passed away later. 170 Influence of Inanition on Metabolism. March 7, 1905: 7h20'" a. m. Slept very well. My tongue is swollen and coated with a brown- ish-white fur; sweet taste in mouth. Eyes are clear and bright. Feel a little weak. 7h30m a. m. Rectal thermometer is com- mencing to irritate me. 9b40m a. m. Took off rectal thermometer. 12h10m p. m. Made an unsuccessful at- tempt to defecate. 2h22m p. m. Commenced to wear rectal thermometer. It does not feel com- fortable. 5h55m p.m. Took off rectal thermometer. Tried the soap suppository, but it caused considerable pain and I was compelled to remove it. Soreness in rectum. 10 p. m. Passed a comfortable day. Feel a little weaker. March 8, 1905: 7h30m a. m. Slept continuously until 5h50m a. m. Feel a little weak and des- pondent. Tongue swollen and thick- ly coated; sweet taste in mouth. Eyes are bright and clear; face does not show any signs of fasting. 10 p. m. Passed a very comfortable day. At times I felt sleepy and managed to get a few naps. March 9, 1905: 7h20m a. m. Slept very well. My tongue is swollen and coated with a thick white fur. Eyes are clear and bright. Feel a little weak as usual, but my nerves are stronger. S''10m a. m. Made an unsuccessful at- tempt to defecate. 3b40m p. m. A slight bilious headache is commencing to come on; my eyes are painful and blurred. 5h20m p. m. Tongue and gums sore. 9h10m p. m. Made another unsuccessful attempt to defecate. 9h18m p. m. Commenced to wear rectal thermometer. Feels all right. 10 p. m. Felt very badly all day, but im- proved in the evening, and feel con- siderably stronger now. March 10, 1905: 7h20m a. m. Did not sleep continuously for one-half hour. The sore gums and swollen tongue made that an impossibility; tongue is coated with a white fur. lh25m p. m. Took off rectal thermometer. 10 p. m. Have passed a very good day, except for swollen tongue. Feel con- siderably stronger than at any other time. Could do some hard work. Pulse-rate — Experiment No. 75.1 Time. Pulse rate. Time. Pulse rate. Time. Pulse rate. Mar. 4, 7h30» a.m. . 68 Mar. 6, 12h00ma.m. . 56 Mar. 8, e^OO" p.m. . 54 8 00 a.m. . 66 2 00 p.m. . 58 8 00 p.m. . 51 10 00 a.m. . 59 4 00 p.m. . 58 10 00 p.m. . 48 12 00 a. m. . 55 6 00 p.m. . 66 Mar. 9, 7 30 a.m. 54 2 00 p.m. . 257 8 00 p.m. . 53 8 00 a.m. . 54 4 00 p.m. . 3 52 10 00 p.m. . 49 10 00 a.m. . 854 6 00 p.m. . 53 Mar. 7, 7 30 a.m. . 59 12 00 a.m. . 3 52 8 00 p.m. . 52 8 00 a.m. . 61 2 00 p.m. . 48 10 00 p.m. . 51 10 00 a.m. . 52 6 00 p.m. . 44 Mar. 5, 7 30 a.m. . *82 12 00 a.m. . 58 8 00 p.m. . 45 8 00 a.m. . 68 2 00 p.m. . 55 10 00 p.m. . 47 10 00 a.m. . 62 4 00 p.m. . 54 Mar. 10, 7 30 a.m. . 57 12 00 a.m. . 64 6 07 p.m. . 52 8 00 a.m. . 55 2 00 p.m. . 61 8 00 p.m. . 53 10 00 a.m. . 54 4 00 p.m. . 67 10 00 p.m. . 49 12 00 a.m. . 51 6 00 p.m. . 461 Mar. 8, 7 35 a.m. . 74 2 10 p.m. . 50 8 00 p.m. . 54 8 00 a.m. 7 58 4 00 p.m. . 52 10 00 p.m. . 353 10 00 a.m. . 53 6 00 p.m. . 50 Mar. 6, 7 30 a.m. . 70 12 00 a.m. . 53 8 05 p.m. . »44 8 00 a.m . . 5 63 2 00 p.m. . 56 10 00 p.m. . 48 10 00 a.m. . «55 4 00 p.m. . 54 1 Pulse taken while sitting. 2 Very Irregular. 8 Very faint. 4 Irregular. B Faint and irregular. 6 Regular. 7 Strong and regular. 8 Regular but faint. 9 Very faint but regular. Metabolism Experiment No. 75. 171 Movements of subject, duration 7 days, from Mar. 4, 7 a. m.t to Mar. 11, 7 a. m., 1905. March 4- p. M. A. M. A. M. 3h 48m rise, food aperture, 7h 30m count pulse. 7b A Am 1 sit, rise, open cur- sit. 7 50 drink. I UU V tain, urinate, 3 50 drink. 8 00 count pulse. 7 uo | weigh self, etc. 3 56 write. 8 02 remove thermome- 7 16 ' food aperture. 4 00 count pulse. ter. 7 22 dress, raise table. 4 10 read. 8 04 food aperture. 7 24 comb hair, tele- 4 28 rise, food aperture, 8 06 food aperture. phone. sit. 8 10 close curtain, at- 7 25 rise, food aperture, 4 32 read. tempt to defecate. drink. 5 00 drink. 8 18 open curtain, sit. 7 26 sit. 5 01 rise, food aperture, 8 20 drink. 7 30 count pulse. sit. 8 38 head on table. 7 44 telephone. 5 04 read. 9 02 food aperture. 7 46 move about, sit. 5 10 move about. 9 04 move about, drink. 7 50 drink. 5 20 stand, doctor count 9 06 food aperture. 7 53 food aperture, sit. pulse. 9 10 close curtain, at- 8 00 count pulse. 5 22 sit, read. tempt to defecate. 8 06 telephone. 6 00 count pulse, drink. 9 12 open curtain. 8 08 read. 6 36 change position. 9 16 sit. 8 15 drink. 6 40 drink. 9 20 adjust thermome- 9 00 drink. 7 02 move about. ter. 9 02 rise, move about. 7 04 urinate, telephone. 9 26 asleep. 9 04 food aperture. 7 06 move about, food 9 32 awake. 9 08 close curtain. aperture, sit. 9 43 drink. 9 10 defecate. 7 22 telephone. 9 44 asleep. 9 15 adjust thermome- 7 35 drink. 9 48 awake. ter. 7 42 rise, food aperture, 10 00 count pulse. 9 20 open curtain, sit. sit. 10 12 rise, food aperture, 9 30 read. 8 00 count pulse. sit. 9 40 drink. 8 18 change position. 10 14 read. 9 43 rise, food aperture, 8 36 drink. 11 35 drink. sit. 9 34 rise, stand. 11 50 move about. 9 53 rise, food aperture. 9 36 rise, telephone. 12 00 count pulse. 10 00 count pulse. 9 58 sit, telephone. p. M. 10 15 drink. 10 00 count pulse. 12" 10m rise, food aperture, 10 22 telephone, read. 10 10 rise, move about. sit. 10 50 drink. 10 12 telephone, sit. 12 12 read. 11 04 rise, move about, 10 14 move about, food 12 30 drink. urinate. aperture. 1 02 rise, food aperture, 11 06 sit. 10 16 open bed, make urinate, sit, read. 11 34 write. motions, write. 1 25 drink. 11 35 drink. 10 20 sit. 2 00 count pulse. 11 42 read. 11 02 rise, urinate, un- 2 28 stop reading. 12 00 count pulse. dress. 2 56 asleep. p. 12f M. 30m drink. A M. March 5. 4 4 04 08 awake, count pulse, asleep, awake, rise, food 1 02 telephone, rise, 7* 02" open curtain, rise. aperture. food aperture. 7 04 urinate, fold bed, 4 10 move about, drink, 1 30 drink, read. telephone. sit. 1 2 44 00 rise, food aperture, count pulse. 7 7 05 09 1 weigh self, etc. 4 4 24 26 rise, food aperture, sit, read. 2 02 rise, stretch self, 7 12 food aperture. 5 16 rise. sit. 7 16 dress. 5 20 drink. 2 04 read. 7 20 raise table, sit, tel- 5 22 stand, doctor count 2 24 head on table, ephone. pulse. asleep. 7 25 drink. 5 26 sit, read. 3 14 awake, stretch self. 7 26 comb hair, tele- 6 00 count pulse. 3 16 asleep. phone. 6 45 drink. 3 44 awake. 7 28 stand, telephone. 6 54 telephone. 172 Influence of Inanition on Metabolism. Movements of subject. — Continued. March 5 (cont.) p. M. 6h 58m rise, stand, sit. 7 04 rise, food aperture, urinate, stand. 7 06 move about, sit. 7 45 drink. 8 00 count pulse. 8 10 telephone. 8 26 rise, lean on table. 8 28 move about, sit. 8 50 rise, lean on table, sit. 9 00 drink. 9 06 rise, food aperture, telephone. 9 45 drink. 10 00 count pulse. 10 08 rise, lower table, open bed, sit. 10 18 telephone. 10 46 lie. 11 04 rise, urinate, un- dress, close cur- tain. March 6. A. M. 7h 00m rise, open curtain. 7 02 urinate, fold bed, weigh self, etc. 7 14 dress, adjust table. 7 16 food aperture, sit, comb hair. 7 25 drink. 7 30 count pulse. 7 36 read. 8 00 count pulse. 8 05 drink. 8 08 move about, food aperture. 8 12 remove thermome- ter. 8 14 attempt to defe- cate. 8 18 sit. 8 45 drink. 9 02 rise, food aperture, move about. 9 14 move about. 9 16 stand, blood sample taken. 9 20 move about. 9 26 stand. 9 34 sit. 9 36 rise, food aperture. 9 50 attempt to defe- cate, drink. 9 52 move about. 9 56 adjust thermome- ter. 10 00 count pulse. A. M. 10" 02m write. 10 34 move about. 10 35 drink. 10 38 sit. 10 46 read. 11 00 move about. 11 07 food aperture. 11 08 work at food aper- ture. 11 26 sit. 11 48 food aperture, move about. 11 50 drink. 12 00 count pulse. p. M. 12h 02m open bed, lie. 12 10 food aperture, lie, asleep. 1 02 awake, move, rise, food aperture. 1 08 urinate, fold bed. 1 14 food aperture, tele- phone. 1 16 raise table, ar- range books, sit, drink. 1 55 drink. 2 00 count pulse. 2 02 read. 3 00 drink. 3 02 move about, blood sample taken, stand. 10 sampling finished. 12 food aperture. 14 sit. 00 count pulse. 05 drink. 50 drink. 02 rise, food aperture. 10 stand. 26 food aperture. 28 stand, doctor count pulse. 34 food aperture. 36 read. 6 00 count pulse. 6 05 drink. 32 rise, lean on table. 00 drink. 02 rise, food aperture. 04 urinate, sit, read. 06 telephone. 26 rise, food aperture. 35 rise, food aperture. 55 drink. 8 00 count pulse. 8 18 rise, move about, sit. 8 20 rise, telephone. 3 3 3 4 4 4 5 5 5 5 5 5 6 7 7 7 7 7 7 7 P. M. 8h24n 8 26 9 00 9 05 9 34 9 55 10 00 10 02 10 04 10 08 10 10 10 30 11 02 11 04 11 06 A. M. 7h00n 7 04 7 08 7 11 7 14 7 16 7 25 7 30 7 34 7 36 8 10 9 00 9 04 9 06 9 14 9 16 9 20 9 40 0 9 9 9 9 42 44 45 48 50 10 00 10 05 10 10 10 12 10 55 11 30 11 32 11 50 12 00 P. M. 12h 02m 12 05 ' telephone, rise, sit (3 times), drink. food aperture, rise, urinate, walk about, sit. drink. count pulse, rise, food aperture, stand, lower table, open bed. move about, move papers, recline, telephone, rise, urinate, undress, close curtain. March 7. rise, open curtain. 1 urinate, weigh self, etc. food aperture. dress, raise table, sit. comb hair. drink. count pulse. telephone. rise, food aperture, sit. drink. drink. move about. sit, blood sample taken. food aperture. move about. sit, read. remove thermome- ter. food aperture. move about. drink. read. food aperture. count pulse. food aperture. food aperture. urinate, sit, write. drink. rise, food aperture. sit, read. drink. count pulse. food aperture, food aperture. Metabolism Experiment No. 75. 173 Movements of subject. — Continued. March 7 (cont.) p M. 12' 1 06" move about. 12 10 attempt to defe- cate. 12 12 sit, write. 12 50 drink. 1 02 move about, food aperture, urinate. 1 06 sit, write. 1 08 stand. 1 15 food aperture. 2 00 count pulse. 2 05 drink. 2 16 move about. 2 18 write. 2 28 adjust thermome- ter. 3 02 move about. 3 04 urinate, sit, read. 3 14 move about. 3 16 food aperture. 3 18 sit, read. 3 20 drink. 4 00 count pulse. 4 05 food aperture. 4 10 move about, uri- nate. 4 12 sit, read. 5 08 stand. 5 14 move about, stand, doctor count pulse. 5 24 move about. 5 28 read. 5 30 remove coat. 5 34 sit. 5 36 adjust sphygmoma- nometer. 5 40 remove apparatus, put on coat. 5 42 stand. 5 44 sit. 5 45 drink. 5 46 read. 5 50 food aperture. 5 52 remove thermome- ter. 5 54 move about. 6 05 sit. 6 07 count pulse. 6 10 food aperture. 6 14 read. 6 40 telephone. 7 01 rise, food aperture. 7 04 sit, read, telephone. 7 08 rise, food aperture. 7 20 rise, move chair, sit. 7 25 drink. P. M. p M. 7h32ir rise, urinate, sit. 121 1 06" stand, sit. 7 36 rise, food aperture. 12 14 food aperture. 8 00 count pulse. 12 28 food aperture. 8 04 rise, stand. 12 55 drink. 8 13 9 02 food aperture, sit. telephone. 1 02 food aperture, uri- nate, count pulse. 9 35 9 40 drink. rise, urinate. 2 00 10 00 count pulse. 2 06 write. 10 10 rise, move about, 2 25 drink. lower table. 3 02 move about. 10 12 open bed, read, lie. 3 06 stand, blood 10 20 drink. sample taken. 10 52 asleep. 3 18 stand. 11 04 rise, urinate, un- 3 20 food aperture. dress. 3 22 sit. 11 06 close curtain. 3 33 drink. March 8. 3 40 food aperture. A. M. 4 00 count pulse. 7h02m rise, open curtain. 4 04 read. 7 04 urinate. 4 50 drink. 7 09 1 we*gn se^> etc- 5 5 04 06 rise, move about, stand, doctor count 7 10 fold bed. pulse. 7 12 dress, raise table, 5 08 sit. sit. 5 10 move about. 7 16 7 20 comb hair. rise, food aperture, sit. drink, count pulse. 5 12 blood pressure test, coat off. 7 32 5 18 put on coat. 7 34 telephone. 5 20 sit. 7 42 food aperture. 5 22 read. 7 50 write. 5 34 food aperture. 7 55 drink. 6 00 count pulse. 8 00 count pulse. 6 40 drink. 8 04 move about. 6 48 doze. 8 06 stand, blood 6 59 food aperture. sample taken. 7 04 rise, urinate, tele- 8 10 drink. phone, food aper- 8 16 move about. ture sit 8 20 8 26 8 46 lean on table, read, food aperture, sit. 7 8 32 00 ■ ill vj UX\j» rise, food aperture, count pulse. 10 00 count pulse. 8 15 drink. 10 02 move about, uri- nate. 9 18 telephone, food ap- erture. 10 06 sit. 9 55 drink. 10 12 asleep. 10 00 count pulse, food 10 54 awake. aperture. 10 56 move about. 10 04 rise, move about. 11 04 asleep. 10 08 lower table, open 11 16 food aperture. bed. 11 19 food aperture. 10 12 read, lie. 11 24 asleep. 10 14 telephone. 11 26 awake. 10 16 move about, adjust 11 30 drink. telephone. 11 44 food aperture. 11 02 rise, urinate, un- 11 46 move about. dress. 12 00 count pulse. 11 04 close curtain. 174 Influence of Inanition on Metabolism. Movements of subject. — Continued. March 9. p. M. A M. A . M. 5l lQm telephone. 9l 14m sit, read. 7 1 02m rise, open curtain, 5 20 count pulse. 10 00 count pulse. urinate. 5 26 food aperture. 10 04 food aperture. 7 04 \ telephone, weigh 5 28 stand, doctor count 10 06 sit. 7 07 / self, etc. pulse. 10 25 drink. 7 16 dress, raise 1 table, 5 30 sit, write. 11 02 food aperture, sit, repair tele- 5 32 coat off, blood pres- sure test. move about. phone. 5 36 11 04 sit. 7 18 food aperture, sit. 5 40 move about, sit, 11 22 food aperture. 7 20 food aperture , re- write. 11 34 drowsy. pair telephone, 5 46 food aperture. 11 52 asleep. comb hair. 5 50 read. 12 00 awake, count pulse. 7 25 drink. 6 00 count pulse. T> 7 7 27 30 food aperture, count pulse. 6 6 14 40 asleep. awake, drink, read. if. ivi.. 12h 02m food aperture. 7 50 food aperture. 7 04 food aperture, uri- 12 U4 read. 7 54 read. nate. 12 15 drink. 8 00 count pulse. 7 10 stand, read, look 12 55 drink. 8 10 attempt to defe- around. 1 02 move about, food cate. 7 14 rise, telephone, sit. aperture. 8 15 drink. 7 35 write. 1 06 urinate. 8 16 move about. 7 38 telephone. 1 08 sit. 8 22 sit. 7 45 food aperture. 1 10 remove thermome- 9 00 move about, drink. 7 50 drink. ter. 9 04' > stand, blood f pled. sam- 8 00 count pulse. 1 25 write. 9 06 8 35 drink. 1 48 food aperture. 9 11 food aperture. 9 08 rise, close curtain, 1 53 food aperture. 9 12 read. attempt to defe- 2 02 write. 9 32 food aperture. cate. 2 08 count pulse. 9 34 move about. 9 10 move about, adjust 2 18 food aperture. 9 44 read. thermometer. 2 35 drink. 10 00 count pulse. 9 12 open curtain. 3 02 ) stand, blood sam- 10 04 food aperture. 10 00 count pulse, write. o 6 04 J pled. 10 08 write. 10 05 drink. 3 10 sit. 10 45 drink. 10 06 stand, lower table, 3 16 read. 11 24 food aperture. open bed, recline, 3 32 rise, move about. 11 28 read. read. 4 00 count pulse. 12 00 count pulse. 11 04 rise, urinate, un- 4 10 drink. p. M. dress. 4 28 telephone. 12h 02m food aperture. 11 06 close curtain. 4 46 stop telephoning, 12 06 read. March 10. read. 12 30 drink. A. M. 5 04 telephone. 1 00 rise, urinate, i move 7h 7 04m 10 1 rise, open curtain, 5 06 stand. about. >■ urinate, fold bed, 5 08 sit. 1 04 food aperture. J weigh self, etc. 5 10 stand, doctor count 1 24 food aperture. 7 14 brush hair, tele- pulse. 2 2 00 02 count pulse, read. 7 15 phone, food aperture. 5 5 12 \ 14 f blood pressure test. 3 00 drink. 7 25 drink. 5 20 put on coat. 3 02 move about. 7 30 count pulse, write. 5 22 stand, telephone. 3 06 stand, blood sam- 7 38 comb hair. 5 25 drink. pled. 7 52 food aperture. 5 26 food aperture. 3 16 sit, read. 8 00 count pulse. 5 28 sit, stand, sit. 3 36 write. 8 15 drink. 5 52 read. 3 48 asleep. 8 20 read. 6 00 count pulse. 5 00 awake. 9 05 drink. 6 10 drink. 5 02 doze. 9 06 ) stand, blood sam- 7 00 urinate. 5 15 drink. 9 08 J pled, move about. 7 08 telephone. Metabolism Experiment No. 75. 175 March 10 (cont.) p. M. 7h 12m rise, food aperture, telephone, sit. 7 24 rise, telephone, move chair, blood pressure test, sit. 7 36 rise, move about, telephone. Movements of subject. — Continued. p. M. 7 46 48 05 12 00 sit, blood pressure test, rise, move chair, stand, read, food aperture, count pulse, telephone, drink. p. M. 10" 00m count pulse. 10 10 drink, rise, lower table, open bed, recline, read. 11 00 rise, undress, uri- nate. 11 04 close curtain. Drinking-water. — Table 122 records the daily consumption of water, and the amounts taken in different 2-hour periods. In this experiment the subject measured and recorded the amount of water drunk. The table shows that on different days, there is no uniformity in the amounts of water drunk in any given period. The amounts consumed were less than in the previous fasting experiment. The average for the 7 days was not far from 1800 grams per day. Table 122 — Record of water consumed 1 — Metabolism experiment Nc . 75. Date. 7to9 9 to 11 11 a.m. lto3 3 to 5 5 to 7 1 to 9 9 to 11 Total for da3'. a. m. a. m. to 1 p.m. p. m. p. m. p. m. p. m. p. m. 1905. Grams. Grams. Grams. Grams. Grams. Grams. Grams. Grams. Grams. 300.0 313.5 172.7 200.0 193.0 395 . 8 323.3 75.0 1973.3 Mar. 5-6 275.0 343.6 326.5 175.0 150.0 172.5 150.0 236.3 1728.9 Mar. 6-7 396.6 300.0 91.4 50.0 399.7 125.0 293.1 361.8 2117.6 Mar. 7-8 275.0 393.1 296.0 200.0 193.0 163.1 200.0 ! 191.1 1911.3 Mar. 8-9 250.0 146.3 394.8 175.0 121.8 138.9 102.3 i 152.5 1581.6 Mar. 9-10... 300.0 289.5 192.8 200.0 328.9 250.0 140.8 1702.0 Mar. 10-11... 300.0 217.8 267.2 150.0 150.0 211 . 1 125.0 [ 272.0 1693.1 1 Period during which water was consumed, was assumed in some instances. (See page 73.) Urine. The usual method of collecting the urine in four periods was followed. In the samples obtained each period, the weight, specific gravity, reaction, nitrogen, phosphoric acid by titration, and creatinine were determined. The results of these determinations are recorded in table 123. The specific gravity was invariably low and the reaction acid. The agreement between the deter- minations of nitrogen, phosphoric acid, and creatinine, in the different periods, with the total determined on the daily composite was in all cases very satis- factory. The results recorded in, column g of the table indicate the amount of preformed creatinine excreted in the urine and do not include the preformed creatine which was subsequently determined and is recorded in table 124, page 178. Weight, composition, and heat of combustion of urine. — In addition to the determinations made on the samples of urine collected each period, a much larger number of determinations were made on the composite urine for each day. From the percentages thus obtained and the weight of urine, the quantities of elements and compounds excreted have been computed and 176 Influence of Inanition on Metabolism. Table 123. — Determinations in urine per period and per day — Metabolism experiment No. 75. Date and period. (a) Amount. CM Specific gravity. (c) Volume (a+6). (a) Reaction. (e) Nitro- gen. (/) Phos- phoric acid by titra- tion (P2O5). (0) Cre- atinine ex- creted (pre- formed) 1905. Mar. 4-5: 7 a.m. to 1 p.m. . . . Grams. 410.9 1.0090 c. c. 407 Grams. 4.10 Grams. 0.383 Grams. 0.340 1 p.m. 7 p.m. . . . 467.1 1.0059 464 Acid 3.30 .394 .330 7 p.m. 11 p.m. . . . 208.7 1.0077 207 Acid 1.63 .207 .182 11 p.m. 7 a.m. . . . Total 421.5 1.0089 418 3.21 .468 .376 1508.2 1508.2 1.0079 1496 1496 12.24 12.20 1.452 1.460 1.228 1.212 Total by composite. Mar. 5-6 : 7 a.m. to 1 p.m. . . . 777.8 1.0054 774 3.82 .591 .313 1 p.m. 7 p.m. . . . 455.2 1.0097 451 Acid 3.30 .802 .256 7 p.m. 11 p.m. . . . 138.9 1.0164 137 1.67 .370 .144 ; 11 p.m. 7 a.m. . . . Total 513.8 1.0073 509 Acid 3.66 .538 .332 ! 1885.7 1885.7 1.0077 1871 1871 12.45 12.40 2.301 2.245 1.045 1.061 Total by composite. Mar. 6-7 : 7 a.m. to 1 p.m. . . . 503.2 1.0066 500 3.55 .453 .236 1p.m. 7 p.m.... 810.7 1.0040 807 Acid 3.73 .635 .222 7 p.m. 11 p.m. . . . 313.6 1.0074 311 2.10 .390 .134 11 p.m. 7 a.m. . . . Total 593.5 1.0061 590 3.64 .626 .281 2221.0 2221.0 1.0057 2208 2208 13.02 12.97 2.104 2.091 .873 .956 Total by composite. Mar. 7-8: Acid 7 a.m. to 1 p.m. . . . 608.9 1 . 0062 605 Acid 3.36 .549 .243 1 p.m. 7 p.m. . . . 821.7 1.0047 818 Acid 3.39 .798 .212 7 p.m. 11 p.m. . . . 184.3 1.0129 182 Acid 1.68 > . > • .111 11 p.m. 7 a.m. . . . Total 384.9 1.0100 381 Acid 3.20 .639 .214 1999.8 1999.8 1.0070 1986 1986 11.63 11.63 2.416 .780 .865 Total by composite. Mar. 8-9 : 7 a.m. to 1 p.m. . . . 314.9 1.0147 310 Acid 3.19 .549 .169 1 p.m. 7 p.m. . . . 572.2 1.0071 568 3.13 .665 .148 7p.m. 11 p.m. . . . 177.9 1.0124 176 1.55 .427 .095 11 p.m. 7 a.m. . . . Total 372.5 1.0095 369 3.00 .550 .237 1437.5 1437.5 l.oioo 1423 1423 10.87 10.89 2.191 2.287 .649 .712 Total by composite. Mar. 9-10: 7 a.m. to 1 p.m. . . . 453.1 1.0104 448 3.26 .561 .157 1 p.m. 7 p.m. . . . 455.1 1.0080 452 2.82 .578 .158 7 p.m. 11 p.m. . . . 260.6 1 . 0085 259 1.73 .397 .091 11 p.m. 7 a.m. . . . Total by composite. 455.6 1 .0072 452 2.93 .499 .244 1624.4 1624.4 1.0081 1611 1611 10.74 10.74 2.035 2.062 .650 .733 Metabolism Experiment No. 75. 177 Table 123. — Determinations in urine per period and per day — Continued. Date and period. (a) Amount (b) Specific gravity. (c) Volume (a -*•&). (d) Reaction. (e) Nitro- gen. (/) Phos- phoric acid by titra- tion (P0O5). (fir) Cre- atinine ex- creted (pre- formed) 1905. Mar. 10-11: 7 a.m. to 1 p.m 1 p.m. 7 p.m. . . . 7 p.m. 11 p.m. . . . 11 p.m. 7 a.m. . . . Total Grams. 393.4 553.2 228.1 421.8 1.0111 1.0063 1.0083 1.0082 c. c. 389 550 227 418 Acid Acid Grams. 2.98 2.83 1.52 2.80 Grams. . 522 .618 .586 Grams. .148 .175 .119 .286 1596.5 1596.5 1.0080 1584 1584 10.13 10.18 1.963 .728 .782 Total by composite. Total, for 7 days. . . 12273.1 12179 81.08 5.953 recorded in table 124. In addition to the elements and compounds determined in experiment No. 73, inorganic and ethereal sulphur, the so-called neutral sulphur, the preformed creatinine, and creatine were determined. Unfor- tunately, determinations of uric acid were impracticable. Phosphoric acid was determined both by fusion and titration and the results by both methods are recorded. The phosphorus, sulphur, and chlorine are expressed as phos- phoric acid, sulphur trioxide, and sodium chloride, respectively. A few determinations could not be made owing to the quantities or condition of the samples. Elimination of Wateb-Vapob. The elimination of water-vapor was measured in the usual manner. The changes in weight of chair, bedding, and other articles are given as usual in a note at the foot of table 125, page 179. Aside from a slight gain in weight of these articles on March 6, there was a continual loss in weight due to the evapo- ration of water in the unusually dry atmosphere inside the respiration chamber. The figures given in column a show that the amount of moisture in the cham- ber was considerably less at the end of the experiment than at the beginning. The total water of respiration and perspiration remained practically constant for the first 3 days of the experiment. On the next 2 days there was a gradual diminution. The lowest elimination was on the sixth day and this was exceeded by only 1 gram on the seventh. Cutaneous excretion of nitrogenous material. — During the 7 days of this experiment and the 3 days of the food experiment following, the subject eliminated through the skin a total of 0.537 gram of nitrogen, a daily average of 0.054 gram. 12 178 Influence of Inanition on Metabolism. Table 124. — Weight, composition, and heat of combustion of urine — Metabolism experiment No. 75. Mar. 4-5. Mar. 6-6. Mar. 6-7. Mar. (a) Weight grams. . (&) Water do (c) Solids, a— b do. . . . (d) Ash do (e) Organic matter, c—d do. . . . (/) Nitrogen do ... . (g) Carbon do. . . . (h) Hydrogen in organic matter. . . .do. . . . (i) Oxygen (by difference) in organic mat- ter, e— (f+g + h) grams.. (j) Phosphorus do ... . Phosphoric acid (P.205) : (k) By fusion grams.. (I) By titration do. . . . (m) Sulphur do. . . . Sulphur trioxide (S03): (n) Total grams . . (o) Inorganic and ethereal do ... . (p) Neutral, n — o do. . . . (q) Creatinine (preformed) do. . . . (r) Total creatinine. ... do. . . . (s) Creatine2 (preformed), r—q do. . . . (0 Chlorine do. . . . (m) Sodium chloride do. . . . (v) Heat of combustion calories. . 1508.2 1469.59 38.61 6.03 32.58 12.24 8.14 2.26 9.94 .625 1.431 1.460 .625 1.559 1.354 .205 1.212 1.237 .025 1.447 2.388 97 1885.7 1839.88 45.82 7.54 38.28 12.45 11.50 2.83 11.50 .984 2.255 2.245 .669 1.669 1.480 .189 1.061 1.294 .233 1.338 2.208 136 2221.0 2175.69 45.31 6.44 38.87 13.02 11.11 2.89 11.85 .897 2.055 2.091 .749 1.871 .956 1.407 .551 .616 1.016 138 1999.8 1953.00 46.80 7.80 39.00 11.63 12.20 2.80 12.37 1.051 2.406 2.416 .722 1.802 1.618 .184 .865 1.325 .460 .245 .405 146 Mar. 8-9. Mar. 9-10. Mar. 10-11. Total for 7 days. (a) Weight grams . (6) Water do . . . (c) Solids, a—b do. . . {d) Ash do . . . (e) Organic matter, c—d do. . . (/) Nitrogen do . . . (g) Carbon do . . . (h) Hydrogen in organic matter. . . .do. . . (i) Oxygen (by difference) in organic mat- ter, e— {f+g + h) grams. (J) Phosphorus do. . . Phosphoric acid (P205) : (k) By fusion grams. (I) By titration do. . . (m) Sulphur do . . . Sulphur trioxide (SO;j) : (n) Total grams. (o) Inorganic and ethereal do. . . (p) Neutral, n—o do... (>30* a.m. . 67 Mar. 12 7''30™a.m. . 68 Mar 13 7>'30ma.m.. 77 8 00 a.m. . 67 8 00 a.m. . 68 8 15 a.m. . 78 10 00 a.m. . 77 10 00 a.m. . 291 10 00 a.m. . 87 12 00 a.m. . 75 12 00 a.m. . 69 12 00 p.m. . 86 2 00 p.m. . 80 2 00 p.m.. 69 2 00 p.m. . 73 4 00 p.m. . 81 4 00 p.m. . 72 4 00 p.m. . 81 6 00 p m. . 64 6 00 p.m.. 76 6 00 p.m. . 76 8 00 p.m. . 75 8 00 p.m. . 77 8 00 p.m. . 72 10 00 p.m. . 70 10 00 p.m. . 76 10 00 p.m. . 70 1 Pulse taken while sitting. 2 Very fast and strong. Movements of subject, duration 3 days, from Mar. 11, 7 a. m., to Mar. 14, 7 a. m., 1905. March 11. A.M. 7h 00m rise, open curtain. 7 02 urinate, move chair. 7 04 c fold bed, dress, 7 09 \ weigh self, etc. 7 11 food aperture. 7 15 food aperture, drink. 7 18 comb hair, tele- phone. 7 25 drink. 7 30 count pulse. 7 35 drink. 7 38 rise, food aperture, sit. 7 44 read. 8 00 count pulse. 8 20 food aperture. 8 22 read, drink. 8 29 food aperture, read. 9 02 move about, sit. 9 08 food aperture. 9 12 food aperture. 9 20 eat, read. 9 44 stand, blood sam- pled. 9 54 sit, read. 9 55 rinse bottle. 10 00 count pulse. 10 04 food aperture. 10 06 read. 11 08 food aperture. 11 10 move about. 11 16 eat, read. 12 00 count pulse. p. M. 12" 0211 12 30 12 1 55 02 food aperture, sit. finish eating, food aperture, drink, move about, food aperture. p. M. lb06m urinate. 1 08 sit. 1 23 food aperture. 1 36 food aperture. 1 38 eat. 1 42 read. 2 00 count pulse. 2 14 food aperture. 2 16 move about. 2 18 sit. 2 32 head on table. 2 36 asleep. 3 48 awake. 3 52 drink. 3 54 write. 4 00 count pulse. 4 02 move about, uri- nate. 4 04 sit. 4 06 asleep. 4 14 awake. 4 32 asleep. 4 40 awake. 5 01 drink. 5 28 stand, telephone. 5 30 move about, sit. 5 32 kneel, doctor count pulse. 5 34 coat off, blood pres- sure test. 5 48 coat on, sit, read. 6 00 count pulse. 6 16 rise, food aperture. 6 24 rise, food aperture. 6 54 rise, food aperture, urinate. 7 02 telephone. 7 15 rise, food aperture. 7 35 finish eating, write. 7 36 food aperture. 8 00 count pulse. 8 02 rise, gymnastics. 8 12 food aperture. p. M. 8h18r 9 9 9 9 06 08 12 14 9 28 9 30 9 40 9 50 10 00 10 12 11 00 11 04 9 04 9 10 9 12 9 14 9 24 drink. food aperture. stand, sit, read. rise, food aperture. close curtain, at- tempt to defecate. open curtain, move about. adjust thermome- ter. telephone. drink. count pulse. rise, lower table, open bed, recline, read. rise, urinate, un- dress. close curtain. March 12. rise, open curtain, 'urinate, move chair, weigh self, etc. k raise table, sit. food aperture. comb hair. drink. count pulse, write. drink. count pulse. read. drink. move about. stand, blood sam- pled. move about, uri- nate. sit. food aperture. eat, read. 13 194 Influence of Inanition on Metabolism. Movements of subject.— Continued. March 12 (cont). A.M. 9" 45"' finish eating, write. 9 50 drink. 10 00 count pulse. 10 04 food aperture. 10 06 sit. 10 12 read. 10 22 move about. 10 24 sit. 10 26 read. 11 00 drink. 11 08 food aperture. 11 22 move about. 11 24 food aperture. 11 26 read. 11 40 drink. 11 42 food aperture. 12 00 count pulse. p. M. 12h 25™ drink P. M. 9h 10m drink. 10 00 count pulse. 10 05 drink. 10 12 rise, open bed, re- cline, read. 11 00 rise, undress, uri- nate, close cur- tain, retire. March 13. p. M. POS" 1 12 1 54 1 1 1 1 1 1 1 2 2 2 2 3 3 4 4 4 4 5 02 04 08 12 14 15 42 00 08 10 30 04 45 04 08 26 28 04 A. M. 7h 00m 02 \ 08 r 14 16 25 30 32 00 30 2 45 move about, telephone, food ap- erture, urinate, lean on table, read, move about, food aperture, sit. drink, read. count pulse, move about, eat, read. finish eating, write food aperture, food aperture, count pulse, read, eat. read. move about, uri- nate, sit, read, food aperture. 1 8 \ stand, blood pres- 22 ( sure test. 26 sit, read. 50 drink. count pulse, food aperture. 02 rise, food aperture. 16 rise, food aperture. 20 eat. 35 finish eating. 40 food aperture. 52 food aperture. 58 telephone. 8 00 count pulse. 8 05 food aperture. 9 00 drink. 06 15 5 5 5 5 5 5 6 00 6 12 7 7 7 7 7 7 7 7 36 7 40 8 08 8 10 8 14 8 15 8 16 8 20 8 30 9 04 9 05 9 10 9 16 9 18 9 20 9 35 9 38 9 42 10 00 10 02 10 05 10 28 10 38 11 00 11 04 11 10 11 12 11 35 12 00 rise, open curtain, urinate, weigh self, etc. raise table, food aperture, sit. drink. count pulse, remove thermome- ter, close curtain, defecate, open curtain. sit, write, food aperture, count pulse, read, food aperture. drink. stand, blood sam- pled. drink. sit. food aperture. food aperture. sit, rest. finish eating. food aperture. read. count pulse. food aperture. drink. move about. head on table, asleep. awake, food aper- ture. move about. sit, read. food aperture, drink, count pulse. p. M. 12h 02m move about, nate, sit. 12 06 read. 12 25 drink. 1 00 urinate. 1 01 food aperture. uri- 02 04 08 12 14 36 00 05 06 08 16 20 28 38 40 02 \ 06 I 08 10 16 20 24 28 3 3 3 4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 6 00 6 04 12 16 6 20 6 22 7 04 7 06 7 20 7 50 7 7 8 8 8 56 58 00 07 20 8 45 9 50 10 00 10 14 11 00 food aperture. sit, drink. move about, food aperture, count pulse, finish eating, drink. food aperture, stand, move about, food aperture, sit. read. move around, read, count pulse, drink. move about, sit, read, move chair, write, telephone, move about, write, move about, blood pressure test, telephone, sit. count pulse, telephone, move about, sit. count pulse, rise, move about, sit, read, food aperture, wave arms, telephone, urinate, food aper- ture, telephone, finish eating, rise, sit. food aperture, move about, stand, food aperture, telephone, count pulse, food aperture, drink, drink, drink. count pulse, rise, lower table, open bed, recline, read, rise, urinate, un- dress, close cur- tain. Metabolism Experiment No. 76. 195 Water and Oxygen Consumed, and Urine, Carbon Dioxide, and Water-Vapor Eliminated. The data showing the amounts of water drunk, the determinations in the urine per period and per day, the elimination of water-vapor, and the oxygen intake and output of carbon dioxide are given in tables 131 to 138. As in experiment No. 75, the water drunk during each period was measured. Table 134. — Record of water consumed — Metabolism experiment No. 76. Date. 7 to 9 a. m. 9 to 11 a. m. 11 a. m. to 1 p.m. lto3 p. m. 3 to 6 p. m. 5 to 7 p. m. 7 to 9 p. m. 9 to 11 Total p. m. for day. 1905. Mar. 11-12 Mar. 12-13 Mar. 13-14 Grams. 220.0 273.0 105.2 Grams. 75.0 73.0 130.0 Grams. 93.0 245.4 155.0 Grams. 144.4 70.0 Grams. 144.3 193.0 Grams. 150.0 244.2 170.5 Grams. 50.0 148.5 Grams. 144.4 207.9 98.3 Grams. 1021.1 1043.5 1070.5 Table 135. -Determinations in urine per period and per day- experiment No. 76. -Metabolism Date and period. (a) Amount. CM Specific gravity. (c) Vol- ume (a-s-6). (d) Reaction. (e) Nitro- gen. u Influence of Inanition on Metabolism. Table 154.— Record of carbon dioxide and oxygen — Metabolis m experim ent No. 77. Date. Period. Carbon dioxide. Oxygen. 1 (a) Amount in | chamber at end of period. (b) Total weight exhaled by subject. (c) Amount in chamber at end of period. (d) Total amount consumed by subject. S 1905. Apr. 8 Preliminary: 1 a. m Grams. 20.4 19.4 Grams. 41.9 Liters. 959.3 956.4 Grows. 37!6 1 a. m. to 3 a. m. . 3 a. m. 5 a. m. . 18.1 42.3 956.9 37.1 Apr. 8-9 5 a. m. 7 a. m. . Total 20.2 45.3 970.3 36.3 . . . • 129.5 110.4 7 a. m. to 9 a. m. . 26.9 65.6 960.5 60.9 9 a. m. 11 a. m. . 27.3 56.4 963.5 51.6 11 a. m. 1 p. m. . 21.6 48.6 968.6 45.9 1 p. m. 3 p. m. . 22.2 52.9 966.9 49.0 3 p.m. 5 p. m. . 23.2 53.6 974.0 47.2 5 p. m. 7 p. m. . 19.9 46.1 973.7 45.7 7 p. m. 9 p.m.. 20.7 50.6 984.0 45.2 9 p. m. 11 p. m. . 23.8 51.9 978.6 52.8 11 p. m. 1 a. m. . 16.6 38.4 985.8 32.5 1 a. m. 3 a. m. . 18.3 39.5 983.0 36.4 3 a. m. 5 a. m. . 19.3 48.2 983.1 45.2 Apr. 9-10 . . . 5 a. m. 7 a. m. . Total 22.9 47.7 984.7 43.6 599.5 556.0 7 a. m. to 9 a. m. . 25.6 65.9 974.4 68.8 9 a. m. 11 a. m. . 23.1 43.6 981.8 40.0 11 a. m. 1 p. m. . 17.4 38.4 982.1 41.3 1 p. m. 3 p.m.. 24.5 54.6 975.1 51.8 3 p. m. 5 p. m. . 25.2 45.4 970.8 52.5 5 p. m. 7 p.m.. 25.1 48.4 976.2 50.5 7 p. m. 9 p. m. . 22.1 52.8 983.3 50.9 9 p. m. 11 p. m. . 27.7 62.5 969.5 61.3 11 p. m. la. m. . 16.1 37.6 977.7 27.5 1 a. m. 3 a. m. . 21.3 42.1 971.5 48.8 3 a. m. 5 a. m. . 19.7 42.3 977.0 33.9 Apr. 10-11... 5 a. m. 7 a. m. . Total 21.8 43.3 968.8 44.2 .... 576.9 .... 571.5 7 a. m. to 9 a. m. . 27.0 63.1 947.5 70.0 9 a. m. 11 a. m. . 25.2 47.5 950.0 43.1 11a. m. 1 p. m. . 18.7 40.9 952.8 30.2 1 p. m. 3 p. m. . 21.0 50.2 936.6 39.1 3 p. m. 5 p.m.. 23.9 50.1 927.0 49.3 5 p. m. 7 p. m. . 18.7 41.8 931.0 40.2 7 p. m. 9 p. m . . 20.9 48.3 931.3 46.4 9 p. m. 11 p. m. . 20.4 47.0 931.2 46.3 11 p. m. 1 a. m. . 20.5 40.1 920.2 35.7 1 a. m. 3 a. m. . 18.3 42.5 926.1 41.6 3 a. m. 5 a. m. . 21.5 41.6 921.3 41.4 5 a. m. 7 a. m . . Total 21.0 43.5 915.0 47.4 .... 556.6 530 . 7 Metabolism Experiment No. 77. 215 Table 154. — Record of carbon dioxide and oxygen — Continued. Date. Period. Carbon dioxide. Oxygen. (a) Amount in (b) Total (fi) Amount in (d) Total chamber weight chamber amount at end of exhaled at end of consumed period. by subject. period. by subject. 1905. Orams. Orams. Liters. Orams. Apr. 11-12... 7 a. m. to 9 a. m. . 27.2 52.8 914.9 44.9 9 a. m. 11 a. m. . 20.0 41.2 924.9 38.7 11 a. m. 1 p. m. . 24.6 49.4 917.3 48.7 1 p. m. 3 p. m. . 18.5 40.5 921.2 37.2 3 p. m. 5 p. m. . 24.0 45.9 914.7 45.9 5 p. m. 7 p. m. . 21.8 48.8 912.2 51.5 7 p. m. 9 p. m. . 17.0 42.8 920.4 36.7 9 p. m. 11 p. m. . 19.7 52.1 913.5 55.3 11 p. m. 1 a. m. . 17.8 43.4 921.2 38.7 1 a. m. 3 a. m. . 18.6 40.2 913.0 41.4 3 a. m. 5 a. m. . 20.5 45.1 915.0 50.7 5 a. m. Total . . 7 a. m. . 19.8 42.5 923.6 41.9 .... 544.7 531.6 Elements Katabolized in the Body. The elements katabolized in the body, obtained by the methods previously described are given in table 155. Table 155. — Elements katabolized in body — Metabolism experiment No. 77. (a) Total weight. (b) Nitro- gen. (c) Carbon. (d) Hydro- gen. (e) Oxygen. Ash. First day, Apr. 8, 1905. Outgo : Water in urine Orams. 556.02 Grams. Grams. Grams. Grams. 556.02 Grams. 2528.15 42.15 669.61 599.46 8'81 7\97 163! 49 282.90 2.06 74.93 2245.25 9.43 594.68 435.97 13^88 Solids in urine Water of respiration ' Carbon dioxide Total 3839.37 3283 . 35 8.81 8.81 171.46 171.46 359.89 359.89 3285.33 2729.31 13.88 13.88 Second day, Apr. 9, 1905. Income: Oxygen from air Outgo: Water in urine 571.55 .... .... .... 571.55 2122.94 54.66 637 . 73 576.90 10^78 13^94 157^34 237 . 56 3.05 71.36 1885.38 14.91 566.37 419.56 li!98 Solids in urine Water of respiration l Carbon dioxide Total 3392 . 23 10.78 10.78 171.28 171.28 311.97 311.97 2886.22 2314.67 11.98 11.98 Loss 2820 . 68 1 Includes also water of perspiration. 216 Influence of Inanition on Metabolism. Table 155. — Elements katabolized in body- Continued. (a) Total weight. (6) Nitro- gen. (c) Carbon. (d) Hydro- gen. (e) Oxygen. (J) Ash. Third day, Apr. 10, 1905. Outgo: Water in urine Grams. 530.72 Grams. Grams. Grams. 1 • • • • | Grams. 530 . 72 Grams. 1520.30 54.00 617.52 556.62 10. 98 13. 70 ni.so 170.12 2.99 69.10 1350.18 15.78 548.42 404.82 10^55 Solids in urine Carbon dioxide Total 2748.44 2217.72 10.98 10.98 165.50 242.21 165.50 1 242.21 2319.20 1788.48 10.55 10.55 Fourth day, Apr. 11, 1905. Outgo: Water in urine 531.59 .... .... .... 531.59 1102.01 54.59 626.56 544.74 li!45 u.hi 148! 56 123.31 3.24 70.11 978.70 16.19 556.45 396.18 9!i4 Solids in urine Carbon dioxide Total 2327.90 1796.31 11.45 11.45 163.13 163.13 196.66 196.66 1947.52 1415.93 9.14 9.14 Loss 1 Includes also water of perspiration. Elements and materials katabolized in the body. — A summary of the elements katabolized and the compounds computed from them is given in table 156. The high carbohydrate katabolism on the third and fourth days is of especial interest as compared with results in other fasting experiments. Table 156. — Elements and materials katabolized in body — Metabolism experiment No. 77. Date. (a) Nitro- gen. (6) Carbon. (c) Hydro- gen. (d) Oxy- gen. (e) Water. if) Protein. (a) Fat. (h) Carbo- hydrates (asjily- cogeu). (i) Ash. 1905. Apr. 8- 9 Apr. 9-10 ... Apr. 10-11 Apr. 11-12 Total, 4 days. Grams. 8.81 10.78 10.98 11.45 Grams. 171.46 171.28 165 . 50 163.13 Grams. 359 . 89 311.97 242.21 196.66 Grams. 2729.31 2314.67 1788.48 1415.93 Grams. 2989.75 2558.28 1945.73 1540.46 Grams. 52.86 64.68 65.88 68.70 Grams. 134.97 171.88 137.65 149.85 Grams. 92.70 14.85 58.92 29.21 Grams. 13.88 11.98 10.55 9.14 42.02 671.37 1110.73 8248.39 9034.22 252.12 594.35 195.68 45.55 Balance of water. — In table 157 the distribution of intake and output of water is shown. Allowance has not been made for water in feces passed on the first and second days of the experiment. Metabolism Experiment No. 77. 217 Table 157. — Distribution of intake and outgo of water — Metabolism experiment No. 77. Date. Outgo from the body. Balance of preformed water. Water of oxida- tion of organic hydro- gen (c-d). (a) Water of urine. CM Water of respira- tion and perspira- tion. (c) Total (a+ft). (d) Pre- formed (katabo- lized) water in outgo.1 (6) Intake in drink. (/) Loss of pre- formed water (rl-e).1 1905. Apr. 8- 9 Apr. 9-10 Orams. 2528.2 2122.9 1520.3 1102.0 Orams. 669.6 637.7 617.5 626.6 Orams. 3197.8 2760.6 2137.8 1728.6 Orams. 2989.8 2558 . 3 1945.7 1540.4 Orams. 2048.2 1592.9 1018.7 962.0 Orams. 941.6 965.4 927.0 578.4 Orams. 208.0 202.3 192.1 188.2 Apr. 10-11 Apr 11-12 Total for 4 days Average per day . . 7273.4 1818.3 2551.4 637.9 9824.8 2456.2 9034 . 2 2258.6 5621.8 1405.5 3412.4 853.1 790.6 197.6 1 No allowance has been made for water In feces passed on the first and second days. Changes in Body-Weight Compared with Balance of Income and Outgo. The balance of income and outgo is compared with the fluctuations in body-weight and as in the other fasting experiments the feces passed during the experiment have not been included in obtaining the outgo. The com- parisons of the measured and computed losses show unsatisfactory agreement on the first 2 days. Table 158. — Comparison of changes of body weight with balance of income and outgo — Metabolism experiment No. 77. Apr. 8-9. Apr. 9-10. Apr. 10-11. Apr. 11-12. Total for 4 days. Average per day. Income : (a) Water consumed (6) Oxygen (c) Total (a + b) Outgo : (d) Urinei (e) Feces2 (/) Carbon dioxide (g) Water of respiration and per- spiration (h) Total (d+f + g) (i) Loss of body material (c— h). . . . ( j) Loss of body weight Grams. 2048.30 556.02 2604.22 2099.30 144.40 599.46 669.61 3368.37 764.15 736 Orams. 1592.90 571.55 2164.45 2205.20 35 . 30 576.90 637.73 3419.83 1255.38 1190 Orams. 1018.70 530.72 1549.42 1879.10 556.62 617.52 3053 . 24 1503.82 1502 Orams. 962.00 531.59 1493.59 1176.70 544.74 626.56 2348.00 854.41 868 Orams. 5,621.80 2,189.88 7,811.68 7,360.30 179.70 2,277.72 Orams. 1405.45 547.47 1952.92 1840.08 44.93 569.43 2,551.42 637.85 12,189.44 3047.36 4,377. 76ll094. 44 4,296 1074 1 The data in this line should not be confounded with urine data in other tables, explanation, p. 66. 2 Not included in the total outgo. See p. 120. See Output of Heat. The total heat production, shown in column d of table 159, was fairly constant throughout the 4 days of this experiment, the difference between the maximum and minimum amounts being but 67 calories. 218 Influence of Inanition on Metabolism. Table 159. — Summary of calorimetric measurements and total heat production — Metabolism experiment No. 77. (a) (Z>) (c) id) Date. Period. Heat meas- ured in terms 0*20 • Heat used in vaporiza- tion of water. Sum of heat correc- tions.1 Total heat produc- tion (tt+b+e). 1905. Preliminary : Calories. Calories. Calories. Calories. Apr. 8 1 a.m. to 3 a.m. . . 77.2 46.2 2 + 7.2 2130.6 3 a.m. 5 a.m. . . 76.2 45.9 2 +12.0 2 134.1 Apr. 8-9 5 a.m. 7 a.m. . . Total 90.0 39.4 2-20.4 2109.0 243.4 131.5 * — 1.2 2373.7 7 a.m. to 9 a.m. . . 180.5 45.1 — 30.0 195.6 '.) a.m. 11 a. m. . . 147.3 37.0 + 1.4 185.7 11 a.m. 1 p.m. . . 130.1 34.7 - 5.9 158.9 1 p.m. 3 p.m. . . 125 . 7 35.4 — .6 160.5 3 p.m. 5 p.m. . . 136.1 33.6 — .5 169.2 5 p.m. 7 p.m. . . 138.5 33.3 — .2 171.6 7 p.m. 9 p.m. . . 129.1 33.0 — 12.6 149.5 9 p.m. 11 p.m. . . 135.3 28.1 -19.8 143.6 11 p.m. 1 a.m. . . 63.6 32.3 + 15.8 111.7 1 a.m. 3 a.m. . . 78.3 29.5 + 12.2 120.0 3 a.m. 5 a.m. . . 98.2 35.3 + 23.7 157.2 Apr. 9-10 5 a.m. 7 a.m. . . Total 106.6 34.0 + 10.3 150.9 1469.3 411.3 — 6.2 1874.4 7 a.m. to 9 a.m. . . 187.6 48.0 — 40.2 195.4 9 a.m. 11 a.m. . . 136.4 30.6 — 13.6 153.4 11 a.m. 1 p.m. . . 101.6 31.3 + 8.3 141.2 1 p.m. 3 p.m. . . 142.1 31.6 - 8.1 165.6 3 p.m. 5 p.m. . . 143.2 29.9 + 9.9 183.0 5 p.m. 7 p.m. . . 135.6 30.5 — 2.7 163.4 7 p.m. 9 p.m. . . 117.1 34.4 -11.9 139.6 9p.m. 11 p.m. . . 158.9 36.4 -35.0 160.3 11 p.m. 1 a.m. . . 81.0 33.3 + 15.9 130.2 1 a.m. 3 a.m. . . 94.8 32.0 + 9.8 136.6 3 a.m. 5 a.m. . . 105.9 32.3 + 22.7 160.9 Apr. 10-11 5 a.m. 7 a.m. . . Total 102.9 30.3 + 16.8 150.0 1507.1 400.6 — 28.1 1879.6 7 a.m. to 9 a.m. . . 192.6 42.9 — 48.0 187.5 9 a.m. 11 a.m. . . 142.7 30.5 - 6.1 167.1 1 1 a.m. 1 p.m. . . 85.9 32.2 — 1.1 117.0 1 p.m. 3 p.m. . . 130.6 34.8 + .7 166.1 3 p.m. 5p.m... 144.0 31. 4 + 4.8 180.2 5 p.m. 7 p.m. . . 99.5 32.1 + 1.4 133.0 7 p.m. 9 p.m. . . 122.8 32.8 -13.2 142.4 9 p.m. 11 p.m. . . 129.4 33.3 -18.8 143.9 11 p.m. 1 a.m. . . 84.2 28.3 + 19.8 132.3 1 a.m. 3 a.m. . . 98.7 33.6 + 6.3 138.6 3 a.m. 5 a.m. . . 121.6 29.1 + 16.3 167.0 5 a.m. 7 a.m. . . Total 100.5 33.7 + 30.7 164.9 1452.5 394.7 - 7.2 1840.0 See pp. 42-49. 2 Not corrected for changes in body temperature and weight. Metabolism Experiment No. 77. 219 Table 159. — Summary of calorimetric measurements and total heat production- Continued. (a) (&) (c) (d) Date. Period. Heat meas- ured in terms C20. Heat used in vaporiza- tion of water. Sum of heat correc- tions.1 Total heat produc- tion (a+b+c). 1905. Apr. 11-12 7 a.m. to 9 a.m. . . Calories. 166.9 Calories. 31.2 Calories. — 52.8 Calories. 145.3 9 a.m. 11a.m... 108.4 31.9 - 5.6 134.7 11 a.m. 1 p.m. . . 139.1 30.5 + 2.1 171.7 1 p.m. 3 p.m. . . 112.3 31.5 - 2.7 141.1 3 p.m. 5 p.m. . . 123.0 29.3 + 16.6 168.9 5 p.m. 7 p.m. . . 135.2 36.6 - 8.1 163.7 7 p.m. 9 p.m. . , 123.6 31.5 -10.6 144.5 9 p.m. 11 p.m. . . 132.1 35.3 — 19.1 148.3 11 p.m. 1 a.m. . . 81.0 32.8 + 20.9 134.7 1 a.m. 3 a.m. . . 85.5 33.8 + 15.1 134.4 3 a.m. 5 a.m. . . 115.2 31.6 + 10.6 157.4 5 a.m. 7 a.m. . . Total 120.5 32.2 + 10.0 162.7 1442.8 388.2 -23.6 1807.4 1 See pp. 42-49. Balance of Energy. A comparison of the energy derived from different sources with the total heat production for the 4 days of the experiment is given in table 160. The agreement is not very satisfactory save on the first day, but owing to the compensating discrepancies the average difference for the whole experiment is but — 0.8 per cent. Table 160. — Comparison of energy derived from katabolized body material with total heat production — Metabolism experiment No. 77. Date. Energy derived from different sources. From body protein. (a) Energy of protein katabo- lized. (&) Poten- tial energy of urine. (c) Net energy (a-b) . (d) From body fat. (e) From body glyco- gen. Total (c + d+e), (a) Total heat pro- duced. Energy from body material grenter (+) or less (— ) than output. (h) Amount (/-y). («) Pro- portion (h~g). 1905. Apr. 8- 9. . Apr. 9-10. . Apr. 10-11. . Apr. 11-12.. Total, 4 days Av. per day. Cals. 299 365 372 388 Cals. 90 157 157 170 1424 356 574 143 Cals. Cals. 209 1288 208 1640 215 1313 218 1430 850 5671 213 1418 Cals. 388 62 247 122 Cals. 1885 1910 1775 1770 819 205 7340 1835 Cals. 1874 1880 1840 1807 Cals. + 11 + SO — 65 — 37 7401 1850 61 15 Perct. + 0.6 + 1.6 — 3.5 —2.0 -0.8 '."><> Influence of Inanition on Metabolism. Table 161. — Oxygen and carbon dioxide thermal quotients and respiratory quotients — Metabolism experiment No. 77. Date and period. (a) Total heat produc- tion. (b) Oxygen con- sumed. (c) Oxygen thermal quotient (100b -»- a). (d) Carbon dioxide elimi- nated. (e) Carbon dioxide thermal quotient (100d-«-a). (f) Volume of carbor dioxide elimi- nated. (dxO.6091 (g) Volumt of oxygen con- sumed (bX0.7). (h) i Respi- ratory quo- tient 1905. Apr. 8: Preliminary : 1 a.m. to 3 a.m. Cals. '1130.6 Grams. 37.0 28.3 Grams. 41.9 32.1 Liters. 21.3 Liters. 25.9 0.82 3 a.m. 5 a.m. 1134.1 37.1 27.7 42.3 31.5 21.6 25.9 .83 5 a.m. 7 a.m. 1109.0 36.3 33.3 45.3 41.6 23.0 25.4 .91 1 1373.7 110.4 29.5 129.5 34.7 65.9 77.2 0.86 Apr. 8-9: 7 a.m. to 9 a.m. 195.6 60.9 31.1 65.6 33.5 33.4 42.6 0.78 9 a.m. 11 a.m. 185.7 51.6 27.8 56.4 30.4 28.7 36.1 .80 11 a.m. 1 p.m. 158.9 45.9 28.9 48.6 30.6 24.8 32.1 .77 1p.m. 3 p.m. 160.5 49.0 30.5 52.9 32.9 26.9 34.3 .79 3 p.m. 5 p.m. 169.2 47.2 27.9 53.6 31.7 27.3 33.1 .83 5 p.m. 7 p.m. 171.6 45.7 26.6 46.1 26.8 23.4 32.0 .73 7 p.m. 9 p.m. 149.5 45.2 30.2 50.6 33.8 25.7 31.6 .81 9p.m. 11 p.m.1 143.6 52.8 36.8 52.0 36.2 26.5 37.0 .72 11 p.m. 1 a.m. 111.7 32.5 29.1 38.4 34.4 19.6 22.7 .86 1 a.m. 3 a.m. 120.0 36.3 30.3 39.5 32.9 20.1 25.5 .79 3 a.m. 5 a.m. 157.2 45.2 28.8 48.2 30.7 24.5 31.6 .78 5 a.m. 7 a.m. 150.9 43.7 29.0 47.6 31.6 24.3 30.6 .79 Total 1874.4 556.0 29.7 599.5 32.0 305.2 389.2 0.78 Apr. 9-10: 7 a.m. to 9 a.m. 195.4 68.8 35.2 65.9 33.7 33.6 48.2 0.70 9 a.m. 11 a.m. 153.4 40.0 26.1 43.6 28.4 22.2 28.0 .79 11 a.m. 1p.m. 141.2 41.3 29.3 38.4 27.2 19.6 28.9 .68 1p.m. 3 p.m. 165.6 51.8 31.3 54.6 33.0 27.8 36.3 .77 3 p.m. 5p.m.' 183.0 52.5 28.7 45.4 24.8 23.1 36.8 .63 5p.m. 7p.m. 163.4 50.5 30.9 48.4 29.6 24.7 35.4 .70 7p.m. 9p.m. 139.6 50.9 36.4 52.8 37.8 26.9 35.6 .71 9p.m. 11 p.m. 160.3 61.3 38.2 62.5 39.0 31.8 42.9 .74 11 p.m. 1 a.m. 130.2 27.5 21.1 37.6 28.9 19.1 19.3 .99 1 a.m. 3 a.m. 136.6 48.8 35.7 42.1 30.8 21.4 34.1 .63 3 a.m. 5 a.m. 160.9 33.9 21.1 42.3 26.3 21.5 23.7 .91 5 a.m. 7 a.m. 150.0 44.2 29.5 43.3 28.9 22.0 30.9 .71 Total 1879.6 571.5 30.4 576.9 30.7 293.7 400.1 0.73 Apr. 10-11: 7 a.m. to 9 a.m.1 187.5 70.0 37.3 63.1 33.7 32.1 49.0 0.66 9 a.m. 11 a.m. 167.1 43.1 25.8 47.5 28.4 24.2 30.1 .80 11a.m. 1p.m. 117.0 30.2 25.8 40.9 34.9 20.8 21.1 .98 1p.m. 3 p.m. 166.1 39.1 23.6 50.2 30.2 25.5 27.4 .93 3 p.m. 5 p.m. 180.2 49.8 27.4 50.1 27.8 25.5 34.5 .74 5p.m. 7p.m. 133.0 40.2 30.3 41.8 31.4 21.8 28.2 .76 7 p.m. 9 p.m. 142.4 46.4 32.6 48.3 33 . 9 24.6 32.5 .76 9 p.m. 11 p.m. 143.9 46.3 32.2 47.0 32.6 23.9 32.4 .74 11 p.m. 1 a.m. 132.3 35.7 27.0 40.1 30.3 20.4 25.0 .82 1 a.m. 3 a.m. 138.6 41.6 30.0 42.5 30.7 21.7 29.1 .74 3 a.m. 5 a.m. 167.0 41.4 24.8 41.6 24.9 21.2 29.0 .73 5 a.m. 7 a.m. Total 164.9 47.4 28.7 43.5 26.4 22.2 33.2 .67 1840.0 530.7 28.8 556.6 30.3 283.4 371.5 0.76 1 See table 159, p. 21S. Metabolism Experiment No. 77. 221 Table 161. — Oxygen and carbon dioxide thermal quotients and respiratory quotients — Continued. Date and period. (a) Total heat produc- tion. (b) Oxygen con- sumed. (c) Oxygen thermal quotient (1006 -s- a). id) Carbon dioxide elimi- nated. (e) Carbon dioxide thermal quotient (100il+ a). •> Influence of Inanition on Metabolism. A Series of Two-day Fasting Experiments with Seven Subjects. Metabolism Experiments Nos. 79-83, 85, and 89. The fasting experiments thus far given in this report furnished data for studying the metabolism during inanition of but 3 persons. It seemed im- portant therefore to make studies with a large number of men, in order to verify some of the deductions rendered possible by the previous experiments. Particularly was it desired to ascertain the length of time required by different individuals for the transition from katabolism with food to fasting katabolism, i. e., the duration of the transitional period. More information was also sought concerning the effect of inanition on the pulse and observations were made of the influence of fasting on the respiration, strength, and the onset of fatigue. Accordingly, 7 fasting experiments of 2 days' duration were made with different subjects during the autumn and winter of 1905-1906, the results of the series being given below. Following experiments Nos. 83, 85, and 89, 24-hour food experiments were made, the data of which are not included in this publication. The 7 men who served as subjects in this series of 2-day fasts were students in Wesleyan University, young men in good health. Two of them had prev- iously been in the calorimeter for short periods of about 24 hours, but the others were inexperienced. It seemed best, therefore, to accustom the men to the environment of the calorimeter and hence each subject entered the chamber approximately 12 hours before the experiment proper began. Measurements of subjects. — In order to give some idea of the physique of the subjects and their muscular development, the measurements of each man, as recorded in the university gymnasium, are shown below. The measurements of H. E. S., A. H. M., and H. C. K., were taken at times very near to that of their respective experiments. The others were measured somewhat earlier, 8 months previous in the case of H. R. D., and the considerable period of about 3 years and 2 months in the case of N". M. P. D. W. was a football and baseball player and had unquestionably developed to an extent beyond that shown by the measurements here given. Notes from diaries. — Each subject kept a diary during his stay in the calori- meter. The diaries have been abstracted, and such notes as relate to the physical and mental condition are shown below. Metabolism Experiments Nos. 79-83, 85, and 89. 223 Measurements of subjects — Metabolism experiments Nos. 79-83, 85, and 89. I. H. E. S. C. It. Y. A. H. M. H.C.K. H.R.D.1 N.M.P. D. W Date -j Age at date Weight kilograms. . Height centimeters.. Srernum do Navel do Pubis do Sitting do Knees do Length : Shoulders to elbow. centim'rs. Elbow to tips do Arm reach do Right foot do Left foot do Girth: Head centimeters. . Neck do Upper chest: Repose centimeters.. Depressed do Inflated do Lower chest : Repose centimeters. . Depressed do Inflated do Waist do Hips do Right biceps do Flexed do Left biceps do Flexed do Right arm do Left arm do Right elbow do Left elbow do Right forearm do Contracted do Left forearm do Contracted do Right wrist do Left wrist do Right thiifh do Left th igh do Right knee do Left knee do Right calf do Left calf do Right instep do Left instep do Breadth : Head centimeters.. Neck do Shoulders do Chest do Waist do Hips do Depth : Chest centimeters.. Abdomen do Oct. 20, 1905. 19' 56 177 33.8 81 78.8 84.9 71.1 66 76 68.7 2L5 26.8 23.5 26 25.8 2L3 47.5 46.1 33 31 39.4 25.7 21.6 32.6 16.8 17.3 Dec. 9, 1904. y. 1Qm. 68.5 159.8 is 36.7 95 86.5 100.5 80.5 79 87.8 75 26^8 30.5 26 28.5 27.8 27^5 57.5 56 36.5 35.5 38.7 28.6 26.3 33.5 18.7 18.3 Oct. 23, 1905. 24y' 62.8 178.6 34.5 85 81.1 89.5 73.2 71.5 80.6 64.5 26^5 28.5 26 28.5 27 53 63 35 34.5 38.8 26.4 21.7 32 17.7 16.6 Oct. 12. I90.r>. :.'l ■ 4m- 72 181.2 35.5 87 86.1 88.9 77.5 76.3 82 73.9 27. 31. 27. 30 28 28' 57.5 64.5 36.5 35.9 42.6 28.3 24.2 32! 3 19.3 18.9 Apr. 10, 1905. 17y" 55.9 171.2 141.4 100.5 88.7 90.7 42.6 36.7 48.5 177.8 26.1 26.2 57.5 35 83.5 87\5 82.5 68 87.5 27 2^5 24 24 25 •ii" i7 " 17 48 48 35.5 35.6 31.5 31.5 22.5 22.6 15.5 10 39.2 23^2 31.2 19.2 17 Oct. 18, 1902. is- 10'" 64.8 177.1 146.6 105.7 88.8 93.5 46.2 38.3 48.1 183 26.6 26.6 67.5 34.1 84 82 88.5 80 86 70.5 92.5 27.3 26.6 22^5 22.3 20.6 20.4 24.3 23^4 15^6 15.5 52.2 53 38.5 38.5 35 35.5 23.4 23.6 15.6 11.2 40.5 26.6 25.4 35 18.6 18.3 Oct. 23, 1903. 19y. nm. 74.3 179.7 147.6 106.5 89.8 95.7 44.8 39.2 46.9 181.2 27.6 28.4 56.5 36.2 93 87.5 97 77.4 86.2 77.9 101.2 32.2 31 27! 5 28.1 23 24.3 28.5 27'. 4 if. 9 17.8 6H.3 5' 39 39 36 36 26 26 15.5 11.6 43.6 28.6 27.2 35 '.4 18.2 17.6 1 Depth of head, 19.9 centimeters. 224 Influence of Inanition on Metabolism. October 13, 1905: 7 a.m. Rose; stomach seems somewhat empty. 7b40m a. m. Was getting very hungry but after I took a drink of water the feeling was not so noticeable. ll"35m a. m. Am getting rather hungry and my belt hangs very loosely about me. lh15m p. m. Do not feel hungry now. 4h14m p. m. All the afternoon I have had a queer feeling on my lungs every time I drew a breath longer than Notes from diaries. Experiment No. 79, H. E. S. usual. Am not especially hungry but feel as if I could eat. Lay down from 5h10m to 7*'2m p. m. October 14, 1905: 7 a. m. Am very sick at my stomach ex- cept when lying down. 7h2m p. m. Have felt weak and tired all day, and have been dizzy when I stood up. llh2m p. m. Went to bed; got up at some time in the night, about 1 a. m. I believe. Thought it was morning. Experiment No October 26, 1905 (preliminary night) : Entered calorimeter chamber about 8 p. m. Attempted to read but was so sleepy that I had to stop; had slept only 5 hours the night before. Re- tired at 11 p. m. October 27, 1905: 7 a.m. Arose at signal; followed the usual morning routine. Sat down and read. 7"45m a. m. Took strength test. Feel all right this morning; not hungry at all. 10"30m a. m. Began to read, sitting near window. 12h20m p. m. Lay down on bed; slept some of the time. llh20m p. m. Drank about one-half glass of cold water before retiring, though not very thirsty. 80, C. R. Y. October 28, 1905: 7 a. m. Do not feel ill, but have bad taste in mouth. Am a little weaker than yesterday, but not especially hungry. 7h20m a. m. Drank some water, which I vomited some time between 8 and 9 a. m. llh20m a. m. Reclined on bed, sleeping part of the time. 6 p. m. Have been lying down most of the afternoon. Felt all right except for bad taste in mouth. 8 p.m. Went to sleep about 8 or 8h30,n p. m. and didn't wake up until 11 p. m. llh20m p. m. Drank some cold water; it tasted good. During the latter part of the night I slept but very little; was anxious for morning to come. Had bad taste in mouth whenever I was awake. Experiment No. 81, A. H. M. November 21, 1905: 7h30ma. m. Slept all through the night; well rested. Awoke at about 6"15m a. m., lay awake until 7. Feel per- fectly normal this morning; have good appetite. Felt somewhat thirsty, drank a little water. lh10m p. m. Studied and read almost all of the morning. Feel very well. Could eat a pretty good dinner now, but fasting does not trouble me very much. 7t'5m p. m. Feel just the same as at noon; just about as hungry. Feel perfectly strong. Read and studied during the afternoon. November 22, 1905: 7h45m a. m. Slept well through the whole night; woke at about 6h15m a. m., rose at 7 a. m. Don't feel quite as strong this morning. Was thirsty when I got up; drank some water and feel better, though somewhat weak when standing. Am all right when sitting down. Head perfectly clear. 1 p. m. Feel stronger now than when I rose this morning, began to feel bet- ter after I had been about an hour and a half. Think perhaps the water made me feel better. Spent nearly all the morning writing an essay. Read a little. 71,10m p. m. Feel all right, not very hun- gry now. Lay down on bed from 1 p. m. to 6; had a few naps. Prob- ably asleep about half the time. Have been reading since 6. llh2m p. m. Have been lying down all the evening, reading a part of the time. Feel well. Metabolism Experiments Nos. 79-83, 85, and 89. 225 November 23, 1905: 7 a.m. Rested well during night; woke up twice, once about 3 a. m. and Notes from diary. — Continued. about 5 a. m. but went to sleep soon each time. Feel well, but a little thirsty. Experiment No. 82, H. C. K. November 23, 1905 (preliminary night) : 11p.m. Made up bed and retired; slept pretty well but woke up rather early. Had a pain in my back which I think was due to lying in one po- sition for so long a time. Also had a pain in my stomach which was more noticeable whenever I took a deep breath. November 24, 1905: 7 a. m. Arose. Mouth and throat were very dry. 7h50m a. m. Taking a deep breath causes a pain in my throat; otherwise I feel well though my mouth is dry. 9h20m a. m. Wrote letters until 2"45m p. m. lh15m p. m. Drank some water. 7h25,n p. m. Slept a little while. Have been lying down and now feel much better. 7h30m p. m. Studied until 10h15m. 10h30m p. m. Studied until 11 p. m. llh2m p. m. Retired. November 25, 1905: 7h2m a. m. This night passed better than the last, though I awoke early. Backache and stomach-ache were not so bad as the night before. Very thirsty. Felt rather warm, on aris- ing, almost like a fever. 7h30m a. m. Defecated. 7h50m a. m. Drank fresh water; feel much better. Feverish feeling has left me and I feel quite comfortable. 1 p. m. Drank some water. Slightly dizzy when I stood up. 3h50m p. m. Lay down. 7h5m p. m. Got up and drank some water. Have a slight headache, also pain in the stomach; otherwise am all right. 7h30m p. m. Feel fairly comfortable. Ly- ing on the respiration apparatus has made my chest and back rather sore. The rectal thermometer does not trouble me. llh5m p. m. Made up bed and retired. Experiment No. 83, H. B. D. December 4, 1905 (preliminary night) : 9h15m p. m. Wrote till 10h25m. 11 p. m. Retired; did not get to sleep for about half an hour, then slept very soundly. Awoke and looked at my watch at 4 a. m.; slept on till about 6"30m, then lay awake till the signal to rise was given. December 5, 1905: 7"30m a. m. Feel very comfortable and contented. Stomach somewhat emp- ty, but not uncomfortable and have no particular desire for food. Feel strong; most of my movements are slow, however, for fear of entang- ling the apparatus attached to me. 7b40m a. m. Read for about 5 minutes, then wrote a short letter, but took considerable time about it as I was in doubt as to what to say. Am be- ginning to notice everything that passes. Am much entertained with watching the people who pass the window. 15 8h15m a. m. Some water was passed in. I drank it all. No strange taste. 9 a. m. Lay down till 11, sleeping a great part of the time. llh10m a. m. Rose, and drank a bottle of water, which tasted very good. Feel perfectly well, and am not hungry. Took dynamometer test. lh10m p. m. Lay down, feeling perfectly well; went to sleep. 3h30m p. m. Awoke, but did not get up. 4h10m p. m. Got up. Feel well, but less ambitious and lively than when I lay down. Slightly unpleasant feel- ing in my head, more like dizziness than headache, but not exactly either; not serious. 41,15m p. m. Received a bottle of water, of which I drank about a third. Feel chilly at times and decidedly unpleasant ever since I got up. 4h30m p. m. Lay down again; more com- fortable when lying down. Have a slight feeling of nausea; hungry. Went to sleep with blanket over me. 226 Influence of Inanition on Metabolism. Notes from diary. — Continued. December 5, 1905 (cont.) 7h10m p. m. Drank some water; it was absolutely tasteless, but cool and re- freshing. Am greatly refreshed by my sleep. Feel perfectly well again, head perfectly clear; less inclined to move rapidly than I was this morn- ing, but am quite as comfortable. Not hungry now except when I think of something in particular to eat. 9h25m p. m. Feel well and comfortable; no pain, nothing to annoy me. 9h30m p. m. Lay down; slept most of time till signal at 11 p. m. 11 p. m. Undressed and retired. It must have been nearly 12 before I fell asleep, then slept soundly. Awoke at 5 a. m., looked at my watch, and went to sleep again. December 6, 1905: 7 a. m. Feel rather faint and weak this morning, but have no pain and am quite comfortable, but hungry. 8h5m a. m. Very unpleasant taste in the mouth. Drank some water and went on writing. 9 a. m. Lay down, after drinking some water; went to sleep almost at once. 10"30m a. m. Awoke, but lay still. 11a.m. Arose; took dynamometer test. 12 a. m. Feel much better than when I arose this morning; that is, feel stronger. Still feel somewhat faint but very comfortable. lp.m. Lay down; feel well. 2h50m p. m. Awoke. 4h10m p. m. Got up. Feel well; a little weak, but no more so than this morning. Quite comfortable. 7 p. m. Feel well and rather stronger than earlier in the day, but some- what faint even yet. 7h 10m p. m. Took dynamometer test. 7h45m p. m. Am not in pain but am de- cidedly faint. It requires very no- ticeable effort to hold up my head, also to write. Left hand weak and rather unsteady, right hand some- what better. Not at all pleasant sit- ting up. 8 p.m. Saw friends at the window; greatly pleased to see them. Made me forget all about how I felt. 9 p. m. Lay down, feeling comfortable, though a little faint and hungry; went to sleep. 11 p. m. Awoke at signal. Retired and went to sleep very shortly. Awoke early, could not tell when, but prob- ably about 5h30m a. m., then went to sleep again. Experiment No. 85, N. M. P. December 8, 1905 (preliminary night) : Read until 9h45m p. m.; light not good. Very comfortable; stomach feels full. December 9, 1905: Did not sleep very well. Woke at lh30m, 3h30m, and 4h30m a. m., staying awake about three-quarters of an hour the last time. The reason for waking was that I kicked off the loose blanket and became chilly. Rose at 7. Drank some water at 7h40m. Slightly cool, but not un- comfortable. 9h15m a. m. Took dynamometer test and drank a little water. 9h30m a. m. Feel somewhat hungry. 121,15m to 12"20m p. m. Walked around the calorimeter and examined the apparatus. 12h20m p. m. Began to study mathemat- ics. lh25m p. m. Stopped studying. Made up bed and lay down, read and rested; slept a little. 4 p. m. Dynamometer test. Everything very comfortable; not feeling hun- gry except when I think about it. Read and rested until 7. 7 p. m. Studied, read, and wrote all the evening. Part of the time in chair and part of the time reclining on bed. 11 p. m. Am not feeling uncomfortable in any way nor particularly hungry. December 10, 1905: Slept better than the previous night; woke up twice, at 2h15m, and 4h30m a. m. or thereabout. The pneumo- graph makes me uncomfortable when I lie down. 7h50m a. m. Felt dizzy when I got up; my stomach seemed somewhat up- set, and I felt weak. Drank some water. After sitting for a few min- utes the dizziness ceased; the bil- ious feeling is also going. Desire to urinate noticeable as it was all day yesterday. Metabolism Experiments Nos. 79-83, 85, and 89. 227 Notes from diary. — Continued. December 10, 1905 (cont.) 8 a. m. Moved around. The water I drink tastes bad. Wrote letters from 8 to 9h45m a. m. 9h45m a. m. Lay down and read. Am feeling considerably better than when I first got up; water tastes better. lPlS"1 a. m. Have been reclining most of the time; began to sit at table and read. 12h15m p. m. Feel chilly, read and wrote at table until nearly 2h30m. 2h30m p. m. Lay down and rested and read until 7; slept a little. 7h15m p. m. Sat and read and wrote. Went to food aperture twice and telephoned some during the evening. My gums feel soft and a little sore; have a peculiar sensation in my ab- domen. Don't feel exactly hungry, but feel empty. My chest is un- comfortable when I lie on a hard part of the pneumograph; feel tired particularly about the muscles in my lower side and back which comes, I suppose, from sitting up. Slight desire to defecate. Lay down about 10h30m but stayed up until then reading and writing. 11 p. m. Went to bed. December 11, 1905: Had great difficulty in getting to sleep last night; probably a little ner- vous. Was hot and sweaty and the pneumograph troubled me until 1 a. m. Woke up repeatedly and dozed again, then slept until 2 a. m., was awake a half hour. Slept until 4, but did not stay awake long; slept until 6h15m. 7 a. m. Rose and went through the usual routine of weighing. Do not feel dizzy, or sick, or uncomfortable in any way. Experiment No. S9, D. W. January 10, 1906: Got into bed at llhllm last night. Slept well until about 4 a. m. and dozed from that time until 7 a. m. 7 a. m. Feeling well except a little hazy, possibly due to loss of sleep. Lay down at 9h10m and had a very com- fortable nap, which lasted until 12" 30m p. m. Still feel a little sleepy, but do not feel especially hungry as yet. 2u45m p. m. Have not felt hungry as yet. Lay down at 3h5m and had a slight nap; got up at 5h30m. Have not felt very hungry nor have I felt uncom- fortable in any way; do not feel thirsty. Lay down at 8h15m until 9h15m. Feel a little hungry for the first time. Re- tired at llh5m. January 11, 1906: Woke up at 3h45m a. m. not feeling very well, and quite thirsty; took a large drink of water, which did not taste very good. Lay down again and dozed until 7 o'clock. Do not feel quite as lively as I did yesterday morning; somehow my stomach feels a litle queer and I feel tired in general. Drank a little water. 9h10m a. m. Was glad to lie down and had a short nap, waking at 11. Feel a little hungry now but water tastes bad, so that I don't care much about eating. Sat and read until 3h5m p. m. when I lay down. Am feeling rather hungry now. At 3h5m I lay down and slept until 5. 8h5m p. m. Felt a little weak after the strength test but otherwise am feel- ing very well. The test seemed a little harder than last night. At 10h23m I had some fresh water passed in; it tasted much better than any of the water I had drunk before. Retired at 11 p. m. but did not sleep very much. Tossed around consid- erably. v.'s Influence of Inanition on Metabolism. Pulse and respiration. — In all experiments previously included in this publi- cation, each subject counted his own pulse and the respiration was not taken. The following records of pulse and respiration were obtained by means of the Fitz pneumograph, the use of which in these researches has already been explained. (See page 10.) Pui.se and Respiration. Metabolism experiment No. 79. Time. Pulse per minute. Respiration ] per minute. 1 Time. Pulse per minute. Respiration per minute. Time. Pulse per minute. Respiration per minute. Oct. 12 Oct. 13 Oct. 14 P. M. P. M. P. M. llh38m 13 8h56m 19 12h08m 19 Oct. 13 9 30 58 17 2 15 67 20 A. M. 10 26 19 3 40 19 12M2m 12 11 50 60 17 5 45 78 18 12 46 11 11 53 66 18 8 35 . . 20 1 23 54 11 11 58 17 9 12 60 18 2 46 14 Oct. 14 9 38 . . 16 3 30 11 A. M. 10 00 59 16 4 14 10 12h50m 18 11 30 63 19 4 46 11 1 25 63 18 11 54 . , 17 5 20 10 2 00 17 Oct. 15 9 46 55 3 10 19 A. M. 11 00 55 13 3 52 19 12h 18m 18 P. M. 4 54 17 1 36 20 lh48m 63 17 5 44 17 2 36 16 3 50 58 19 5 45 62 19 2 40 16 7 14 19 6 26 17 3 30 20 8 12 • • 18 6 30 57 18 6 06 6 45 17 18 Metabolism Experiments Nos. 79-83, 85, and 89. 229 Pulse and Respiration — Continued. Metabolism experiment No. 80. Pulse Respiration Pulse Respiration Pulse Respiration Time. per per Time. per per Time. per per minute. minute. minute. minute. minute. minute. Oct. 26 Oct. 27 Oct. 2S P. M. P. M P. M. llb45m 73 16 71.35m 62 llh30m 74 Oct. 27 9 00 52 13 11 44 is A. M. 10 05 47 14 11 52 76 17 12u20m 14 10 35 . . 14 Oct. 29 12 50 14 11 45 59 18 A. M. 12 59 73 13 Oct. 28 12h06'n 77 17 1 24 14 A. M. 12 20 75 16 1 54 16 12"25m 59 14 12 23 79 1 58 67 20 12 55 63 13 12 24 75 2 16 59 13 1 25 65 17 12 37 77 i6 2 26 . . 14 2 00 58 14 12 48 81 20 2 52 13 2 32 58 15 12 50 86 3 02 58 14 3 00 63 20 12 59 75 15 3 28 14 3 40 60 14 1 21 72 15 3 52 14 4 23 17 1 34 71 16 3 59 62 13 4 32 78 19 1 40 73 15 4 26 15 5 00 65 15 1 55 91 19 4 53 50 13 5 30 68 15 2 11 72 17 5 10 , , 13 6 10 68 14 2 24 77 16 5 28 14 6 40 70 14 2 37 71 16 6 00 r t 13 8 10 87 19 2 50 71 15 6 08 52 13 8 13 97 . . 3 00 73 16 6 15 68 14 8 42 90 17 3 17 81 21 6 54 14 9 56 64 16 3 36 73 15 8 45 17 10 27 80 16 3 53 97(?) 15 9 28 16 11 41 64 14 4 08 81 21 10 49 16 P. M. 4 15 86 11 32 17 12h 10"' 67 15 4 24 84 19 P. M. 3 33 . . 20 4 36 89 18 12h 17m 14 3 41 87 . , 4 49 73 16 1 17 20 4 35 73 16 4 59 74 17 2 10 13 5 27 82 22 5 18 75 14 2 47 12 6 22 72 24 5 22 73 16 3 23 13 9 14 87 14 5 34 71 15 3 32 56 9 48 63 14 5 38 75 . . 4 23 17 10 28 66 15 5 50 76 16 5 01 19 10 57 76 19 6 02 . , 17 5 45 56 11 25 20 6 06 80 6 15 17 11 28 85 20 6 07 75 . . 7 27 16 230 Influence of Inanition on Metabolism. Pulse and Respiration — Continued. Metabolism experiment No. 81. Pulse Respiration Pulse i Respiration ! Pulse Respiration Time. per per Time. per per Time. per per minute. minute. minute. minute. minute. minute. Nov. 20 Nov. 21 Nov. 22 P. M. P. M. P. M. llh27m 43 14 11b37m 40 13 3h44m 48 17 11 43 42 13 11 52 41 14 4 00 44 14 11 58 14 Nov. 22 4 18 47 17 Nov.21 A. M. 4 32 48 17 A. M. 12" 09m 41 13 4 58 52 18 12"04m 42 12 23 41 13 5 14 48 17 12 17 42 13 12 39 39 15 5 50 50 21 12 26 42 14 12 55 40 14 6 41 45 22 12 39 43 13 1 12 39 14 7 12 53 12 55 45 15 1 28 39 14 9 41 50 19 1 13 41 12 1 43 39 14 10 20 48 19 1 26 41 13 1 56 41 14 10 47 51 20 1 41 40 13 2 01 39 11 24 47 16 1 58 41 13 2 12 43 14 11 39 47 15 2 10 41 13 2 25 . . 15 11 55 48 15 2 24 40 12 2 31 40 12 Nov. 23 2 39 41 12 2 40 39 14 A. M. 2 55 45 13 2 54 40 14 12hllm 49 16 3 10 46 19 2 59 13 12 24 49 15 3 24 40 11 3 14 42 14 12 45 53 16 3 39 42 12 3 28 41 14 12 55 49 15 3 55 42 12 3 45 41 13 1 00 . , 15 4 12 42 12 3 58 43 14 1 09 15 4 34 42 15 4 13 42 13 1 15 48 15 4 45 41 14 4 28 42 13 1 27 49 15 4 55 44 13 4 42 42 15 1 40 48 15 5 15 44 14 4 55 42 15 1 53 . . 16 5 29 13 5 12 45 13 1 56 15 5 35 42 13 5 28 48 17 1 59 47 15 5 49 42 13 5 32 , , 13 2 09 . . 15 6 04 13 5 42 13 2 14 47 15 6 16 45 17 5 52 44 14 2 20 47 15 6 28 42 6 01 45 2 28 48 . . 6 37 43 12 6 13 45 is 2 40 55 18 6 49 43 12 6 27 46 15 2 54 49 17 8 37 47 16 6 41 46 17 3 09 46 15 9 00 18 6 56 47 14 3 24 48 15 9 37 44 17 8 17 55 17 3 39 48 15 10 12 47 11 8 42 59 15 3 54 50 16 P. M. 8 57 64 t m 4 08 . . 17 12u 12m 46 9 26 60 4 11 15 1 38 43 20 9 46 17 4 14 53 21 2 04 39 9 58 6i 4 27 49 16 2 16 48 i3 10 19 57 20 4 31 51 13 2 48 59 11 10 34 58 4 41 . . 13 4 02 44 13 10 57 53 20 4 45 49 14 4 26 42 11 11 20 54 4 55 50 14 5 01 45 12 11 41 55 16 5 12 51 18 5 28 39 12 11 56 50 17 5 22 15 6 00 47 P. M. 5 29 49 . . 9 00 44 l"06m 46 14 5 41 52 15 9 38 is 2 05 53 14 5 56 53 14 9 48 45 2 20 47 6 10 53 15 10 06 42 a 2 30 41 14 6 25 56 17 10 36 . . 15 2 46 40 14 6 29 . . 16 10 43 56 t t 3 08 43 15 6 38 17 10 44 46 3 20 44 18 6 50 55 19 11 24 38 13 3 27 44 •• 6 57 58 Metabolism Experiments Nos. 79-83, 85, and 89. 231 Pulse and Respiration — Continued. Metabolism experiment No. 82. Pulse Respiration Pulse Respiration Pulse Respiration Time. per per Time. per per Time. per per minute. minute. minute. minute. minute. minute. Nov. 23 Nov. 2^ Nov. 25 P. M. A. M. A. M. 6u59m 59 23 5h25m 53 16 lh39m 50 17 7 13 64 25 5 39 54 17 2 02 50 18 7 26 66 22 5 57 53 18 2 13 55 21 7 47 67 20 6 08 . , 15 2 33 51 18 7 58 60 21 6 25 56 17 2 45 52 19 8 13 22 6 29 , , 16 2 56 54 19 8 17 23 6 38 , t 17 3 18 65 18 8 23 65 21 6 43 51 16 3 25 57 20 8 30 61 6 55 55 15 3 41 56 18 8 43 73 22 8 14 64 23 3 55 55 18 8 57 68 20 8 27 52 19 4 10 56 19 9 10 64 18 8 41 69 20 4 26 55 19 9 26 61 23 8 57 61 23 4 38 56 19 9 31 58 9 20 70 24 4 56 55 19 9 48 53 ie 9 53 68 24 5 12 56 19 10 00 52 17 10 08 59 23 5 27 63 19 10 15 . , 17 10 18 72 23 5 38 17 10 26 59 21 10 30 63 25 5 45 57 10 34 58 20 10 54 72 23 5 54 55 17 10 45 59 21 11 12 69 23 6 09 58 18 10 56 58 22 11 52 72 24 6 23 59 19 11 16 49 20 P. M. 6 37 18 11 29 48 16 ^"og1" 68 23 6 42 59 16 11 45 48 16 1 52 65 6 53 19 11 54 . . 17 2 20 73 28 6 57 ei 18 Nov. 24 2 46 65 24 8 09 73 24 A. M. 3 03 67 23 8 24 55 26 12h07m 48 , t 3 21 68 26 8 44 78 27 12 19 49 16 3 35 59 19 9 01 73 24 12 30 . , 17 3 58 63 19 9 23 67 23 12 33 , . 16 4 09 58 19 9 39 66 23 12 38 51 16 4 29 52 21 9 55 69 25 12 46 51 16 4 43 52 18 10 07 60 23 12 56 51 16 5 01 63 17 10 15 58 21 1 10 50 16 7 54 64 10 35 66 23 1 24 48 15 9 35 2i 10 43 68 22 1 28 17 9 42 69 23 10 57 68 24 1 39 50 17 9 47 75 11 24 79 26 1 55 15 9 59 2i 11 38 68 25 2 04 52 17 10 07 64 11 44 66 25 2 17 57 16 10 22 67 22 11 54 71 24 2 27 51 16 10 39 57 23 P. M. 2 42 52 17 10 43 22 12h01m 63 25 2 56 52 17 10 58 59 21 12 25 23 3 10 52 17 11 21 53 21 12 47 72 22 3 25 53 17 11 37 52 19 2 01 67 23 3 29 54 11 52 53 19 2 07 67 23 3 38 17 Nov. 25 2 16 63 22 3 44 58 18 A. M. 2 24 64 20 3 55 56 16 12"07m 53 18 2 47 67 27 4 17 54 17 12 22 54 18 3 01 74 25 4 32 59 16 12 38 15 3 12 74 26 4 36 54 12 42 56 3 24 80 26 4 44 57 17 12 53 52 17 3 39 83 25 4 56 52 17 1 13 49 18 4 11 86 20 5 12 53 17 -: 1 24 49 19 4 19 68 24 232 Influence of Inanition on Metabolism. Pulse and Respibation — Continued. Metabolism experiment No. 82 — Continued. Time. Pulse per minute. Respiration per minute. Time. Pulse per minute. Respiration per minute. Time. Pulse per minute. Respiration per minute. Nov. 25 Nov. 25 Nov. 26 P. M. P. M. A. M. 4>'30m 62 20 llh30m 76 3h49m 59 4 37 67 19 11 40 60 20 3 57 62 20 4 45 64 20 11 55 60 20 4 15 62 19 5 08 61 20 Nov. 26 4 25 59 18 6 11 67 23 A. M. 4 40 68 17 6 26 22 12h09m 60 19 4 56 78 20 6 31 81 12 25 60 19 4 59 84 6 44 70 23 12 41 67 16 5 00 74 7 00 72 12 43 63 5 09 20 7 15 80 26 12 47 58 5 14 71 8 35 70 12 55 56 18 5 16 79 9 40 85 28 12 59 , 18 5 17 67 , B 9 43 76 1 10 59 19 5 25 73 21 9 53 25 1 27 56 20 5 28 66 9 57 si 24 1 41 57 18 5 39 19 10 08 27 1 55 60 19 5 43 69 10 10 84 25 2 17 64 20 5 54 u m 21 10 13 88 2 25 19 6 00 83 10 23 27 2 29 60 6 01 77 t g 10 29 77 2 39 60 19 6 14 69 20 10 31 86 m 2 56 62 19 6 25 70 18 10 41 25 3 04 63 19 6 40 83 23 10 44 71 . , 3 13 64 # t 6 43 75 . . 10 54 23 3 14 68 6 44 70 . . 10 58 82 23 3 27 62 20 6 57 73 21 11 27 63 21 3 40 • • 18 Metabolism Experiments Nos. 79-83, 85, and 89. 233 Pulse and Respiration — Continued. Metabolisvi experiment No. 83. Pulse Respiration Pulse Respiration Pulse Respiration Time. per per Time. per per Time. per per minute. minute. minute. minute. minute. minute. Dec. 4 Dec. 5 Dec. 6 P. M. P. M. P. M. 10h 19m 69 21 6h36m 61 18 lh31m 77 18 10 44 59 19 6 49 74 17 1 40 56 10 54 68 7 04 . , 18 1 53 54 ie 11 27 80 19 7 20 82 2 00 71 11 43 77 20 7 43 79 is 2 13 73 17 Dec. 5 8 12 85 18 2 32 77 18 A. M. 8 39 76 18 2 49 75 18 12b04ra 67 18 9 05 56 17 3 13 80 21 12 20 72 20 9 15 83 3 24 99 19 12 39 64 18 9 23 19 3 33 92 12 49 65 17 9 43 74 18 3 46 89 21 1 07 65 17 10 04 19 3 56 20 1 28 65 16 10 11 78 4 16 87 21 2 05 63 16 10 32 70 i<3 4 30 81 21 2 24 18 10 48 74 18 4 47 83 2 54 63 17 11 17 76 20 5 06 83 is 3 11 63 16 11 31 , t 18 5 24 20 3 35 63 16 11 52 , . 17 5 39 82 18 3 53 66 17 Dec. 6 6 00 81 17 4 34 65 17 A. M. 6 12 80 4 55 65 12h01m 59 6 25 73 16 5 40 67 ie 12 32 17 6 30 72 5 53 64 18 12 57 73 16 6 44 69 is 6 04 . . 18 1 25 63 16 7 31 71 21 6 17 63 17 1 52 63 16 7 55 66 17 6 39 17 2 10 66 15 8 15 81 22 7 30 . . 21 2 37 63 16 8 35 71 21 8 23 . . 19 2 53 67 17 9 07 19 8 42 56 19 3 16 72 18 9 38 75 18 9 04 , . 22 3 52 69 17 9 49 76 19 9 26 84 16 4 19 58 17 10 04 69 17 9 52 21 4 35 69 17 10 28 62 16 10 11 72 21 5 05 66 16 10 57 63 15 10 36 85 . t 5 27 70 17 11 15 75 18 10 43 17 6 09 71 18 11 48 75 19 10 53 70 18 6 39 72 18 Dec. 7 11 17 75 23 6 48 71 17 A. M. 11 48 . . 21 6 54 76 19 12h04m 73 16 P. M. 7 34 94 21 12 19 70 17 12b27m 80 18 7 54 81 19 12 58 17 12 38 82 22 8 15 19 1 03 66 1 07 88 26 8 24 94 1 24 61 15 1 44 74 17 8 37 87 i9 1 59 68 17 2 05 60 17 8 53 91 18 2 19 64 16 2 27 . , 19 9 17 81 18 2 50 63 16 2 42 76 16 9 30 17 3 04 65 16 3 06 62 9 44 72 , . 3 19 59 14 3 18 18 9 55 75 16 3 42 69 18 3 27 73 10 07 17 4 02 89 23 3 38 , , 19 10 24 88 22 4 20 58 15 3 56 , , 18 10 42 83 19 4 31 89 16 4 15 , m 21 10 52 87 , 4 51 73 18 4 32 95 . , 11 31 18 5 03 74 18 4 42 88 19 11 44 87 , , 5 19 72 18 5 10 78 16 P. M. 5 32 72 19 5 23 , . 16 12h 10m . . 21 5 47 74 18 5 33 81 17 12 20 87 , , 6 23 72 20 6 10 73 17 12 33 82 23 6 40 77 20 6 18 78 12 42 . . 20 6 55 75 19 6 27 .. 18 ■>:) 1 Influence of Inanition on Metabolism. Pulse and Respibation — Continued. Metabolism experiment No. 85. I ' «fi| Pulse Respiration Pulse Respiration Pulse Respiration Time. per per Time. per per Time. per per minute. minute. minute. minute. minute. minute. Dec. 8 Dec. 9 Dec. 10 P. M. P. M. A. M. 8h38m 62 13 5b32m 46 , # llh46m 67 14 8 48 63 13 5 47 57 14 P. M. 9 04 62 , t 5 56 . , 12 12h07m . m 12 9 20 64 6 14 59 12 16 13 9 56 , . U 6 23 . t i4 12 31 58 , . 10 13 53 6 44 12 12 39 71 , , 10 45 48 12 7 27 62 12 43 13 11 15 . . 13 7 47 ii 12 54 67 13 11 52 . . 12 7 54 61 . . 2 06 67 . . Dec. 9 8 34 12 2 19 m t 14 A. M. 8 49 63 2 27 14 12h07m 57 8 54 12 2 35 51 . m 12 23 . . 14 9 23 48 14 2 45 14 1 21 . . 11 9 37 54 14 3 17 59 14 1 50 46 10 03 13 3 29 59 13 2 12 ii 10 22 51 3 45 63 13 2 29 45 12 10 31 i2 4 37 62 . . 3 23 , , 11 10 43 59 5 29 69 13 4 08 11 11 39 ii 5 38 13 4 23 56 11 43 11 6 07 69 , . 5 08 i2 11 51 . . 11 6 15 71 5 22 44 Dec. 10 6 32 is 5 31 . . i2 A. M. 7 07 13 6 09 60 12 12"32m , , 12 7 15 75 13 6 26 57 13 12 45 66 7 30 94 15 6 48 54 11 1 15 . , i2 7 50 79 . . 8 24 71 13 1 24 56 7 59 16 8 34 62 1 43 i2 8 13 77 14 8 46 t . i2 2 17 50 11 8 37 68 17 8 53 59 11 2 39 12 9 16 73 13 9 32 68 13 2 54 67 11 9 33 80 16 9 48 67 15 3 05 69 14 10 14 14 10 07 62 16 3 29 57 11 10 30 65 16 10 23 57 3 50 66 10 45 65 10 43 . . 12 4 16 52 12 11 11 73 is 10 50 61 13 4 34 13 11 30 15 11 15 13 4 55 12 11 45 73 . . 11 29 51 12 5 03 64 12 Dec. 11 11 50 10 5 26 69 A. M. P. M. 5 51 i4 12h02m 64 . , 12h08m 64 6 10 63 13 12 23 63 12 29 56 i2 6 31 69 13 12 45 ii 12 38 61 10 7 30 77 1 22 , . 14 12 48 . 12 8 23 i4 1 41 74 16 12 59 65 8 32 83 14 1 55 15 1 34 54 i3 8 43 78 2 03 56 16 1 50 54 9 11 75 2 50 79 2 07 64 9 30 18 2 59 i3 2 20 ii 9 43 13 3 11 80 15 2 50 si 10 20 60 13 3 56 83 16 3 37 56 i2 10 28 61 4 40 82 14 4 10 52 10 39 i3 5 11 73 14 4 37 55 i2 10 45 73 5 36 73 16 4 46 58 12 10 55 i3 6 17 79 15 5 24 9 11 27 70 6 41 84 14 Metabolism Experiments Nos. 79-83, 85, and 89. 235 Pulse and Respiration — Continued. Metabolism experiment No. 89. Pulse Respiration Pulse Respiration Pulse Respiration Time. per per Time. per per Time. per per minute. minute. minute. minute. minute. minute. Jan. 9 Jan. 10 Jan. 11 P. M. P. M. P. M. 10h51m , , 19 10h46m 67 3h14m 59 11 34 50 , , 11 02 46 3 28 ie> 11 51 , , 17 11 18 i9 3 44 59 17 Jan. 10 11 30 48 p m 4 18 52 16 A. M. Jan. 11 4 39 54 15 lh14m 52 . . A. M. 4 53 16 1 34 . . 18 12" 25™ . t 15 5 50 18 2 06 51 m . 12 47 50 6 30 22 2 48 13 1 15 . . 17 6 50 19 4 19 44 . , 1 46 50 7 11 77 21 4 40 , , 11 2 24 54 1.6 7 19 75 20 5 31 . . 14 2 55 50 m 7 35 23 6 15 15 3 15 17 7 46 22 6 48 47 . . 3 34 48 8 09 21 6 59 , , 16 4 23 51 8 19 63 20 7 30 , . 17 4 34 u m 15 8 34 20 7 42 71 . . 5 38 , . 17 8 44 58 8 16 64 . . 6 15 46 8 49 62 is 8 35 . , 20 6 31 . t 14 9 08 55 19 8 54 63 . , 6 47 54 , B 9 30 20 9 49 59 17 7 27 63 9 39 53 10 09 64 16 7 38 . t i6 10 19 61 i9 10 35 13 8 08 66 10 38 24 11 07 65 8 24 69 19 11 06 22 11 44 47 . m 8 46 57 11 22 19 P. M. 9 28 51 14 Jan 12 12" 16m 48 t t 10 46 47 16 A M. 12 44 51 . . 10 54 15 12h25m m # 16 1 51 62 , . 11 11 # t 17 1 10 50 2 08 66 11 23 15 1 19 50 2 53 22 11 31 62 1 42 16 3 19 78 , m 11 53 60 3 26 16 3 49 18 P. M. 3 38 49 4 21 47 17 12h17m 17 5 13 15 6 54 , . 20 12 30 55 5 29 46 7 19 20 12 48 62 i(3 5 38 50 7 35 59 u m 2 05 67 21 6 10 i6 7 59 75 2 26 68 6 23 54 8 56 51 18 2 35 20 6 38 , . 16 9 59 55 19 2 46 e>i 6 48 49 , . 10 34 22 236 Influence of Inanition on Metabolism. Record of body movements. — The subjects of these 2-day experiments followed no very definite routine, except that the times of rising, weighing, and retiring were the same for all. Their activity varied with their tempera- ment and inclinations though in no case was there evidence of much muscular exertion, and all the experiments were essentially " rest experiments." An index of the muscular activity of the different men may be obtained from the " movements of subjects " recorded below. The men were all cautioned to attempt to secure like muscular activity on the 2 days of the experiments. In some instances this was accomplished by having the record of body movements for the first 24 hours prepared during the night and requiring that the subject duplicate this record as nearly as possible on the second day. Movements of Subjects. Experiment No. 79, H. E. S., Oct. 13, 7 a. m., to Oct. 15, 7 a. m., 1905. October 13. A. M. P. M. A. M. 11" 35m write. 5h44m telephone. r 02m rise, dress. 11 54 telephone. 7 04 rise, urinate. 7 06 } • weigh self, etc. 11 56 sit, food aperture. 7 06 telephone, sit. 7 13] 11 58 rise. 7 08 food aperture. 7 20 sit, write. p. M. 7 10 sit. 7 24 telephone. 12" 02m raise table, sit, 7 16 lie, read. 7 29 food aperture. write. 8 02 telephone. 7 40 drink, write. 12 07 urinate. 8 05 food aperture. 7 42 lie, read. 12 24 telephone, write. 9 02 rise, urinate. 7 52 move about, food 1 04 read, seal letter. 9 04 lie. aperture. 1 08 stand, telephone. 10 02 sit, write. 7 54 read. 1 12 drink. 10 08 rise, coat off. 8 28 telephone. 1 14 write. 10 10 lie. 8 32 sit, food aperture, 1 16 food aperture, 10 12 asleep. rise. read. 11 02 rise, urinate, write 8 33 urinate. 1 20 stand, read. 11 04 close curtain, un 8 34 sit, drink. 1 22 sharpen pencil. dress, retire. 8 8 36 38 write, lie, read. 1 2 26 10 read, rise. October 1%. 8 40 telephone, read. 2 12 food aperture, tele- A. M. It li C\ A hi 9 08 rise, strength test. phone. 2" 24 rise. 9 12 telephone, strength 2 14 open bed. 7 02 n -i 4 \ rise, dress, urinate test. 2 16 stand, write. 7 14 1 7 18 j weigh self, etc. 9 18 write. 2 18 lie, read. 9 20 lie, read. 2 24 sit, food aperture. 7 22 food aperture. 9 36 write. 2 25 lie, read. 7 24 telephone. 9 45 drink. 2 34 sit, food aperture. 7 25 O -t A food aperture. 9 10 52 07 telephone, read, drink. 2 2 36 38 telephone, lie. 8 14 8 28 sleep, telephone. 10 24 sit, telephone. 4 04 rise, urinate, sit, 8 30 lie. 10 26 rise, food aperture. write. 8 34 asleep. 10 28 sit, rise, sit. 4 08 telephone. 9 02 telephone. 10 30 lie, read. 4 20 comb hair. 9 04 rise. 10 36 urinate. 4 28 read. 9 06 strength test. 11 18 telephone. 5 02 write. 9 08 sit, write. 11 22 rise, food aperture. 5 04 rise, urinate. 9 14 food aperture. 11 24 sit, drink, food ap- 5 08 lie, read. 9 18 telephone. erture. 5 12 sit, food aperture. 9 24 food aperture. 11 26 rise, lie, read. 5 14 lie, read. 9 25 urinate. Metabolism Experiments Nos. 79-83, 85, and 89. 237 Movements of Subjects. — Contin ued. October H (cont.) p. M. P. M. A. M. lh 10m write. 5h08ra lie, read. 9h 28m lie, study. 1 12 drink. 5 28 stop reading. 9 52 telephone. 1 14 look at watch, lie. 5 52 read. 9 54 sit, food aperture. 1 24 telephone, lie. 6 08 stop reading. 9 58 rise, move about. 1 34 read. 7 02 rise, urinate. 10 00 lie, read. 2 20 sit. 7 06 sit, write. 10 18 telephone. 2 25 lie, read. 7 08 food aperture. 10 20 sit, food aperture. 3 02 asleep. 7 10 drink, write. 11 24 lie. 3 12 sit, food aperture. 7 18 lie. 11 36 sit, food aperture. 3 14 lie. 7 40 restless. P. M. 3 24 read. 8 06 strength test. 12h 08m sit, read. 3 46 doze. 8 10 open bed, write. 12 36 read. 3 50 count pulse. 8 12 drink, lie. 12 38 sit erect. 4 40 doze. 8 24 doze. 1 02 telephone. 5 02 sit, urinate, tele- 11 02 rise, urinate. 1 04 food aperture. phone. 11 04 undress, close cur 1 08 read. 5 06 food aperture. tain, retire. Experiment No. 80, C. R. T., Oct. 27, 7 a. m., to Oct. 29, 7 a. m., 1905. October 27. p. M. A. M. A. M. 7" 06m rise, telephone. 11" 20m move. 7h 00m rise, urinate. 7 08 urinate, lie. 11 22 arrange bed, lie. 7 7 06} 14) ■ weigh self, etc. 11 11 00 02 rise, urinate, arrange bed. 11 p. 34 M. asleep. 7 26 dress, sit. 11 04 strength test. 12" 32m asleep. 7 33 food aperture. 11 10 close curtain, un- 1 00 awake, telephone. 7 34 wipe absorbers. dress, retire. 1 02 read. 7 54 food aperture. 11 18 telephone, food ap- 1 04 urinate. 9 02 telephone. erture, drink. 1 06 sit, food aperture. 9 08 stand, strength 1 08 lie. test. October 28. 1 24 asleep. 9 06 sit, read. A. M. 2 08 awake, turn over, 9 26 telephone. 7h 00m rise, urinate. asleep. 9 10 30 08 read, telephone. 7 7 1 ft 1 weigh self, etc. 3 3 04 10 awake. restless. 10 10 rise, food aperture. 7 20 drink. 3 26 doze. 10 14 sit. 7 40 sit. 3 34 awake. 10 38 read. 8 08 write. 5 04 sit, telephone. p. M. 8 16 count pulse. 5 06 strength test, sit, 12* 20m rise, open bed, uri- 8 36 asleep. telephone. nate. 8 42 awake. 5 12 lie. 12 22 arrange bed, lie, 9 04 rise, strength test. 6 20 sit, telephone. read. 9 08 rise, food aperture. 7 02 rise, urinate. 1 04 sit, telephone. 9 10 sit. 7 06 sit, take tempera- 1 44 stop reading, re- 9 18 asleep. ture. cline. 9 48 awake. 7 08 lie. 2 02 asleep. 9 54 asleep. 8 00 asleep. 4 06 sit, telephone, food 10 02 telephone, rise, sit. 11 00 awake, rise, uri- aperture. 10 34 asleep. nate. 4 08 rise, strength test. 11 04 telephone. 11 04 strength test. 4 10 read. 11 08 rise, food aperture. 11 07 food aperture. 4 40 lie. 11 12 sit. 11 12 close curtain. 5 22 doze. 11 14 elbow on knee. 11 20 drink, retire. 238 Influence of Inanition on Metabolism. Movements of Subjects. — Continued. Experiment No. 81, A. H. M., Nov. 21, 7 a. m., to Nov. 23, 7 a. m., 1905. November 21. p. M. A M. A M. 41 26,r stand, adjust ther- 8l 24™ telephone. 7' 02m rise, telephone, mometer. 8 32 read, write. open curtain. 4 28 move about, sit. 8 40 read. 7 04 urinate, dress. 4 32 read, elbow on 9 02 telephone, rise, 7 06 telephone. knees. food aperture. 7 7 08 18 [ weigh self, etc. 5 5 02 04 stand, telephone. 9 04 sit, rise, strength test. 7 22 sit, write. 5 06 sit, read. 9 08 sit, telephone. 7 24 food aperture, sit, 5 14 rise, food aperture. 9 12 elbows on knees. drink. 5 16 sit. 9 40 telephone, adjust 7 42 read. 5 20 stand, sit. thermometer. 8 34 stop reading. 5 32 stand. 10 16 food aperture. 8 42 read. 5 34 sit, read. 10 20 write. 9 02 rise, food aperture. 6 04 read. p. M. 9 04 strength test. 7 00 telephone. 12" 08m telephone, move 9 06 telephone. 7 02 food aperture, uri- about. 9 10 sit, read. nate. 12 10 adjust thermome- 9 24 rise, food aperture. 7 06 telephone, stand, ter, sit. 9 30 stand, read. food aperture. 12 28 read. 9 40 sit, read. 7 10 sit. 12 46 rest, elbows on 11 02 rise, move about. 7 20 read. knees. 11 04 sit. 7 54 stop reading. 1 02 rise, urinate. 11 08 read. 8 10 read. 1 04 write. 11 26 turn around. 8 50 stop reading. 1 06 telephone. 11 28 read. 9 04 stand, strength 1 08 arrange bed, lie. 11 44 move chair. test. 1 44 asleep. p. M. 9 08 sit. 2 16 sit, arrange blan- 12h 08" ' read. 9 48 read. ket, lie. 1 02 telephone. 10 20 stop reading. 4 48 awake. 1 04 rise, food aperture, 10 26 read. 5 02 telephone. urinate, stand. 11 00 telephone. 5 04 read. 1 12 stand. 11 02 rise, urinate, open 6 08 sit, read. 1 1 14 24 sit. arrange clothes, sit. sit, chair tilted. 11 04 bed. close curtain, un- dress, retire. 6 6 6 12 14 16 rise, urinate, sit, read. 1 28 November 22. 7 7 02 04 rise, urinate, strength test. 2 14 asleep. A. M. 7 06 telephone. 2 16 sit. 7h 02m telephone, rise, 7 08 food aperture. 2 22 read. open curtain. 7 12 sit, read. 3 02 telephone, rise, 7 06 urinate, dress, fold bed. ► weigh self, etc. adjust chair, sit, 7 32 lie. strength test. 7 7 7 08^ 19 7 44 asleep. 3 04 telephone. 8 20 read. 3 06 sit. 20 9 02 stop reading. 3 10 rise, food aperture. telephone. 10 08 read. 3 12 sit, read. 7 26 food aperture. 10 24 telephone. 3 40 read. 7 28 stand, adjust ther- 11 02 rise, urinate. 4 02 rise, sit, wave mometer. 11 04 open bed, undress, arms. 7 30 sit. close curtain, re- 4 24 telephone. 7 45 drink, write. tire. Experiment No. 82, E r. C K., Nov. 2k, 7 a. m., to 1 Vov . 26, 7 a. m., 1905. November 2J/. A. M. A. M. A. M. 7" 23m food aperture. 7h 30m write. 71 02" 1 rise, urinate, fold 7 24 move about, adjust 7 35 adjust table. bed, dress. p n e u m o graph 8 04 sit, read. 7 7 09 17 i weigh self, etc. and thermome- ter. 8 9 42 04 stop reading, telephone. 7 22 telephone. 7 28 sit, write. 9 06 rise, food aperture. Metabolism Experiments Nos. 79-83, 85, and 89. 239 Movements of Subjects. — Continued. November 24 (cont.) A. M. 9" 08m sit. 9 12 telephone. 9 14 rise, strength test. 9 16 sit, write. 9 36 telephone. 9 38 rise, food aperture, sit. 9 40 sit. 9 42 write. 11 12 telephone, rise, sit. 11 16 write. p. M. Is 04m telephone. 1 06 sit. 1 10 rise, urinate, food aperture, drink. 1 14 telephone. 1 16 remove thermome- ter. 1 18 food aperture. 1 22 sit, write. 1 24 food aperture. 1 26 sit, adjust ther- mometer. 1 30 sit, write. 1 48 sit, read, write. 3 04 telephone, rise, food aperture, sit. 3 06 rise, food aperture, drink. 3 08 sit, telephone. 3 10 strength test. 3 14 sit. 3 16 lower table, open bed. 3 18 write. 3 20 lie. 3 26 asleep. 5 30 sit, drink, lie. 7 02 rise, telephone. 7 04 food aperture. 7 06 urinate. p. M. A. M. 7h 12m fold bed, brush 10" 26m telephone. mercury from 10 30 study. floor. 11 02 telephone, rise. 7 15 raise table. 11 04 strength test. 7 22 sit, write. 11 14 food aperture. 7 30 study. 11 20 play cards. 9 05 drink. p. M. 9 55 strength test. lh 00m telephone, urinate, 10 25 drink. drink. 10 30 sit, study. 1 02 move, sit, play 11 02 rise, u r i n a te , cards. drink. 1 28 sit. 11 04 open bed. 2 10 doze. 11 08 undress, retire. 2 2 32 34 write, play cards. November 25. 3 04 drink. A. M. 3 10 write. 7> 02"1 rise, urinate. 3 38 rest, strength test. 7 04 ) dress, weigh self, 3 40 food aperture, sit. 7 15 f etc. 6 44 write. 7 20 sit, write. 3 46 fold table, open 7 26 food aperture. bed. 7 32 close curtain, defe- 3 50 lie. cate. 3 52 sit, telephone, food 7 42 adjust thermome- aperture, drink. ter, connect res- 4 04 doze. piration tube. 7 04 rise, urinate, drink. 7 42 sit, raise table. 7 06 telephone. 7 48 food aperture. 7 12 sit, write. 7 50 drink. 7 16 telephone. 7 54 write. 7 20 food aperture, 8 00 sit. read. 8 10 food aperture. 7 30 fold bed, raise 8 20 read. table, sit, write. 8 30 write. 8 15 drink. 9 02 telephone. 8 20 play cards. 9 04 food aperture. 9 25 food aperture, 9 06 move, sit. drink. 9 10 write. 9 26 strength test. 9 14 sit. 9 28 sit, play cards. 9 48 telephone. 11 05 rise, urinate, drink. 10 02 head on arms table. on 11 06 fold table, open bed. 10 06 doze. 11 12 close curtain, re- 10 22 sit. tire. Experiment No. 83, H. R. D., Dec. 5, 7 a. m., to Dec. 7, 7 a. m., 1905. A. M. 9h 16m rise, open bed, lie. 9 44 sit, telephone. 9 46 rise, food aperture. 9 48 lie, sit, telephone. 9 50 rise, food aperture. sit. 9 56 food aperture. 9 58 lie. 10 22 doze. 11 02 sit, telephone December 5. A. M. A. M. 8h12ra rise, food aperture, 7h00m rise. drink. 7 02 open curtain, uri- 8 16 sit. nate. 8 18 write. 7 04 dress. 8 38 read. 7 06 ) fold bed, weigh 9 00 urinate. 7 22 f self, etc. 9 04 telephone. 7 28 sit, write. 9 08 rise, food aperture. 7 40 read, write. 9 13 rise, food aperture. 8 08 telephone. 9 14 sit. 240 Influence of Inanition on Metabolism. Movements of Subjects. — Continued. December 5 (cont.) A. M. llh 04m rise, strength, test. 11 06 fold bed. 11 08 urinate, drink. 11 10 raise table, sit, write. p. M. 12"05m 12 20 12 24 12 44 00 02 06 08 10 12 30 04 08 12 15 18 20 22 24 32 34 06 02 04 06 12 14 24 26 30 10 urinate. telephone. write. stop writing, sit. urinate. telephone, read. write. sit. rise, open bed, lie. asleep. awake. sit, food aperture. raise table. sit, write. urinate. sit. rise, food aperture. sit, drink. write. open bed, lie. cover self with blanket, asleep, telephone, urinate, strength test, sit, telephone, food aperture, drink, rise, move about, fold bed. sit, write, restless, food aperture. p. M. 8b 15" 9 10 9 25 9 36 11 00 11 02 11 04 A. M. 5h00r 02 04 06 09 16 18 24 26 28 32 34 38 15 02 9 04 9 08 9 10 9 12 11 02 11 04 11 12 11 20 7 7 7 7 7 7 7 7 7 7 7 7 8 9 read. drink. urinate. fold table, open bed, lie, asleep, rise. urinate, telephone, open bed, undress, retire. December 6. rise, urinate, lie, asleep, rise, urinate, fold clothes, dress, hang chair. I weigh self, etc. telephone. food aperture, drink, raise table, telephone, sit, write, food aperture, sit, write, drink, write, rise, food aperture, sit, write, drink, rise, fold table, open bed, lie. cover self, asleep, rise, strength test, urinate, fold bed, raise table, sit. drink, write. p. M. 12h 28m 12 36 12 40 1 02 1 06 1 08 1 10 1 14 2 50 3 24 3 30 4 08 sit. sit, write. sit, feet on table. urinate. write, telephone. fold table, sit. lie, cover self. asleep. awake. sit, lie. food aperture. rise, urinate, fold bed, raise table, write, rise, food aperture, sit, drink, lie. lean on elbow, lie, asleep. rise, raise table, sit, urinate, drink, write, strength test, read, write, turn chair, lean on table, fold table, open bed, lie, asleep, rise, urinate, open bed, undress, close curtain, re- tire. December 7. A.M. 5h 30m rise, urinate, lie, asleep. 22 25 32 40 22 02 04 7 16 7 34 7 44 8 35 9 02 11 02 11 04 Experiment No. 85, N. M. P., Dec. 9, 7 a. m., to Dec. 11, 7 a. m., 1905. A. M. 7h00" 7 02 7 04 08 10 12 \ 20 , 22 24 26 30 7 32 December 9. rise. telephone. remove thermome- ter. urinate. dress, fold bed. weigh self, etc., tel- ephone. telephone. raise table. food aperture. adjust thermome- ter and pneumo- graph. sit, write. A. M. 7h 36m rise, sit. 7 40 drink. 7 44 stretch arms. 7 50 write. 8 28 sit. 8 34 write. 8 48 sit, arms over head. 06 telephone. 08 rise, food aperture. 10 sit, rise, strength test. 12 write. 15 drink. 26 sit. 30 write. A. M. 9h 50m telephone. 9 52 rise, food aperture. 9 54 sit. 10 04 read. 10 24 turn chair, read. 10 28 telephone. 10 32 read. 10 52 stretch arms over head. 11 04 telephone. 11 12 telephone, food ap- erture. 11 14 sit, read. 11 26 study. 11 44 rise, move about. 11 48 sit. Metabolism Experiments Nos. 79-83, 85, and 89. 241 Movements of Subjects. — Continued. December 9 (cont.) p. M. p. M. 11' ' 08m urinate. 12' . 04>" sit erect. 11 12 close curtain, re- 12 06 read. tire. 12 1 48 00 study. telephone. December 10. 1 02 stand, urinate. A M. 1 04 food aperture. 71 00" 1 rise. 1 06 drink, sit, study. 7 02 dress, urinate. 1 14 rise. 7 04 ) telephone, weigh 1 16 write. 7 11 f self, etc. 1 18 fold table, open 7 12 telephone. bed. 7 14 drink. 1 20 lie, read. 7 16 telephone. 1 46 telephone. 7 17 food aperture. 2 04 asleep. 7 22 sit. 2 08 turn over. 7 24 adjust thermome- 3 02 turn over. ter. 3 08 rise, urinate, 7 34 raise table. drink. 7 36 sit, write. o 10 move about, lie, 7 40 telephone. read. 7 46 rise, remove ther- 3 38 sit, strength test. mometer. 3 40 telephone. 7 48 food aperture. 3 42 write. 7 50 sit. 3 50 lie, read. 7 54 rise, stand. 4 40 stop reading. 7 56 food aperture. 5 00 turn over. 8 02 sit, write. 5 56 sit, lie. 8 08 telephone. 7 02 sit. 8 10 rise, move about. 7 03 food aperture. food aperture. 7 06 move about. 8 12 sit, read. 7 08 close curtain, uri- 8 32 read. nate. 9 02 telephone, write. 7 10 open curtain. 9 08 rise, food aperture. 7 12 move about. 9 10 sit, take tempera- 7 16 sit. ture. 7 24 stand. 9 16 rise, food aperture. 7 26 sit. 9 18 sit, telephone. 7 28 stand. 9 20 rise, move about. 7 32 telephone. 9 24 rise, food aperture. 7 34 raise shield, sit, 9 32 drink. read. 9 34 lie. 7 38 raise table, sit, 9 36 read. read. 9 46 telephone. 9 06 strength test. 9 48 telephone. 9 12 rise, open bed. 9 50 sit, food aperture. 9 14 lie, read. 9 52 lie, telephone. 9 16 sit, read. 9 54 sit, food aperture. 11 00 telephone. 9 56 clean food aper- 11 02 undress. ture. P. M. 9h 58m 10 08 10 24 10 44 11 02 11 04 11 10 11 16 11 20 11 28 11 34 11 36 P. M. lh021' 1 1 1 1 1 1 2 2 2 2 3 4 4 4 7 7 7 7 7 7 7 9 9 9 09 10 12 20 38 46 20 24 44 56 18 02 04 08 00 04 06 14 28 42 44 20 24 26 10 18 10 20 10 22 10 30 11 04 11 06 11 08 lie. telephone. doze. read. telephone, sit, food aperture, rise, sit, adjust thermometer, sit, read, telephone, sit. read. telephone, strength test, sit, read. telephone, urinate. food aperture. sit. write. read. telephone. read. rise, food aperture, move about. lie. doze. read. doze. telephone. sit, strength test. lie, read. urinate. telephone, food ap- erture. sit, read. rise, sit. telephone. telephone. food aperture. strength test. sit. write. fold table. lie. move about. lie. urinate. undress. close curtain, re- tire. Experiment No. 89, D. W., Jan. 10, 7 a. m., to Jan. 12, 7 a. m., 1906. January 10. a. m. a. M. 7" 07m urinate. 7h 00m rise, fold bed. 7 08 ) . . ... . 7 04 urinate, adjust 7 15 } weigh self, etc. chair. 7 16 adjust chair, tele- 7 06 dress, telephone. phone. 16 A. M. 7b20m arrange bed, ad- just table, sit, write. 7 26 read. 7 52 telephone. 242 Influence of Inanition on Metabolism. Movements of Subjects. — Continued. January 10 (cont.) A. M. 7" 54m rise, remove ther- mometer. 7 56 food aperture. 8 02 sit, read. 8 40 stop reading, play cards. 9 08 telephone. 9 10 food aperture. 9 12 open bed, lie. 10 48 turn over. p. M. 12h 22™ awake. 12 30 read. 12 50 sit, telephone. 12 52 lie, read. 1 00 urinate. 1 04 sit. 1 06 adjust thermome- ter, write. 1 08 move about. 1 12 read. 1 36 telephone. 1 38 strength test. 1 44 sit, write. 1 46 read. 2 48 write. 2 50 play cards. 3 02 telephone. 3 06 rise, food aperture. 3 08 lie, sit. 5 04 awake. 5 26 rise, fold bed, ad- just table, sit, write. 6 08 read. 7 02 urinate. 7 06 telephone. 8 00 strength test. 8 10 lie, read. 8 56 turn over. 9 40 telephone. 9 42 stop reading, play cards. 46 food aperture. 48 sit, play cards. stand, strength test. 10 08 sit, play cards. 10 26 stop playing. 9 9 10 00 P. M. 10h 28m arrange pneumo- graph, food aper- ture. 10 30 write. 10 42 stop writing. 10 50 sit erect. 11 02 telephone, close table, urinate. 11 04 undress, open bed. 11 06 retire. January 11. A.M. 3h50m wake, rise, arrange pneumograph. 3 52 sit, write. 3 56 food aperture, drink. 3 58 retire. 7 00 rise, fold clothes. 7 04 adjust chair. 7 06 7 11 [•weigh self, etc. 7 12 food aperture, tele- phone. 7 14 urinate. 7 16 adjust table, tele- phone. 7 18 food aperture. 7 20 sit, write. 8 00 drink. 8 30 sit erect. 8 32 read. 9 02 telephone. 9 04 rise, food aperture. 9 06 open bed. 9 08 ) lie, asleep, awake, \ asleep. 11 00 P. M. 12h 06™ awake. 1 02 telephone. 1 04 urinate. 1 06 move about. 1 08 drink, sit, write. 1 10 rise, food aperture, drink. 1 12 sit. 1 28 write. 1 42 strength test. 1 46 write. 1 56 rise, food aperture 2 02 sit, read. P. M. 3h02m fold table, open bed. 3 04 lie, asleep. 5 00 awake. 5 24 fold bed, adjust table. 5 26 sit, write. 5 30 food aperture. 5 32 sit, mend bed, drink. 5 34 sit, write. 5 41 food aperture. 6 02 play cards. 6 52 telephone. 7 02 urinate. 7 04 food aperture. 7 06 telephone. 7 08 food aperture. 7 10 read. 7 14 stop reading. drink, write. 7 30 telephone. 8 00 stop writing. 8 02 strength test. 8 06 sit, write. 8 10 stop writing, drink. 8 12 remove table, ar- range bed, lie. 9 14 rise, adjust table, write. 9 16 write. 9 17 drink, read. 9 44 stop reading, write. 10 10 stop writing, stand. 10 12 strength test. 10 14 sit, write. 10 20 telephone, food ap- erture. 10 22 play cards, food ap- erture, drink. 10 24 play cards. 10 42 stop playing. 10 46 sit quietly. 11 04 drink, telephone, urinate, undress. 11 06 remove table, ar- range bed, close curtain, retire. Metabolism Experiments Nos. 79-83, 85, and 89. 243 Drinking-water. — As in the earlier experiments the subjects were allowed drinking-water whenever desired with no restriction as to the amounts. The quantities consumed were not accurately measured except for each 24 hours, but the estimated amounts per period recorded in table 162 are probably not far from correct. An examination of table 162 shows wide variation in the daily intake of water for the different experiments. The minimum amount consumed during 24 hours by any subject was 115.1 grams, and the maximum was 1467.1 grams. In general these subjects drank less water than the 3 men who were the subjects of experiments Nos. 59 and 68 to 77. Water from the city supply was used and no allowance was made for the small amounts of organic matter and salts which it contained. Table 162. — Record of water consumed — Metabolism experiments Nos. 79-83, 85, and 89. Experiment number, subject, and date. 7 to 9 a. m. 9 toll a. m. 11a.m. to 1 p. m. 1 to 3 p. m. 3 to 5 p. m. 5 to 7 p. m. 7 to 9 p. m. 9 to 11 p. m. 11p.m. to la.m. Total tor day. Experiment No. 79, H.E.S.: Oct. 13-14. 1905 Oct. 14-15, 1905 Experiment No.80,C.R.Y.: Oct. 2 -28.1905 Oct. 28-29, 1905 Experiment No. 81, A. H.M.: Nov. 21-22, 1905 Nov. 22 23, 1905 Experiment No. 82.H.C.K.: Nov. 24-25. 1905 Nov. 25-26, 1905 Experiment No.83, H.R.D.: Dec. 5-6, 1905 Dec. 6-7. 1905 Experiment No. 85, N.M. P.: Dec. 9-10,1905 Dec. 10 11,1905 Experiment No. 89, D. W.: Jan. 10-11, 1906 Jan. 11-12, 1906 Gms- 224.8 147.5 163.3 146.1 124.3 103.1 148-0 448.4 147.2 125.7 288.6 ^115.1 48.8 Gm 224 147 125 183 Gms Gms. 161.1 38.4 219.6 147.2 104.5 148.0 219.5 140.2 39! 2 Gms. 104.5 294.3 154.0 221.3 140.1 64.4 Gms. 40.9 104.6 39.1 Gms. 102.5 164.0 104.1 69.6 292.7 212.8 221.3 86.5 85.5 73.0 Gms. 295.8 105.0 212.8 86.4 85.6 115.1 Gms. '132 »143 145 104 42.0 Gms. 782.5 339.9 132.8 206.4 291.1 193.8 857.8 1092.9 1467.1 884.2 704.6 707.7 115.1 357.2 1 Amounts known. In other experiments one or more of the amounts per period are esti- mated. 2 3 to 5 a. m. Ukine. The urine was collected daily at 7 a. m., 1 p. m., 7 p. m., and 11 p. m. Unfortunately the subject of experiment No. 79 did not collect the urine on the first morning until an hour and a half after the experiment began. It was impossible to make any satisfactory allowance for the excess included in the preliminary urine and no attempt has been made to correct for it. The error would probably be proportional to the time of the experiment and hence from 6 to 10 per cent would represent the error. During experiment No. 85, a small quantity of the urine for the second period of December 9 was spilled in the food aperture and on the floor of the chamber. This was immediately taken up with cheese cloth, and the nitrogen content determined and added to the total nitrogen for the day. The total weight of the urine 244 Influence of Inanition on Metabolism. spilled was obtained by the ratio of its nitrogen content to the nitrogen of the measured urine. The same method was employed in calculating the carbon, hydrogen, ash, and solid matter in the urine spilled. While it is impossible that all the urine was taken up by the cheese cloth, it is believed that the error introduced is inappreciable. As stated above, the amount of urine was obtained for each period in all experiments. The specific gravity of the urine for the different periods, how- ever, was not taken, except during the first day of experiment No. 79, and the Table 163. — Amounts and specific gravity of urine — Metabolism experiments Nos. 79-83, and 85. Experiment number, subject, and date. 8h33m a. m. to 1 p. m. 7 a. m. to 1 p. m. 1 to 7 p. m. 7 to 11 | p. m. 11 p. m. to 7 a. m. Total for day Specific grav- ity. Experiment No. 79, H. E. S. : Oct. 13-14, 1905 Oct. 14-15, 1905 Experiment No. 80, C. R. T. Oct. 27-28, 1905 Oct. 28-29, 1905 .. Experiment No. 81, A. IT. M. Nov. 21-22, 1905 Nov. 22-23, 1905 Experiment No. 82, H. C. K. Nov. 24-25, 1905 Nov. 25-26, 1905 Experiment No. 83, H. R. D. Dec. 5-6, 1905 Dec. 6-7, 1905 Experiment No. 85, N. M. P. Grams. J591.4 Dec. 9-10, 1905. Dec. 10-11, 1905. Orani8. 214 . 5 186.2 254.2 199.4 191.4 193.3 161.3 558.2 335.5 332.7 192.3 Grams. Grams. 2 171.5 3 76.7 186.6 132 210.4 168.7 133.5 213.4 124.5 343.7 246.5 306.2 618.3 5 34.6 267.2 180.9 111.0 82.9 138.9 62.6 931.9 124.4 159.7 106.0 80.3 Grams. 4187.6 325.6 569.9 243.5 244.2 287.5 195.3 358.1 275.4 263.6 131.0 139.6 Grams. 1027.2 859.1 1147.4 777.4 660.0 831.2 575.7 1795.0 1204.5 1065.0 1222.6 679.4 1.0118 1.0223 1.0169 1.0237 1.0264 I . 0230 1.0261 1.0127 1.0146 1.0186 1.0154 1.0203 1 Specific gravity, 1.0057. 2 Specific gravity, 1.0169. * Specific gravity, 1.0220. 4 Specific gravity, 1.0233. 0 Calculated as spilled. This Is included In the total for the day. See p. 243. 2 days of experiment No. 89. Except in the latter experiment, determinations of nitrogen and other constituents of the urine were made only on the daily composites. The amounts of urine by periods and the specific gravity for each day are given in table 163. The corresponding data for experiment No. 89 are given in a separate table, for the reason that in this experiment the nitrogen was determined on the urine for each period. Weight and composition of urine. — In table 165 are recorded the determi- nations of elements and compounds in the urine for each day of the series, together with the heat of combustion of the organic matter of each daily sample. In considering the table, special attention should be paid to the note on experiment No. 85. Metabolism Experiments Nos. 79-83, 85, and 89. 245 Elimination of Water- Vapob. The total water of respiration and perspiration eliminated per period and per day and the total amount of water-vapor remaining in the calorimeter chamber at the end of each period are shown in table 166. In the notes at the foot of the table are shown the changes in weight of the chair, bed, bedding, etc., in the chamber. The object of recording these data has already been explained. (See p. 30.) Elimination of Carbon Dioxide and Absobption of Oxygen. Table 167 records the total weights of carbon dioxide exhaled and of oxygen consumed. The method of obtaining the data has been previously explained. Table 164. — Determinations in urine per period and per day — Metabolism experiment No. 89. Date. Period. (a) Amount. (6) Specific gravity. (o) Volume (a+b). (d) Nitro- gen. 1906. Jan. 10-11. Jan. 11-12. 7 a.m. to 1 p.m 1 p.m. 7 p.m 7p.m. 11 p.m 11 p.m. 7 a.m Total Total by composite 7 a.m. to 1 p.m 1 p.m. 7 p.m 7 p.m. 11 p.m 11 p.m. 7 a.m Total Total by composite Total for 2 days. . . Grams. 225.2 200.6 84.3 134.3 644.4 644.4 126.5 142.5 91.4 179.2 539.6 539.6 1184.0 1.0264 1.0280 1.0303 1.0326 1.0296 c. c. 219 195 82 130 626 626 1.0319 123 1.0340 138 1.0329 88 1.0318 173 522 1.0338 522 1148 Grams. 2.82 2.73 1.49 2.95 9.99 10.47 3.18 3.69 2.54 5.05 14.46 14.43 24.45 Since the measurement of these gases during the preliminary night furnishes a part of the data regarding the transition from the katabolism with food to katabolism during inanition, the amounts for the preliminary period have been included whenever possible. Elements Katabolized in the Body. The elements katabolized in the body during each of the experiments are recorded in table 168. The usual method of computation was followed in obtaining the data. Preparatory to substitution in the formulas used in computing the materials katabolized, the results have been carried to the second decimal 32 place. 32 See discussion of the use of significant figures, p. 17. 246 Influence of Inanition on Metabolism. Table 165. — Weight, composition, and heat of combustion of urine — Metabolism experiments Nos. 79-83, 85, and 89. Experiment No. 79. H. E. S. Oct. 13-14, 1905. Oct. 14-15. 1905. Experiment No. 80. C. K. Y. Oct. 27-28, 1906. Oct. 28-29, 1905. (a) Weight grams (b) Water do. . (c) Solids, a — b do.. (d) Ash do. . (e) Organic matter, c—d do.. (/) Nitrogen do. . (g) Carbon do . . (h) Hydrogen in organic matter. . . .do. . (i) Oxygen (by difference) in organic mat ter, e — (f+g + h) grams (j ) Phosphorus do. . Phosphoric acid (P205) : (k) by fusion grams (I) by titration do (in) Sulphur do. . Sulphur trioxide (S03) : (n) Total grams (o) Inorganic and ethereal do. . (p) Neutral, n—o do. . (q) Creatine (preformed). do . . (r) Total creatinine do. . (s) Creatinine l (preformed), r—q . . .do. . (t) Chlorine do. . (w) Sodium chloride do. . (v) Heat of combustion calories 1027.2 996.08 31.12 8.73 22.39 8.11 6.27 1.75 6.26 .472 1.081 1.124 .543 1.355 1.162 .193 1.212 1.225 .013 2.917 4.817 72 859.1 809.96 49.14 11.17 37.97 14.35 10.22 3.09 10.31 .726 1.664 1.765 .777 1.940 1.701 .239 1.254 1.418 .164 3.622 5.976 116 1147.4 1105.75 41.65 18.93 22.72 7.78 7.11 2.07 5.76 .517 1.184 1.267 .389 .969 .827 .142 1.505 1.505 8.898 14.685 75 777.4 734.80 42.60 12.52 30.08 9.95 9.02 2.57 8.54 1.057 2.421 2.754 .685 1.711 1.483 .227 1.398 1.538 .140 4.028 6.646 97 Experiment No. 81. A. H. M. Nov 21-22, Nov. 22-23, 1905. 1905. Experiment No. 82. H. C. K. Nov. 24-25, 1905. Nov. 25-26, 1905. (a) Weight grams. (b) Water do . . . (c) Solids, a — b do. . . (d) Ash do. . . (e) Organic matter, c — d do. . . (/) Nitrogen do. . . (g) Carbon do. . . (h) Hydrogen in organic matter do. . . (i) Oxygen (by ditlerence) in organic mat- ter, e — (f+g+h) grams. (j) Phosphorus do. . . Phosphoric acid (P205) : (k) by fusion grams. (I) by titration do... (m) Sulphur ... .do. . . Sulphur trioxide (SO.,): (n) Total grams. (o) Inorganic and ethereal do . . . (p) Neutral, n—o do... (q) Creatinine (preformed) do. . . (r) Total creatinine do. . . («) Creatine1 (preformed), r — q do. . . (t) Chlorine do. . . («) Sodium chloride do . . . (v) Heat of combustion calories. 660.0 621.26 38.74 12.61 26.13 9. 7. 9, .11 .52 .18 7.32 .643 1.472 1.470 .572 .427 .295 .132 184 .226 .042 3.880 6.402 80 831.2 783.16 48.04 11.22 36.82 13.05 9.48 2.91 11.38 1.060 2.429 2.505 .817 2.036 1.891 .145 1.229 1.357 .128 2 . 794 4.612 109 575.7 538 . 57 37.13 9.67 27.46 9.38 7.71 2.07 8.30 .486 1.113 1.132 .570 1.423 1.230 .193 1.643 1.726 .083 3.451 5.694 88 1795.0 1740.07 54.93 16.33 38.60 14.36 10.77 3.59 9.88 .873 000 017 768 1.915 1.746 .169 1.730 1 . 795 .065 6.714 11.079 122 1 In terms of creatinine. Metabolism Experiments Nos. 79-83, 85, and 89. 247 Table 165. — Weight, composition, and heat of combustion of urine — Continued. Experiment No. 83. H. R. D. Dec. 5-6, 1905. Dec. 6-7, 1905. Experiment No. 85. N M. P. Dec. 9-10, 1905. Dec. 10-11, 1906. Experiment No. 89. D.W. Jan. 10-11, 1906. Jan. 11-12, 1906. (a) Weight grams (b) Water do.. (c) Solids, a — b do.. (d) Ash do. . (e) Organic matter, c—d.. do. . (/) Nitrogen do . . (g) Carbon do . . {h) Hydrogen in organic mat- ter grams (i) Oxygen (by difference) in organic matter, e—(f + g + h) grams (J) Phosphorus do . . Phosphoric acid (P205): (k) by fusion grams (0 by titration do.. (m) Sulphur do.. Sulphur trioxide (S03) : («) Total grams (o) Inorganic and ethereal, srrams (p) Neutral, n — o.. grams (q) Creatinine (preformed), grams (r) Total creatinine. . .grams (s) Creatine4 (preformed), r — q grams. . (t) Chlorine do.. (u) Sodium chloride. . . .do. . , (v) Heat of combustion. cals. . 1204.5 1159.21 45.29 8.19 37.10 13.25 9.76 2.65 11.44 1.051 2.407 2.414 .871 2.173 1.961 .212 1.217 1.317 .100 .517 .853 116 1065.0 1012.39 52.61 8.73 43.88 13.53 12.78 3.20 14.37 1.219 2.793 2.829 .703 1.755 1.580 .175 1.129 1.363 .234 .628 1.036 151 i 34.6 1188.0 1 33.37 1145.11 1 1.25 42.89 1 .36 12.47 1 .89 30.42 2 .32 11.05 1 .22 7.48 1 .06 1.90 1 .29 9.99 3 .549 3 1.257 3 1.283 3 .604 1.507 1.219 .288 1.525 1.543 .018 4.582 7.562 3 98 i 679.4 I 642.44 | 36.96 i 5.57 t 31.39 I 11.35 I 8.22 I 2.11 9.71 .519 1.189 1.516 .603 1.506 1.331 .175 1.541 1.703 .162 1.461 2.411 {" 4 } 644.4 599.55 44.85 13.34 31.51 9.99 8.38 2.26 10.88 1.027 2.354 .693 1.731 1.472 .258 2.028 2.091 .063 5.857 9.666 99 93 539.6 492 22 47.38 7.45 39.93 14.46 9.87 2.81 12.79 1.368 3.134 .861 2.148 1.831 .317 1.922 1.966 .044 1.805 2.978 114 1 Calculated. See p. 243. 2 Amount of nitrogen determined for urine lost. See p. 243. 3 Does not include possible amount in urine lost. 4 In terms of creatinine. 248 Influence of Inanition on Metabolism. Table 166. — Record of water of respiration and perspiration — Metabolism experi- ments Nos. 79-83, 85, and 89.1 Date and period. (a) Total amount of vapor in chamber at end of period. (6) Total water of respira- tion and perspira- tion. Date and period. (a) Total amount (b) Total water of of vapor in chamber respira- tion and at end of period. perspira- tion. EXPERIMENT No. 79. H. E. S. 1905. Oct. 13: Preliminary : 5 a.m. to 7 a.m.. Oct. 13-14: 7 a.m. to 9 a.m. 9 a.m. 11 a.m. 11 a.m. I p.m. lp.m. 3 p.m. 3 p.m. 5 p.m. 5 p.m. 7 p.m. 7 p.m. 9 p.m. 9 p.m. 11 p.m. 11 p.m. 1 a.m. 1 a.m. 3 a.m. 3 a.m. 5 a.m. 5 a.m. 7 a.m. Total Oct. 14-15: 7 a.m. to 9 a.m. 9 a.m. 11 a.m. 11 a.m. 1 p.m. 1 p.m. 3 p.m. 3 p.m. 5 p.m. 5 p.m. 7 p.m. 7 p.m. 9 p.m. 9 p.m. 11 p.m. 11 p.m. 1 a.m. 1 a.m. 3 a.m. 3 a.m. 5 a.m. 5 a.m. 7 a.m. Total Grams. 51.6 53.8 50.4 51.2 48.6 50.0 49.4 48.3 47.4 49.0 53.6 53.0 52.3 57.2 54.2 57.2 53.9 54.7 50.8 50. S 49.7 53.3 54.4 54.0 50.4 Grams. 53.0 55.5 55.6 55.6 54.8 55.7 52.9 54.4 54.1 63.0 54.2 58.2 667.0 55.2 57.7 63.9 57.9 58.8 59.8 61.2 61.4 59.8 61.0 54.2 53.2 704.1 EXPERIMENT No. 80. C. R. Y. 1905. Oct. 26-27: Preliminary : 11 p.m 11 p.m. to 1 a.m. 1 a.m. 3 a.m. 3 a.m. 5 a.m. 5 a.m. 7 a.m. Total, 8 hours Oct. 27-28 : 7 a.m. to 9 a.m. 9 a.m. 11 a.m. 11 a.m. 1 p.m. lp.m. 3 p.m. 3 p.m. 5 p.m. 5 p.m. 7 p.m. 7 p.m. 9 p.m. 9 p.m. 11 p.m. 11 p.m. 1 a.m. 1 a.m. 3 a.m. 3 a.m. 5 a.m. 5 a.m. 7 a.m. Total Oct. 28-29: 7 a.m. to 9 a.m. 9 a.m. 11 a.m. 11 a.m. lp.m. lp.m. 3 p.m. 3 p.m. 5 p.m. 5 p.m. 7 p.m. 7 p.m. 9 p.m. 9 p.m. 11 p.m. 11 p.m. 1 a.m. 1 a.m. 3 a.m. 3 a.m. 5 a.m. 5 a.m. 7 a.m. Total Grams. Grams. 51.7 .... 51.1 66.1 56.4 83.2 58.2 76.9 54.2 75.6 .... 2 301.8 55.7 67.2 49.6 47.5 46.0 84.2 41.2 74.3 43.5 75.7 38.4 76.1 37.8 70.1 34.8 71.0 40.2 73.2 47.2 95.2 52.2 88.6 53.9 103.7 .... 926.8 57.3 92.2 47.1 77.4 50.1 88.0 46.4 95.4 45.2 89.5 42.9 91.9 42.9 83.0 40.2 83.2 51.5 95 . 2 46.9 92.8 46.4 85.0 50.4 86.9 1060.5 1 Allowance has been made for water gained or lost by chair, bedding and miscellaneous articles as follows: Experiment No. 79, October 13-14, 17.0 grams lost; October 14-15, 6.8 grams lost ; experiment No. 80, October 27-28, 17.8 grams lost. 2 Does not include corrections for changes in weight of chair and bedding. Metabolism Experiments Nos. 79-83, 85, and 89. 249 Table 166. — Record of water of respiration and perspiration ' — Continued. Date and period. (a) (0) Total Total amount water of of vapor respira- in tion chamber and at end perspira- of period. tion. Date and period. (a) Total amount of vapor in chamber at end of period. (b) Total water of respira- tion and perspira- tion. EXPERIMENT No. 81. A. H. M. 1905. Nov. 21: Preliminary : 1 a.m 1 a. m to 3 a.m. 3 a.m. 5 a.m. 5 a.m. 7 a.m. Total, 6 hours. Nov. 21-22: 7 a.m. to 9 a.m. 9 a.m. 11 a.m. 1 p.m. 3 p.m. 5 p.m. 7p.m. 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. Total 11 a.m.. 1 p.m. , 3 p.m. . 5 p.m., 7 p.m. 9 p.m. , 11 p.m. - 1 a.m., 3 a.m. 5 a.m. , 7 a.m. Nov. 2i-23: 7 a.m. to 9 a.m. 9 a.m. 11 a.m. 1 p.m. 3 p.m. 5 p. m. 7 p.m. 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. Total. 1 1 a. m. . 1 p.m. 3 p.m. . 5 p.m. . 7 p.m. . 9 p.m. . 11 p.m. . 1 a. m. . 3 a.m. . 5 a.m.. 7 a.m. . Grams. 33.3 30.3 27.9 25.4 Grams. 60.0 55.6 53.9 ^ 169.5 31.5 32.1 29.0 28.1 26.3 25.6 25 3 23.9 21.1 24.7 25.5 26.5 56.1 63.4 54.6 51.8 51.6 52.6 48.3 43.6 46.6 45.6 45.3 49.1 31.3 30.1 27.8 25.1 25.6 26.9 27.3 27.5 27.7 27.1 26.3 28.6 608.6 56.5 64.3 55.1 50.9 49.7 58.2 56.4 57.5 55 . 2 55.7 54.7 56 . 5 670.7 EXPERIMENT No. 82. H. C. K. 1905. Nov. 23-24 : Preliminary : 7p.m 7p.m. 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. to 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. 7 a.m. Total, 12 hours. Nov. 24- 7 a.m. 9 a.m. 11 a.m. 1 p.m. 3 p.m. 5 p.m. 7 p.m. 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. 35: to 9 a.m. 11 a.m. 1 p.m. 3 p.m. 5 p.m. 7 p.m. 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. 7 a.m. Total. Nov. 25- 7 a.m. 9 a.m. 11 a.m. 1 p.m. 3 p.m. 5 p.m. 7 p.m. 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. 26: to 9 a.m. 11 a.m. 1 p.m. 3 p.m. 5 p.m. 7 p.m. 9p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. 7 a.m. Total. Grams. 47.1 36.1 31.0 32.1 29 . 4 31.0 34.0 45.0 39. 38 35.9 33.8 30.4 32.1 29.6 31.3 33.1 31.9 31.8 Grams. 73.7 67.5 69.6 64.5 65.2 77.0 2417.6 48.5 40.1 38.8 34.7 33.8 31.7 38.0 35.3 35.2 34.2 35.0 35.2 99.4 87.0 59.8 72.8 73.8 67.7 70.5 65.3 64.8 50.4 63.6 67.0 842.1 1 10.2 85.8 78.3 75.8 74.4 74.0 84.2 80.2 72.9 57.8 69.5 77.4 940.5 1 Allowance has been made for water gained or lost by chair, bedding, and miscellaneous articles as follows: Experiment No. 81, November 21-22, 47.9 grams lost; November 22-23, 4.9 grams lost ; experiment No. 82, November 24-25, 10.0 grams lost ; November 25-26, 24.0 grams gained. 2 Does not include corrections for changes in weight of chair and bedding. 250 Influence of Inanition on Metabolism. Table 166. — Record of water of respiration and perspiration1 — Continued. (a) (ft) (a) (ft) Total Total Total Total amount water of amount water of Date and period. of vapor in respira- tion Date and period. of vapor in respira- tion chamber and chamber and at end perspira- at end perspira- of period. tion. of period. tion. experiment no. i 53. H. R. D. EXPERIMENT No. 85. N. M. P. 1905. 1905. Dec. 4-5 : Dec. 8-9 : Preliminary : Grams. Grams. Preliminary : Grams. Grams. 36.7 34.5 11 p.m. to 1 a.m 40.2 47.1 36.1 70.2 35.4 73.0 34.9 60.7 34.2 62.4 30.5 59.0 Dec. 5-6: 30.9 60.3 Dec. 9-10: 31.4 57.7 2 272. 8 2247.6 - 38.6 68.3 7 a.m. to 9 a.m 38.1 76.8 31.8 57.4 36.6 68.4 32.2 57.9 11 a.m. 1 p.m 30.6 66.5 28.2 53.2 31.5 59.6 3 p.m. 5 p.m 27.3 51.7 28.3 59.6 5 p.m. 7 p.m. 31.3 60.5 31.3 65.4 29.5 58.6 32.7 73.6 26.8 52.8 30.4 61.3 28.3 53.3 29.9 62.8 30.7 61.2 30.5 60.0 30.1 53.3 30.9 59.9 5 a.m. 7 a.m Total 29.7 56.3 32.0 61.9 .... 684.5 .... 775.8 Dec. 6-7: Dec. 10-11: 38.6 78.8 40.2 73.7 9 a.m. 11 a.m 29.3 52.5 35.9 63.4 29.8 59.9 35.5 65.5 27.6 52.3 34.4 61.2 3 p.m. 5 p.m 28.1 54.6 33.4 59 9 26.7 54.0 34.3 68.7 7 p.m. 9 p.m 28.8 58.1 7 p.m. 9 p.m 36.1 70.1 26.8 53.6 33.9 68.9 29.4 55.4 35.8 68.0 28.0 51.8 1 a.m. 3 a.m 37.5 72.9 3 a.m. 5 a.m 27.8 48.1 3 a.m. 5 a.m 38.2 67.9 Total 29.9 52.9 37.0 73.2 672.0 813.4 1 Allowance has been made for water gained or lost by chair, bedding, and miscellaneous articles as follows : Experiment No. 83, December 5-6, 44.0 grams lost ; December 6-7, 13.0 grams lost ; experiment No. 85, December 9-10, 38.0 grams gained ; December 10-11, 7.0 grams lost. 2 Does not include corrections for changes in weight of chair and bedding. Metabolism Experiments Nos. 79-83, 85, and 89. 251 Table 166. — Record of water of respiration and perspiration * — Continued. Date and period. (a) Total amount of vapor in chamber at end of period. (6) Total water of respira- tion and perspira- tion. Date and period. (a) Total amount of vapor in chamber at end of period. (&) Total water of respira- tion and perspira- tion. EXPERIMENT No. 89. D. W. 1906. Jan. 9-10: Preliminary : 11 p.m Grams. 37.7 39.2 41.3 40.1 39.6 Grams. 44.6 44.4 46.1 42.4 1906. Jan. 10-11 : Total Grams. 32.0 35.1 Grams. 68.0 68.2 1 a.m. 3 a.m 3 a.m. 5 a.m Jan. 10-11: 7 a.m. to 9 a.m 3 p.m. 5 p.m 7 p.m. 9 p.m 9 p.m. 11 p.m .... 819.6 Jan. 11-12 : Total 34.9 32.1 34.4 36.8 32.7 37.2 37.2 37.7 36.2 33.5 32.3 31.6 65.7 61.2 68.5 64.5 61.9 73.7 79.9 69.2 69.9 61.6 64.2 62.4 2177.5 36.5 33.7 33.0 36.5 31.5 33.5 35.7 38.8 36.2 33.2 68.1 64.1 66.1 71.0 i 64.6 71.1 72.5 73.2 69.2 63.5 802.7 1 Allowance has been made for water gained or lost by chair, bedding, and miscellaneous articles as follows: Experiment No. 89, January 10-11, 9.5 grams lost; January 11-12, 0.3 gram lost. 2 Does not include corrections for changes in weight of chair and bedding. 252 Influence of Inanition on Metabolism. Table 167. — Record of carbon dioxide and oxygen — Metabolism experiments Nos. 79-83, 85, and 89. EXPERIMENT No. 79. H. E. S. Date. Period. Carbon dioxide. Oxygen. (a) Amount in (b) Total (c) Amount in W) Total chamber weight chamber amount at end of exhaled at end of consumed period. by subject. period. by subject. 1905. Oct. 12-13 . . . Preliminary: Grams. 48.9 Grams. Liters. Grams. 11 p. m. to 1 a. m. . 46.6 52.7 1 a. m. 3 a. m. . 39.5 41.0 3 a. m. 5 a. m. . 44.4 48.1 Oct. 13-14... 5 a. m. 7 a. m. . Total, 8 hours 7 a. m. to 9 a. m. . 39.9 41.2 893.7 .... 183.0 .... 55.4 67.8 881.9 67.3 9 a. m. 11 a. m. . 50.5 54.2 884.6 42.7 11 a. m. 1p.m.. 51.1 55.1 877.2 49.7 1 p. m. 3 p. m. . 51.2 57.4 878.3 48.5 3 p. m. 5 p. m. . 49.4 50.7 877.2 47.9 5 p.m. 7 p. m. . 47.8 51.7 882.2 49.6 7 p. m. 9 p. m. . 46.3 50.4 889.1 42.0 9 p. m. 11 p. m. . 46.5 51.2 887.7 52.0 lip. m. 1 a. m. . 46.8 51.0 886.9 41.8 1 a. m. 3 a. m. . 49.8 52.0 886.1 51.3 3 a. m. 5 a. m . . 47.0 43.4 894.7 36.7 Oct, 14-15. .. 5 a. m. 7 a. m. . Total 46.0 47.1 903.0 46.4 .... 632.0 575.9 7 a. m. to 9 a. m. . 62.9 65.7 899.5 66.4 9 a. m. 11 a.m.. 58.6 54.9 903.2 45.2 11a. m. 1p.m.. 57.6 54.1 894.7 53.2 1 p. m. 3 p. m. . 59.2 56.3 896.0 55.0 3 p. m. 5 p. m. . 57.5 51.3 898.7 51.5 5 p. m. 7 p.m.. 55.6 54.6 909.3 46.9 7 p. m. 9 p. m. . 48.5 56.8 913.6 52.4 9 p. m. 11 p. m. . 40.1 45.3 919.2 47.8 11 p. m. 1 a. m. . 43.7 51.3 908.8 51.8 1 a. m. 3 a. m. . 45.4 52.1 903.2 48.8 3 a. m. 5 a. m . . 44.3 45.5 907.5 43.4 5 a. m. 7 a. m. . Total 43.2 47.3 912.7 42.9 .... 635.2 605.3 EXPERIMEJ JT No. 80. ( 3. R. Y. 1905. Preliminary: Oct. 26-27 . . . 11 p. m 52.2 40.2 61.6 935.8 941.3 32 .2 11 p. m. to 1 a. m. . 1 a. m. 3 a. m . . 32.3 47.5 932.4 40.4 3 a. m. 5 a. in. . 32.3 50.2 933.8 39.1 5 a. in. 7 a. m. . Total, 8 hours 32.7 52.4 948.9 33.0 211.7 144.7 Metabolism Experiments Nos. 79-83, 85, and 89. 253 Table 167. — Record of carbon dioxide and oxyget , — Continued. EXPERIMENT No. 80. C. R. Y. (Cont'd). Date. Period. Carbon dioxide. Oxygen. (o) Amount in chamber at end of period. (6) Total weight exhaled by subject. (c) Amount in chamber at end of period. (d) Total amount consumed by subject. 1905. Oct. 27-28... 7 a. m. to 9 a. m. . Grams. 45.4 Grams. 73.6 Liters. 938.7 Grams. 70.8 9 a. m. 11 a. m. . 42.1 58.7 950.6 43.9 11a. in. 1 p. m. . 32.7 55.2 950.8 46.3 1 p.m. 3 p. m. . 24.7 49.0 955.8 39.3 3 p. m. 5 p. m. . 30.9 51.3 945.6 50.8 5 p. m. 7 p. m. . 22.5 46.2 960.4 43.9 7 p. m. 9 p. m. . 25.0 43.3 960.9 42.2 9 p. m. 11 p. m. . 19.5 43.6 966.8 40.3 lip. m. 1 a. m. . 24.2 58.7 956.3 55.8 1 a. m. 3 a. m. . 23.8 47.0 944.2 43.7 3 a. m. 5 a. m . . 27.3 50.8 936.3 51.8 Oct. 28-29... 5 a. m. 7 a. in. . Total 24.1 50.0 937.4 47.4 627.4 576.2 7 a. m. to 9 a. m. . 35.6 64.3 928.5 68.7 9 a. m. 11 a. m. . 27.2 55.0 946.7 48.1 11 a. m. 1 p. m. . 30.9 49.5 938.1 46.6 1 p. m. 3 p. m. . 24.4 52.0 941.2 53.2 3 p. m. 5 p. m. . 28.2 48.2 948.7 49.6 5 p. m. 7 p. m. . 25.0 56.5 957.4 59.5 7 p. m. 9 p. m. . 22.2 51.6 960.3 48.8 9 p. m. 11 p. m. . 23.5 49.3 966.5 44.1 11 p. m. 1 a. m. . 23.4 57.0 952.1 59.0 1 a. m. 3 a. m . . 21.9 47.7 957.1 43.7 3 a. m. 5 a. m. . 24.0 52.5 963.0 53.7 5 a. m. 7 a. m. . Total 32.3 56.6 958.5 53.5 640.3 628.5 EXPERIMENT No. 81. 1 L H. M. 1905. Nov. 21 Preliminary: 1 a. m 26.1 946.4 1 a. m. to 3 a. m. . 15.6 34.7 956.0 28.1 3 a. m. 5 a. m. . 21.0 37.9 956.7 30.9 Nov. 21-22... 5 a. m. 7 a. m. . Total, 6 hours 7 a. m. to 9 a. m. . 18.7 39.9 957.6 32.2 .... 112.5 91.2 28.2 61.6 952.1 52.6 9 a. m. 11 a. m. . 29.5 53.1 945.5 52.7 11 a. m. 1 p. m. . 27.4 45.7 941.5 41.6 1 p. m. 3 p. m. . 28.1 42.4 934.4 42.4 3 p. m. 5 p. m. . 29.4 45.8 938.3 46.3 5 p. m. 7 p. m. . 29.1 44.0 942.6 45.5 7 p.m. 9 p. m. . 31.7 44.3 941.2 44.9 9 p. m. 11 p. m. . 21.0 44.4 942.9 41.0 11 p. m. 1 a. m. . 25.3 37.9 935.5 36.6 1 a. m. 3 a. m. . 18.4 35.5 936.4 38.9 3 a. m. 5 a. m. . 19.3 40.1 937.3 35.4 5 a. m. 7 a. m. . Total 21.3 39.9 925.2 38.9 534.7 516.8 254 Influence of Inanition on Metabolism. Table : 167. — Record of carbon dioxide and oxygen — Continued. EXPERIMENT No. 81. A. H. M. (Cont'd). Date. Period. Carbon dioxide. Oxygen. (a) . Amount in (b) Total (c) Amount in (d) Total chamber weight chamber amount at end of exhaled at end of consumed period. by subject. period. by subject. 1905. Grams. Grams. Liters. Grams. Nov. 22-23... 7 a. m. to 9 a. m. . 29.0 59.7 914.4 62.3 9 a. m. 11 a. m. . 30.8 51.7 909.5 50.5 11 a. m. 1 p. m. . 27.4 49.2 898.8 46.7 1 p. m. 3 p. m. . 25.0 38.3 900.4 35.1 3 p. m. 5 p. m. . 22.1 37.3 894.3 33.3 5 p.m. 7 p. m. . 29.3 44.8 882.4 52.8 7 p. m. 9 p. m. . 28.1 46.9 878.9 43.1 9 p. m. 11 p. m. . 30.1 39.6 877.2 45.9 11 p. m. 1 a. m. . 27.5 41.6 877.0 40.8 1 a. m. 3 a. m. . 29.7 33.9 870.4 32.2 3 a. m. 5 a. m. . 26.5 36.3 874.6 38.8 5 a. m. 7 a. m. . Total 22.0 45.0 872.0 45.6 .... 524.3 527.1 EXPERIMEN T No. 82. 1 [. C. K. 1905. Preliminary: Nov. 23-24... 7 p. m. " 44.4 34.9 69.9 885.4 900.1 55^3 7 p. m. to 9 p. m. . 9 p. m. 11 p. m. . 29.6 62.1 904.3 50.8 11 p. m. 1 a. m. . 25.5 60.2 897.6 46.0 1 a. m. 3 a. m. . 23.5 46.5 902.5 33.9 3 a, m. 5 a. m. . 24.1 51.5 896.8 46.7 Nov. 24-25... 5 a. m. 7 a. m. . Total, 12 hours 7 a. m. to 9 a. m. . 27.2 52.0 889.0 .... 342.2 35.5 82.7 868.2 80.2 9 a. m. 11 a. m. . 39.3 72.1 863.9 65.9 11 a. m. 1 p. m. . 43.3 65.8 844.9 54.3 1 p. m. 3 p. m. . 43.6 71.3 840.8 63.7 3 p. m. 5 p. m. . 33.1 56.0 848.0 47.2 5 p. m. 7 p. m. . 34.2 56.7 848.0 47.1 7 p. m. 9 p. m. . 42.7 70.4 834.2 69.2 9 p. m. 11 p. m. . 32.1 62.9 840.8 58.4 11 p. m. 1 a. m. . 36.9 55.8 827.1 47.3 1 a. m. 3 a. m. . 37.9 47.7 821.4 44.6 3 a.m. 5 a. m. . 25.7 48.3 826.4 41.5 5 a. m. 7 a. m. . Total 29.4 51.2 824.2 43.9 740.9 663.3 Metabolism Experiments Nos. 79-83, 85, and 89. 255 Table 167. — Record of carbon dioxide and oxygen — Continued. EXPERIMENT No. 82. H. C. K. (Cont'd). Date. Period. Carbon dioxide. Oxygen. (a) Amount in « Total (c) Amount in (d) Total chamber weight chamber amount at end of exhaled at end of consumed period. by subject. period. by subject. 1905. Grams. Grams. Liters. Grams. Nov. 25-26... 7 a. m. to 9 a. m. . 40.7 88.8 803.5 85.6 9 a. m. 11 a. m. . 36.3 63.0 814.1 59.4 11 a. m. 1p.m.. 42.5 72.0 801.6 70.9 1 p. m. 3 p. m. . 42.0 62.8 812.1 59.3 3 p. m. 5 p. m. . 42.2 64.2 819.2 51.8 op. m. 7 p. m. . 39.8 56.7 822.9 57.1 7 p. m. 9 p. m. . 53.0 71.1 809.1 75.4 9 p. m. 11 p. m. . 53.0 71.3 813.3 68.9 11 p. m. 1 a. m. . 34.2 55.1 812.3 50.4 1 a. m. 3 a. m. . 43.5 50.5 803.4 44.8 3 a. m. 5 a. m. . 38.1 52.7 805.2 52.8 5 a. m. 7 a. m. . Total 34.0 59.1 800.2 57.4 767.3 733.8 EXPERIMEIs [T No. 83. H . R. D. 1905. Preliminary. Dec. 4-5 11 p. m 25.8 26.8 49!4 924.6 925.4 45!8 lip. m. to 1 a. m. . 1 a. m. 3 a. m. . 21.7 36.6 942.4 32.4 3 a. m. 5 a. m. . 25.7 40.2 946.1 41.9 Dec. 5-6 5 a. m. 7 a. in . . Total, 8 hours 7 a. m. to 9 a. m. . 23.4 42.6 960.6 32.6 .... 168.8 152.7 33.1 67.8 952.6 76.9 9 a. m. 11 a. m. . 29.7 56.1 961.6 48.1 11 a. m. 1 p. m. . 32.1 58.4 956.5 58.8 1 p. m. 3 p. m. . 24.9 44.8 967.0 35.5 3 p. m. 5 p. m. . 28.4 51.1 970.3 51.8 5 p. m. 7 p. m. . 26.6 45.0 957.6 41.4 7 p.m. 9 p.m.. 30.6 54.4 960.4 61.6 9 p. m. 11 p. m. . 29.0 50.5 963.1 38.4 11 p. m. 1 a. m. . 28.6 48.4 956.1 49.0 1 a. m. 3 a. m. . 26.9 41.3 957.2 36.0 3 a. m. 5 a. m. . 28.2 42.9 946.2 45.5 5 a. m. 7 a. m. . Total 29.7 46.0 945.5 42.2 606.7 585.2 256 Influence of Inanition on Metabolism. Table 167. — Record of carbon dioxide and oxygen — Continued. EXPERIMENT No. 83. H. R. D. (Cont'd). Date. Period. Carbon dioxide. Oxygen. (a) Amount in chamber at end of period. (6) Total weight exhaled by subject. (c) Amount in chamber at end of period. id) Total amount consumed by subject. 1905. Dec. 6-7 7 a. m. to 9 a. m. . 9 a. m. 11 a. m. . 11 a. m. 1p.m.. 1p.m. 3 p. m. . 3 p. m. 5 p. m. . 5 p. m. 7 p. m. . 7 p. m. 9 p. m. . 9 p.m. 11 p. m. . 11 p. m. 1 a. m. . 1 a. m. 3 a. m. . 3 a. m. 5 a. m. . 5 a. m. 7 a. m. . Total Grams. 41.6 34.8 38.3 34.6 39.6 35.4 29.0 34.3 23.6 21.6 23.4 26.1 Grams. 67.6 46.4 55.3 44.6 51.6 42.2 56.3 45.2 42.6 38.9 41.5 47.0 Liters. 926.4 941.3 927.1 925.2 923.7 925.0 922.3 918.7 920.5 919.6 913.8 912.4 Grams. 74.2 32.8 59.3 32.0 55.6 38.9 65.4 29.9 36.1 43.9 39.0 47.3 .... 579.2 554.4 EXPERIMENT No. 85. N. M. P. 1905. Dec. 8-9 Dec. 9-10.... Preliminary: 11 p. m 31.2 29.3 28.1 24.9 27.5 58.3 50.2 47.2 51.9 911.6 908.6 912.3 916.8 916.5 53.0 41.7 33.1 47.7 11 p. m. to 1 a. m. . 1 a. m. 3 a. m. . 3 a. m. 5 a. m. . 5 a. m. 7 a. m. . ! 207.6 | 175.5 7 a. m. to 9 a.m . . 9 a. m. 11 a. m. . 11 a. m. 1 p. m. . 1 p. m. 3 p. m. . 3 p. m. 5 p. m. . 5 p. m. 7 p. m. . 7 p.m. 9 p. m. . 9 p. m. 11 p. m. . 11 p. m. 1 a. m. . 1 a. m. 3 a. m. . 3 a. m. 5 a. m. . 5 a. m. 7 a. m. . 37.4 35.7 39.8 33.3 31.0 31.1 34.0 29.4 27.9 26.6 31.5 28.4 78.1 62.4 70.1 55.5 55.4 57.2 65.4 54.7 52.0 46.9 49.9 49.0 904.0 896.1 890.3 888.1 883.1 870.7 854.7 841.8 828.3 808.9 789.4 778.7 69.7 55.8 57.0 47.8 52.1 49.4 67.1 51.1 39.9 46.2 46.0 45.4 Total 696.6 627.5 Metabolism Experiments Nos. 79-83, 85, and 89. 257 Table 167. — Record of carbon dioxide and oxygen — Continued. EXPERIMENT No. 85. N. M. P. (Cont'd). Date. Period. Carbon dioxide. Oxygen. («) Amount in (b) Total (c) Amount in (d) Total chamber weight chamber amount at end of exhaled at end of consumed period. by subject. period. by subject. 1905. Dec. 10-11... 7 a. m. to 9 a. m. . 39.9 77.0 756.6 67.3 9 a. m. 11 a. m. . 34.2 62.2 759.3 50.9 11 a. m. 1 p. m. . 34.3 61.8 753.0 61.9 1 p. m. 3 p. m. . 34.4 61.9 760.2 61.9 3 p. m. 5 p. m. . 34.5 56.6 775.4 50.2 5 p.m. 7 p. m. . 30.4 55.7 789.0 56.9 7 p. m. 9 p. m. . 35.0 67.5 788.8 67.1 9 p. m. 11 p. m. . 30.3 61.5 802.3 59.3 11 p. m. 1 a. m. . 28.9 56.2 801.5 51.2 1 a. m. 3 a. m. . 26.3 50.7 798.5 48.7 3 a. m. 5 a. m. . 28.7 52.9 795.6 50.5 5 a. m. 7 a. m. . Total 28.7 55.3 793.0 49.8 719.3 675.6 EXPERIMI ]NT No. 89. D. W. 1906. Preliminary: Jan. 9-10 11 p. m 42.6 33.4 66.2 938.3 943.3 54!8 11 p. m. to 1 a. m. . 1 a. m. 3 a. m. . 29.7 51.5 946.6 45.9 3 a. m. 5 a. m. . 30.4 55.2 957.1 45.0 Jan. 10-11... 5 a. m. 7 a. m. . Total, 8 hours 7 a. m. to 9 a. m. . 31.2 52.7 957.3 50.9 225.6 196.6 35.9 75.1 965.7 62.4 9 a. m. 11 a. m. . 28.9 57.3 975.4 37.4 11 a. m. 1 p. m. . 28.9 53.2 959.9 46.8 1 p. m. 3 p. m. . 33.8 65.6 950.4 67.1 3 p. m. 5 p.m.. 29.1 54.4 964.6 40.0 5 p.m. 7 p. m. . 30.6 63.5 958.8 68.4 7 p. m. 9 p. m. . 38.4 69.4 953.9 56.3 9 p. m. 11 p. m. . 39.7 74.6 949.8 79.5 11 p. m. 1 a. m. . 33.5 50.3 956.0 38.9 1 a. m. 3 a. m. . 25.6 48.2 966.2 46.0 3 a. m. 5 a. m. . 34.9 56.8 966.6 47.7 Jan. 11-12... 5 a. m. 7 a. m . . Total 29.6 54.0 965.1 54.9 .... 722.4 645.4 7 a. m. to 9 a. m. . 45.8 65.7 964.1 67.3 9 a. m. 11 a. m. . 29.6 54.5 976.7 44.4 11 a. m. 1 p. m. . 43.7 58.6 957.4 52.6 1 p. m. 3 p. m. . 38.8 66.0 951.4 74.7 3 p. m. 5 p. m. . 44.7 50.8 942.0 49.7 5 p. m. 7 p. m. . 36.0 66.9 942.8 74.6 7 p. m. 9 p. m. . 61.4 75.1 922.2 65.7 9 p. m. 11 p. m. . 40.3 67.7 921.2 76.1 11 p. m. 1 a. m. . 54.7 53.0 907.9 40.4 1 a. m. 3 a. m. . 30.5 49.5 917.9 49.8 3 a. m. 5 a. m. . 28.7 50.3 912.3 48.2 5 a. m. 7 a. m. . Total 25.2 47.4 908.2 37.8 705.5 681.3 17 258 Influence of Inanition on Metabolism. Table 168. — Elements Jcatabolized in body — Metabolism experiments Nos. 79-83, 85, and 89. (a) Total weight. (6) Nitro- gen. (c) Carbon. (d) Hydro- gen. (e) Oxygen. (/) Ash. Experiment No. 79. H.E.S. First day, Oct. 13, 1905. Income: Oxygen from air. . . . Outgo: Grams. 575.90 Grams. Grams. Grams. Grams. 575.90 Grams. 996.08 31.12 666.98 632.02 siii 6.27 172.38 111.46 1.75 74.64 884.62 6.26 592 . 34 459.64 8*. 73 Total 2326.20 1750.30 8.11 8.11 178.65 178.65 187.85 187.85 1942.86 1366.96 8. 8. 73 73 Second day, Oct. 14, 1905. Income : Oxygen from air. . . . Outgo : Water in urine 605.27 ■ • • • .... .... 605.27 • • ■ ■ 809.96 49.14 704.07 635.18 14.35 10.22 173.24 90.63 3.09 78.79 719.33 10.31 625.28 461.94 11. 17 Total 2198.35 1593.08 14.35 14.35 183.46 183.46 172.51 172.51 1816.86 1211.59 11 11 17 17 Experiment No. 80. C.R.Y. First day, Oct. 21, 1905. Income : Oxygen from air. . . . Outgo : 576.21 576.21 1105.75 41.65 926.76 627.38 7.78 7.11 171.11 123.73 2.07 103.70 982.02 5.76 823.06 456 . 27 18 93 Solids in urine Total 2701.54 2125.33 7.78 7.78 178.22 178.22 229.50 229 . 50 2267.11 1690.90 18 18 93 93 Second day, Oct. 28, 1905. Income : Oxygen from air. . . . Outgo : 628.55 62S.55 * • • ■ 734.80 42.60 1060.53 640.27 9.95 9.02 174 63 82.22 2.57 118.67 652.58 8.54 941.86 465.64 12 52 Solids in urine Water of respiration ' Total 2478.20 1849.65 9.95 9.95 183.65 183.65 203.46 203.46 2068.62 1440.07 12 12 52 52 Experiment No. 81. A.H.M. First day, Nov. 21, 1905. Income : Oxygen from air. . . . Outgo : 516.78 516.78 621.26 38.74 608.59 534.68 9.11 7.52 145.82 69.52 2.18 68.10 139.80 139.80 551.74 7.32 540.49 388.86 12 61 Carbon dioxide Total 1803.27 1286.49 9.11 9.11 153.34 153.34 1488.41 971.63 12 12 .61 61 1 Includes also water of perspiration. Metabolism Experiments Nos. 79-83, 85, and 89. 259 Table 168. — Elements katabolized in body — Continued. (a) Total weight. (6) Nitro- gen. (c) Carbon. (d) Hydro- gen. (e) Oxygen. (/) Ash. Experiment 81. A.H.M. (cont.) Second day, Nov. 22, 1905. Outgo : Grams. 527.06 Grams. Grams. Grams. Grams. 527.06 Grams. 783.16 48.04 670 . 74 524.33 13.05 9* 48 142 '. 98 87.64 2.91 75.06 695.52 11.38 595.68 381.35 11.22 Total 2026.27 1499.21 13.05 13.05 152.46 152.46 165.61 165.61 1683.93 1156.87 11.22 11.22 Experiment No. 82. H.C.K. First day, Nov. 24, 1905. Outgo : 663.26 663.26 538.57 37.13 842.10 740.87 9.38 7'71 202.04 60.27 2.07 94.23 478.30 8.30 747.87 538.83 9.67 Total 2158.67 1495.41 9.38 9.38 209.75 209.75 156.57 1 56 . 57 1773.30 1110.04 9.67 9.67 Loss Second day, Nov. 25, 1905. Outgo : Water in urine 733.77 .... .... 733.77 • • • * 1740.07 54.93 940.50 767.26 14.36 10.77 209.25 194.71 3.59 105.24 1545.36 9.88 835.26 558.01 16.33 Total 3502.76 2768.99 14.36 14.36 220.02 220.02 303.54 303.54 2948.51 2214.74 16.33 16.33 Experiment No. 83. H.R.D. First day, Dec. 5, 1905. Income : Oxygen from air Outgo : 585.22 585.22 1159.21 45.29 684.52 606.70 13*25 9.76 165.47 129.72 2.65 76.60 1029.49 11.44 607.92 441.23 8.19 Total 2495.72 1910.50 13.25 13.25 175.23 175.23 208.97 208.97 2090.08 1504.86 8.19 8.19 Second day, Dec. 6, 1905. Outgo : 554.37 .... • • • • 554.37 • • • * 1012.39 52.61 672.01 579.17 13.53 12.78 157.95 113.29 3.20 75.20 899 . 10 14.37 596.81 421.22 8.73 Solids in urine Carbon dioxide Total 2316.18 1761.81 13.53 13.53 170.73 170.73 191.69 191.69 1931.50 1377.13 8.73 8.73 1 Includes also water of perspiration. 260 Influence of Inanition on Metabolism. Table 168. — Elements katabolized in oody- -Continued. (a) Total weight. (b) Nitro- gen. (c) Carbon. (d) Hydro- gen. (e) Oxygen. (f) Ash. Experiment No. 85. N.M.P. First day, Dec. 9, 1905. Income : Oxygen from air. . . . Grams. 627.54 Grams. Grams. Grams. Grams. 627.54 Grams. Outjro : U178.48 2 44.14 775.79 696.59 2695.00 2067.46 » 11.37 2 7. 70 189.96 2131.87 2 1.96 86.81 '1046.61 2 10.28 688.98 506.63 2 12.83 Water of respiration l Carbon dioxide Total 11.37 11.37 197.66 197.66 220.64 220.64 2252.50 1624.96 12.83 12.83 Second day, Dec. 10, 1905. Income : Oxygen from air. . . . 675.57 ■ • 657.57 . . . • Outgo: 642.44 36.96 813.43 719.33 11.35 8.22 196.18 71.89 2.11 91.02 570.55 9.71 722.41 523.15 5.57 Carbon dioxide Total 2212.16 1536.59 11.35 11.35 204.40 204.40 165.02 165.02 1825.82 1150.25 5.57 5.57 Experiment No. 89. D.W. First day, Jan. 10, 1906. Income : Oxygen from air. . . . Outgo : 645.36 645 . 36 599.55 44.85 819.57 722.40 9.99 8.38 197.01 67.09 2.26 91.71 532.46 10.88 727.86 525.39 13.34 Total 2186.37 1541.01 9.99 9.99 205.39 205.39 161.06 161.06 1796.59 1151.23 13.34 13.34 Second day, Jan. 11, 1906. Income: Oxygen from air. . . . Outgo : 681.32 1 .... 681.32 • ■ . • 492.22 47.38 802.74 705 . 54 14.46 9.87 192.43 55.08 2.81 89.83 437.14 12.79 712.91 513.11 7.45 Solids in urine Total 2047.88 1366.56 14.46 14.46 202.30 202.30 147 . 72 147.72 1675.95 994.63 7.45 7.45 1 Includes also water of perspiration. - Includes estimated portion in urine lost. See p. 243. 8 Includes 0.32 gram nitrogen determined in urine lost. Materials katabolized in body. — Table 169 shows the amounts of body protein, fat, and glycogen derived by formulae from the quantities of the elements katabolized as shown in table 168. Balance of water. — The amounts of preformed water lost from the body by the several subjects of these experiments, together with the water of oxidation of organic hydrogen are shown in table 170. From this table the sources of intake and output of water may also be found. The method of obtaining the data has already been explained. Metabolism Experiments Nos. 79-83, 85, and 89. 261 Table 169. — Materials katabolized in body — Metabolism experiments Nos. 79-83, 85, and 89. Experiment number, subject, and date. (a) Water. (b) Protein. (c) Fat. (d) Carbohy- drates (as glycogen). (e) Ash. Experiment No. 79. Oct. 13-14, 1905. . Oct 14 15 1905. . H. E. S. : Grams. 1443.52 1299.01 1815.23 1571.11 1044.41 1275.44 1124.60 2419.54 1635.07 1487.61 1714.01 1204.32 1171.22 1051.44 Orams. 48.66 86.10 46.68 59.70 54.66 78.30 56.28 86.16 79.50 81.18 68.22 68.10 59.94 86.76 Orams. 133.58 158.17 141.58 190.09 146.93 161.17 140.10 303.56 156.18 143.93 127.43 168.00 131.80 183.61 Orams. 117.55 39.95 103.62 17.14 28.72 1 25.72 165.56 44.73 32.78 41.60 146.02 91.64 165.63 39.63 Orams. 8.73 11.17 18.93 13.52 12.61 11.22 9.67 16.33 8.19 8.73 12.83 5.57 13.34 7.45 Experiment No. 80. Oct 37 28 1905. C. R. Y. : Oct. 38-29, 1905. Experiment No. 81. Nov. 21-22, 1905 Nov. 22-23, 1905. Experiment No. 82. Nov. 24-25, 1905. Nov. 25-26, 1905. Experiment No. 83. Dec 5 6, 1905 . A. H. M. : H. C. K. : H. R. D. : Dec 6 7 1905. . - Experiment No. 85. Dec. 9-10, 1905 Dec. 10 11, 1905 Experiment No. 89. Jan. 10-11, 1906. Jan. 11-12, 1906. N. M. P. : i D. W. : 1 Glycogen gained. a Amounts for first day, Dec. 9-10, are calculated, with elements of estimated urine lost included. See p. 243. Table 170. — Distribution of intake and outgo of water — Metabolism experiments Nos. 79-83, 85, and 89. Experiment number, subject, and date. Outgo from the body. Balance of preformed water. (a) Water of oxida- tion of organic hydro- gen (c-d). (a) Water of urine. (b) Water of respira- tion and perspira- tion. (c) Total (a+o). (d) Pre- formed (katabo- lized) water in outgo. (e) Intake in drink. (/) Loss of pre- formed water Ul-e). No. 79. H.E.S. : Orams. \ Orams. Orams. Orams. Orams. Orams. Grams. Oct. 13-14, 1905 996.1 667.0 1663.1 1443.5 782.5 661.0 219.6 809.9 704.1 1514.0 1299.0 339.9 959.1 215.0 No. 80. C.R.Y. : Oct. 27-28, 1905 1105.7 926.8 2033.5 1815.2 132 . 8 1682.4 217.3 734.8 1060.5 1795.3 1571.1 206.4 1364.7 224.2 No. 81. A.H.M. : 621.3 608.6 1229.9 1044.4 291.1 753.3 185.5 Nov. 22-23,1905 783.2 670.7 1453.9 1275.4 193.8 1081.6 178.5 No. 82. H.C.K. : Nov. 24-25,1905 538.6 842.1 1380.7 1124.7 857.8 266.9 256.0 Nov. 25-26,1905 1740.1 940.5 2680.6 l2419.5 1092.9 11326.6 261.1 No. 83. H.R.D. : 1159.2 684.5 1843.7 1635.1 1467.1 168.0 208.6 1012.4 672.0 1684.4 1487.6 884.2 603.4 196.8 No. 85. N.M.P. : Dec. 9-10, 1905.... 1178.5 775 . 8 1954.3 1714.0 704.5 1009.5 240.3 Dec. 10-11, 1905 642.4 813.4 1455.8 1204.3 707.7 496.6 251.5 No. 89. D.W. : Jan. 10-11, 1906 599.5 819.6 1419.1 1171.2 115.1 1056.1 247.9 Jan. 11-12, 1906 492.2 802.7 1294.9 1051.4 357.2 694.2 243.5 1 Does not include water of feces passed on this day. See p. 120. 362 Influence of Inanition on Metabolism. Changes in Body-Weight Compared with Balance of Income and Outgo. In table 171 the changes of body-weight obtained by weighing the subjects each morning are compared with the losses of body material, i. e., the deter- mined weights of the different factors of the outgo less those of the income. Although the comparisons in table 171 do not show exact balances, neverthe- less, they are in general fairly satisfactory and for the most part there is a Table 171. — Comparison of changes in tody weight with balance of income and outgo — Metabolism experiments Nos. 79-83, 86, and 89. Experiment number, subject, and date. Income. Outgo. (ft) Loss of body material (C-fif). (i) Loss of body- weight. (a) Water con- sumed. (6) Oxy- gen. (c) Total (a+W. (d) Urine.1 (e) Carbon dioxide. (f) Water of respi- ration and perspi- ration. (g) Total (d+e+f). No. 79. H.E.8.: Oct. 13-14,1905. Oct. 14-15, 1905. No. 80. C.R.Y.: Oct. 27-28,1905. Oct. 28-29, 1905. No. 81. A.H.M.: Nov. 21-22,1905. Nov. 22-23,1905. No. 82. H.C.K.: Nov. 24-25,1905. Nov. 25-26,1905. No. 83. H.R.D.: Dec. 5-6, 1905.. Dec. 6-7, 1905.. No. 85. N.M.P.: Dec. 9-10,1905. Dec. 10-11,1905. No. 89. D.W. : Jan. 10-11,1906. Jau. 11-12,1906. Q-rams. 782.50 339.90 132.80 206.40 291 . 10 193.80 857 . 80 1092.90 1467.10 884.20 704.50 707.70 115.10 357.20 Grams. 575.90 605.27 576.21 628.55 516.78 527.06 663.26 733.77 585.22 554.37 627.54 675.57 645.36 681.32 Orams. 1358.40 945.17 709.01 834.95 807.88 720.86 1521.06 1826.67 2052.32 1438.57 1332.04 1383.27 760.46 1038.52 Orams. 1018.10 883.90 879.20 1103.80 621 . 70 787.90 559.40 1632.20 1295.80 1070.90 U379.72 670.80 749.05 494.70 Orams. 632.02 635.18 627.38 640.27 534.68 524.33 740.87 767.26 606.70 579.17 696.59 719.33 722.40 705.54 Orams. 666.98 704.07 926.76 1060.53 608.59 670.74 842.10 940.50 684.52 672 . 01 775.79 813.43 819.57 802.74 Orams. 2317.10 2223.15 2433.34 2804.60 1764.97 1982.97 2142.37 2 3339.90 2587.02 2322.08 2852.10 2203.56 2291.02 2002.98 Grams. 958.70 1277.98 1724.33 1969.65 957.09 1262.11 621.31 1513.29 534.70 883.51 1520.06 820.29 1530.56 964.46 Orams. 1089.00 1258.00 1725.00 1935.00 969.00 1237.00 605.00 1528.00 539.00 864.00 1498.00 830.00 1502.00 980.00 1 The data in this column should not be confounded with urine data in other tables. See explanation, p. 66. 2 On Nov. 25, 118.0 grams of feces were excreted. This amount Is not Included in the total outgo. See p. 120. 3 Includes 34.62 grams urine calculated as lost. See p. 243. tendency for the errors in the individual days of each experiment to compensate each other. A particularly poor balance was obtained on the first day of experiment No. 79, the reason for which is not known. The average error in terms of the calculated loss is for experiment No. 79, -{-55 grams or -f-4.9 per cent of the average daily loss; for experiment No. 80, — 17 grams or — 0.9 per cent; for experiment No. 81, — 7 grams or — 0.6 per cent; for experiment No. 82, — 1 gram or — 0.1 per cent; for exp. No. 83, — 8 grams, or — 1.1 per cent; for exp. No. 85, — 6 grams or — 0.5 per cent; for exp. No. 89, — 7 grams or — 0.6 per cent. The wide variation in exp. No. 79 is of course due to the large error, 130 grams, on the first day. In all but two experiments the larger loss of body-weight occurred on the second day. Metabolism Experiments Nos. 79-83, 85, and 89. 263 Output of Heat. The total heat production per period and per day for this series of experi- ments is shown in column d of table 172. Columns a, b, and c contain the data essential to obtaining column d. Table 172. — Summary of calorimetric measurements and total heat production — Metabolism experiments Nos. 79-83, 85, and 89. Experiment number, subject, and date. Period. (a)! Heat measured in terms C20- (6) Heat used in vaporiza- tion of water. (c)j Sum of heat correc- tions.1 Total heat pro- duction (a + b + c). 79. H. E. S. 1905. Calories. Calories. Calories. Calories. Oct. 13-14.. 7 a. m. to 9 a. m. . 158.8 32.2 -17.9 173.1 9 a. m. 11 a. m. . 130.0 33.7 + .5 164.2 11 a. m. 1 p. m. . 136.9 33.7 + 18.3 188.9 1 p. m. 3 p. m. . 136.9 33.8 -15.5 155.2 3 p.m. 5 p. m. . 113.7 33.3 + 27.1 174.1 5 p. m. 7 p. m. . 117.6 33.8 - 8.7 142.7 7 p. m. 9 p. m. . 114.4 32.1 + 2.3 148.8 9 p. m. 11 p. m. . 136.6 33.0 - 2.4 167.2 11 p. m. 1 a. m. . 109.7 32.9 + 34.0 176.6 1 a. m. 3 a. m. . 139.2 38.1 -19.1 158.2 3 a. m. 5 a. m. . 101.9 32.9 + 12.9 147.7 Oct. 14-15.. 5 a. m. 7 a. Total m. . 117.0 35.4 + 2.3 154.7 1512.7 404.9 + 33.8 1951.4 7 a. m. to 9 a. m. . 199.1 33.0 -44.6 187.5 9 a. m. 11 a. m. . 155.9 34.4 -27.2 163.1 11a. m. 1 p. m. . 125.9 38.2 + 16.7 180.8 1 p. m. 3 p. m. . 141.2 34.6 + 3.3 179.1 3 p. m. 5 p. m. . 125.1 35.2 - 1.4 158.9 5 p. m. 7 p. m. . 138.0 35.8 + 13.2 187.0 7 p. m. 9 p. m. . 133.8 36.6 + 8.3 178.7 9 p. m. 11 p. m. . 120.9 36.7 -21.4 136.2 11 p. m. 1 a. m. . 119.1 35.7 + 34.3 189.1 1 a. m. 3 a. m. . 141.3 36.4 - 3.0 174.7 3 a. m. 5 a. m. . 134.1 32.4 -16.1 150.4 80. C. R. Y. 5 a. m. 7 a. Total m. . 118.3 31.8 + 11.4 161.5 1652.7 420.8 -26.5 2047.0 Preliminary: Oct. 26-27.. 11 p. m. to 1 a. m. . 101.2 39.1 2+12.6 2 152.9 1 a. m. 3 a. m. . 99.6 49.3 .... 2 148.9 3 a. m. 5 a. m. . 96.0 45.5 2 + 8.4 2 149.9 Oct. 27-28.. 5 a. m. 7 a. Total, 8 hours. . m. . 91.2 44.8 2-24.0 2112.0 388.0 178.7 2- 3.0 2 563.7 7 a. m. to 9 a. m. . 191.2 40.6 -17.5 214.3 9 a. m. 11 a. m. . 161.8 29.0 -19.3 171.5 11 a. m. 1 p. m. . 113.7 50.7 -22.3 142.1 1 p. rn. 3 p. m. . 92.9 44.9 - 4.4 133.4 3 p. m. 5 p. m. . 97.2 45.7 + 38.6 181.5 5 p. m. 7 p. m. . 100.1 45.9 -11.7 134.3 7 p. m. 9 p. m. . 110.1 42.4 - 5.5 147.0 9 p. m. 11 p. m. . 109.3 42.9 + 1.7 153.9 11 p. rn. 1 a. m. . 107.0 44.3 + 41.3 192.6 1 a. m. 3 a. m. . 101.7 57.2 - 3.1 155.8 3 a. m. 5 a. m. . 99.0 53.3 + 9.1 161.4 5 a. m. 7 a. Total m. . 97.6 62.3 + 6.4 166.3 1381.6 559.2 + 13.3 1954.1 1 See pp. 42-49. 2 Does not include correction for change in body temperature. 264 Influence of Inanition on Metabolism. Table 172. — Summary of calorimetric measurements and total heat production- Continued. Experiment number, subject, and date. Period. (o) Heat measured in terms C20- (b) Heat used in vaporiza- tion of water. (c) Sum of heat correc- tions.1 («0 Total heat pro- duction (a + b + c). 80. C.R.Y.(coat. 1905. Calories. Calories. Calories. Calories. Oct. 28-29.. 7 a. m. to 9 a. m. . 193.2 54.6 + 33.0 280.8 9 a. m. 11 a. m. . 145.9 45.8 -20.1 171.6 11 a. m. 1 p. m. . 97.4 52.1 - 9.6 139.9 1 p. m. 3 p. m. . 101.4 56.4 + 12.0 169.8 3 p. m. 5 p. m. . 108.3 53.0 + 13.8 175.1 5 p. m. 7 p. m. . 132.4 54.4 - 0.3 186.5 7 p. m. 9 p. m. . 116.8 49.1 -11.2 154.7 9 p. m. 11 p. m. . 118.4 49.3 + 2.6 170.3 11 p. m. 1 a. m. . 113.5 56.4 - 8.9 161.0 1 a. m. 3 a. m. . 105.2 55.0 - 6.3 153.9 3 a. m. 5 a. m. . 121.1 50.4 -11.5 160.0 81. A. H. M. 5 a. m. 7 a. Total m. . 115.5 51.4 + 9.0 175.9 1469.1 627.9 + 2.5 2099 . 5 Preliminary: Nov. 20-21.. 1 a. m. to 3 a. m. . 70.3 35.5 2- 3.6 2 105.2 3 a. m. 5 a. m. . 69.3 33.0 2 + 6.0 2 108.2 Nov. 21-22. . 5 a. m. 7 a. Total, 6 hours . . m. . 65.9 31.9 2+ 9.6 2 107.5 208.5 100.4 2+12.0 2 320 . 9 7 a. m. to 9 a. in. . 150.4 35.6 - 3.8 182.2 9 a. m. 11 a. m. . 135.2 39.9 -25.4 149.7 11a. m. 1 p. m. 130.0 34.7 -10.7 154.0 1 p. m. 3 p. m. . 110.2 33.0 + 43.9 187.1 3 p. m. 5 p. m. . 130.6 32.9 -23.5 140.1 5 p. m. 7 p. m. . 120.3 33.5 + 10.7 164.5 7 p. m. 9 p. m. . 119.5 31.0 -10.2 140.3 9 p. m. 11 p. m. . 122.6 28.2 -17.3 133.5 11 p. m. 1 a. m. . 65.0 29.9 + 27.1 122.0 1 a. m. 3 a. m. . 70.4 29.3 + 2.8 102.5 3 a. m. 5 a. m. . 84.9 29.1 -12.4 101.7 Nov. 22-23.. 5 a. m. 7 a. Total 7 a. m. to 9 a. m. . m. . 72.3 31.4 + 47.2 150.9 1311.4 388.7 + 28.4 1728.5 171.0 33.7 -26.1 178.6 9 a. m. 11 a. m. . 154.1 38.3 - 7.7 184.7 11 a. m. 1 p. m. . 149.8 32.8 - 5.8 176.8 l.p. m. 3 p. m. . 89.7 30.4 - 8.5 111.6 3 p. m. 5 p. m. . 68.9 29.6 + 22.9 121.4 5 p. m. 7 p. m. . 126.2 34.7 + 18.5 179.3 7 p. m. 9 p. m. . 106.5 33.6 -23.7 116.5 9 p. m. 11 p. m. . 120.6 34.3 + 2.7 157.7 11 p. m. la. m. . 97.3 32.9 + 23.3 153.6 1 a. m. 3 a. m. . 93.8 33.2 + 5.8 132.8 3 a. m. 5 a. m. . 93.0 32.6 - 4.5 121.1 5 a. m. 7 a. Total m. . 99.0 33.7 + 13.9 146.6 1369.9 399.8 + 10.8 1780.5 1 See pp. 42-49. 2 Does not include correction for change in body temperature. Metabolism Experiments JSTos. 79-83, 85, and 89. 265 Table 172.- -Summary of calorimetric measurements and total Jieat production — Continued. Experiment number, subject, and date. Period. (a) Heat measured in terms C20- i (6) Heat used in vaporiza- tion of water. (c) 1 Sum of heat correc- tions.1 (d) Total heat pro- duction (a+b + c). 82. H. C. K. 1905. Preliminary: Calories. Calories. Calories. Calories. Nov. 23-24.. 7 p. m. to 9 p. m. . 168.9 43.7 2- 6.0 2 206.6 9 p. m. 11 p. m. . 168.8 40.0 2- 1.2 2 207.6 11 p. m. 1 a. m. . 117.0 41.2 2+11.4 2 169.6 1 a. m. 3 a. m. . 89.3 38.2 2- 6.0 2 121.5 3 a. m. 5 a. m. . 93.7 38.6 2+ 6.0 2 138.3 Nov. 24-25. . 5 a. m. 7 a. m. . Total, 12 hours 7 a. m. to 9 a. m. . 116.2 45.6 2- 5.4 2 156.4 753.9 247.3 2- 1.2 2 1000.0 210.7 59.3 -19.8 250.2 9 a. m. 11 a. m. . 167.6 52.0 + 21.8 241 A 11 a. m. 1p.m.. 165.8 35.9 -51.2 150.5 1 p. m. 3 p. m. . 164.8 43.6 + 7.4 215.8 3 p.m. 5 p. m. . 111.4 44.2 - 9.5 146.1 5 p. m. 7 p. m. . 117.7 40.5 + 7.7 165.9 7 p. m. 9 p. m. . 172.1 42.3 + 10.5 224.9 9 p. m. 11 p. m. . 169.0 39.2 - 8.6 199.6 11 p. m. 1 a. m. . 101.0 38.9 + 20.3 160.2 1 a. m. 3 a. m. . 122.2 30.3 - 7.2 145.3 3 a. m. 5 a. m. . 105.8 38.1 + 20.3 164.2 Nov. 25-26.. 5 a. m. 7 a. m. . Total 106.6 40.2 + 10.7 157.5 1714.7 504.5 + 2.4 2221.6 7 a. m. to 9 a. m. . 235.6 64.1 -28.9 270.8 9 a. m. 11 a. m. . 171.4 49.6 -12.7 208.3 11 a. m. 1 p. m. . 165.9 45.2 + 18.9 230.0 1 p. m. 3 p.m.. 161.0 43.7 - 7.9 196.8 3 p. m. 5 p. m. . 135.5 42.8 + 10.2 188.5 5 p. m. 7 p.m.. 173.3 42.6 + 6.2 222.1 7 p. m. 9 p.m.. 192.2 48.7 + 15.8 256.7 9 p. m. 11 p. m. . 191.0 46.3 -12.0 225.3 - 11 p. m. 1 a. m. . 108.2 42.0 + 2.5 152.7 1 a. m. 3 a. m. . 111.7 33.0 + 18.2 162.9 3 a. m. 5 a. m . . 130.4 40.0 + 3.8 174.2 83. H. R. D. 5 a. m. 7 a. m. . Total 128.7 44.6 + 15.6 188.9 1904.9 542.6 + 29.7 2477.2 Preliminary : Dec. 5 11 p. m. to 1 a. m. . 94.9 45.7 2- 3.6 2 137.0 1 a. m. 3 a. m. . 89.7 43.2 2- 4.8 2 128.1 3 a. m. 5 a. m. . 85.2 36.9 2+ 1.2 2 123.3 5 a. m. 7 a. m. . 80.6 35.7 2+ 2.4 2 118.7 350.4 161.5 2- 4.8 2 507.1 1 See pp. 42-49. 2 Does not include correction for change in body temperature. 266 Influence of Inanition on Metabolism. Table 172. — Summary of calorimetric measurements and total heat production — Continued. Experiment (a) (6) (c) (d) number, subject, and date. Period. Heat measured in terms C20- Heat used in vaporiza- tion of water. Sum of heat correc- tions.1 Total heat pro- duction (a+b+c). 83. H.R.D. (cont. 1905. Calories. Calories. Calories. Calories. Dec. 5-6. . . 7 a. m. to 9 a. m. . 174.4 42.5 - 7.4 209.5 9 a. m. 11 a. m. . 131.4 36.2 + 4.1 171.7 11 a. m. 1 p. m. . 159.4 36.5 -11.3 184.6 1 p. m. 3 p. m. . 106.9 33.7 -12.9 127.7 3 p. m. 5 p. in. . 120.1 32.8 - 6.4 146.5 5 p. m. 7 p. m. . 68.0 38.0 + 6.6 112.6 7 p. m. 9 p. m. . 176.0 36.9 + 10.0 222.9 9 p. m. 11 p. m. . 119.7 33.4 - 2.9 150.2 11 p. m. 1 a. m. . 97.8 33.7 + 31.9 163.4 1 a. m. 3 a. m. . 101.2 38.4 - 5.5 134.1 3 a. m. 5 a. m. . 88.7 33.7 + 14.5 136.9 Dec. 6-7... 5 a. m. 7 a. Total m. . 106.2 35.5 + 11.8 153.5 1449.8 431.3 + 32.5 1913.6 7 a. m. to 9 a. m. . 201.4 47.3 -28.0 220.7 9 a. m. 11 a. m. . 104.0 31.8 - 5.9 129.9 11 a. m. 1 p. ni. . 186.0 36.1 + 13.2 235.3 1 p. m. 3 p. m. . 83.6 31.6 - 1.7 113.5 3 p. m. 5 p. m. . 129.3 33.0 + 6.1 168.4 5 p. m. 7 p. m. . 103.7 32.5 - 7.7 128.5 7 p. m. 9 p. m. . 182.2 35.0 -14.8 202.4 9 p. m. 11 p. m. . 90.6 32.4 -10.5 112.5 11 p. m. la. m. . 98.7 33.4 + 31.8 163.9 1 a. m. 3 a. m. . 114.9 31.3 - 5.0 141.2 3 a. m. 5 a. m. . 85.6 29.1 + 13.0 127.7 85. N. M. P. 5 a. m. 7 a. Total m. . 116.3 32.0 + 14.5 162.8 1496.3 405.5 + 5.0 1906.8 Preliminary: Dec. 9 11 p. m. to 1 a. m. . 123.8 41.5 2+ 6.0 2171.3 1 a. m. 3 a. m. . 134.5 36.0 2- 5.4 2 165.1 3 a. m. 5 a. m. . 99.2 34.9 2+ 7.8 2141.9 Dec. 9-10.. 5 a. m. 7 a. Total, 8 hours . 7 a. m. to 9 a. m. . 89.7 34.2 2+ 3.0 2 126,9 447.2 146.6 2+11.4 2 605.2 m. . 202.6 43.6 -28.6 217.6 9 a. in. 11 a. m. . 158.3 38.6 - 0.4 196.5 11 a. m. 1 p. m. . 156.1 37.5 -13.4 180.2 1 p. m. 3 p. m. . 112.0 33.4 + 7.4 152.8 3 p. m. 5 p. m. . 132.7 33.4 + 26.5 192.6 5 p. m. 7 p. m. . 121.6 36.8 + 1.6 160.0 7 p. m. 9 p. m. . 180.1 41.7 + 4.9 226.7 9 p. m. 11 p. m. . 150.8 34.4 -23.2 162.0 11 p. m. 1 a. m. . 93.8 35.3 + 16.6 145.7 1 a. m. 3 a. m. . 131.6 33.7 - 7.7 157.6 3 a. m. 5 a. in. . 104.5 33.6 + 19.7 157.8 5 a. m. 7 a. Total m. . 104.3 34.8 + 20.2 159.3 1648.4 436.8 + 23.6 2108.8 1 See pp. 42-49. ; Does not include correction for change in body temperature. Metabolism Experiments Nos. 79-83, 85, and 89. 267 Table 172. — Summary of calorimetric measurements and total heat production — Continued. Experiment (a) (&) (c) (d) number, subject, and date. Period. Heat measured in terms C20- Heat used in vaporiza- tion of water. Sum of heat correc- tions.1 Total heat pro- duction (a + b + c). 85. N.M.P.(cont.) 1905. Calories. Calories. Calories. Calories. Dec. 10-11.. 7 a. m. to 9 a. m. . 216.0 44.0 -29.0 231.0 9 a. m. 11 a. ni. . 162.8 37.9 - 0.5 200.2 11 a. m. 1 p. m. . 161.5 39.2 + 6.5 207.2 1 p. m. 3 p. ni. . 150.6 36.6 + 11.6 198.8 3 p. m. 5 p. m. . 110.1 35.8 + 20.6 166.5 5 p. m. 7 p. m. . 144.5 41.0 + 9.1 194.6 7 p. m. 9 p. m. . 185.4 41.9 + 3.6 230.9 9 p. m. 11 p. m. . 158.4 41.1 -15.3 184.2 11 p. m. 1 a. m. . 109.0 40.6 + 27.9 177.5 1 a. m. 3 a. m. . 132.4 43.5 - 2.8 173.1 3 a. m. 5 a. m. . 114.3 40.5 + 8.0 162.8 89. D. W. 1906. 5 a. m. 7 a. Total m. . 127.7 43.6 + 6.6 177.9 1772.7 485.7 + 46.3 2304.7 Preliminary: Jan. 9 11 p. ni. to 1 a. m. . 102.7 44.6 2+10.8 2158.1 1 a. m. 3 a. m. . 128.9 44.4 2 + 3.6 2 177.0 3 a. ni. 5 a. m. . 121.8 46.1 2-10.8 2 157.1 Jan. 10-11.. 5 a. m. 7 a. Total, 8 hours . 7 a. m. to 9 a. m. . 101.8 42.4 2+ 7.2 2151.4 455.3 177.5 2+10.8 2 643.6 m. . 207.8 40.7 -17.3 231.2 9 a. m. 11 a. m. . 100.5 38.5 -28.9 110.1 11 a. m. 1 p. m. . 97.4 39.6 + 2.2 139.2 1 p. m. 3 p. m. . 186.5 42.5 + 27.0 256.0 3 p. m. 5 p. in. . 89.9 38.7 -19.9 108.7 5 p. m. 7 p. m. . 180.0 42.6 + 19.0 241.6 7 p.m. 9 p. in. . 141.4 43.4 - 2.6 182.2 9 p. m. 11 p. m. . 211.1 43.8 -18.0 236.9 11 p. m. 1 a. ni. . 91.3 41.4 + 19.5 152.2 1 a. m. 3 a. m. . 134.2 38.1 -18.3 154.0 3 a. m. 5 a. m. . 117.8 40.7 + 16.5 175.0 Jan. 11-12.. 5 a. m. 7 a. Total m. . 118.6 40.8 + 3.6 163.0 1676.5 490.8 -17.2 2150.1 7 a. m. to 9 a. m. . 204.9 38.9 + 1.3 245.1 9 a. m. 11 a. m. . 101.5 36.2 -18.6 119.1 11 a. m. 1 p. ni. . 118.4 40.5 + 7.3 166.2 1 p. m. 3 p. m. . 208.7 38.2 + 17.5 264.4 3 p. m. 5 p. m. . 87.7 36.7 -13.1 111.3 5 p. m. 7 p. m. . 202.0 43.7 + 9.0 254.7 7 p. m. 9 p. m. . 151.9 47.4 + 1.1 200.4 9 p. m. 11 p. m. . 204.0 41.0 -10.5 234.5 11 p. m. 1 a. m. . 102.3 41.4 + 47.6 191.3 1 a. m. 3 a. m. . 139.9 36.5 -42.8 133.6 3 a. m. 5 a. ni. . 131.2 38.0 - 1.0 168.2 5 a. m. 7 a. Total m. . 108.8 36.9 + 19.8 165.5 1761.3 475.4 + 17.6 2254.3 1 See pp. 42-49. 2 Does not include correction for change in body temperature. 268 Influence of Inanition on Metabolism. Balance of Energy. The comparison between the total heat production as measured and the completed energy of katabolized body material is given in table 173. Table 173. — Comparison of energy derived from katabolized body material ivith total heat production — Metabolism experiments Nos. 79-83, 85, and 89. Experiment number, subject, and date. Energy derived from different sources. From body protein. (a) Energy of protein katabo- li/cd. (6) Poten- tial energy of urine. (o) Net energy (a-b). (d) From body fat. (e) From body glyco- gen. if) Total (c+d+e). Energy from body material greater (+) or less (— ) than output. (g) Total heat pro- duc- tion. (ft) Amount d) Pro- portion (ft-*-flf). No. 79. Oct. 13- Oct. 14- No. 80. Oct. 27- Oct. 28- No. 81. Nov. 21- Nov.22- No. 82. Nov. 24 Nov. 25- No. 83. Dec. 5- Dec. 6- No. 85. Dec. 9- Dec. 10- No. 89. Jan. 10- Jan. 11- H.E.8.: Gals. Cals. Cals. Cals. Cals. Cals. Cals. Cals. -14,1905. 275 72 203 1265 493 1961 1951 + 10 -15,1905. 486 116 370 1509 167 2046 2047 - 1 C.R.Y.: -28,1905. 264 75 189 1351 434 1974 1954 + 20 -29,1905. 337 97 240 1813 72 2125 2099 + 26 A.H.M.: -22,1905. 309 80 229 1402 120 1751 1729 + 22 -23,1905. 442 109 333 1538 U08 1763 1781 -18 H.C.K. : -25,1905. 318 88 230 1337 694 2261 2222 + 39 -26,1905. 487 122 365 1942 187 2494 2477 + 17 H.R.D. : 6, 1905.. 449 116 333 1490 137 1960 1914 + 46 7, 1905. . 459 151 308 1373 174 1855 1907 — 52 N.M.P. : -10,1905. 385 96 289 1216 612 2117 2109 + 8 -11,1905. 385 99 286 1603 384 2273 2305 -32 D.W. : -11,1906. 339 93 246 1257 694 2197 2150 + 47 -12,1906. 490 114 376 1742 166 2284 2254 + 30 Per ct. + 0.5 + 1.0 + 1.2 + 1.3 -1.0 + 1.8 + 0.7 + 2.4 -2.7 + 0.4 -1.4 + 2.2 + 1.3 1 Glycogen gained. The average discrepancy for each experiment is as follows : Experiment No. 79, +5 calories or +0.3 per cent; for experiment No. 80, +23 calories or +1.1 per cent; experiment No. 81, +2 calories or +0.1 per cent; experiment No. 82, +28 calories or +1.2 per cent; experiment No. 83, — 3 calories or — 0.2 per cent; experiment No. 85, — 12 calories or — 0.5 per cent; experi- ment No. 89, +39 calories or +1.8 per cent. Average of 7 experiments +12 calories or +0.5 per cent. Relations between Oxygen Consumption, Cabbon Dioxide Elimination, and Heat Production. The oxygen and carbon dioxide thermal quotients and the respiratory quotients for experiments Nos. 79-83, 85, and 89 are shown in table 174. Metabolism Experiments Nos. 79-83, 85, and 89. 269 Table 174. — Oxygen and carbon dioxide thermal quotients and respiratory quotients — Metabolism experiments Nos. 79-83, 85, and 89. Experiment number, subject, and date. Period. (a) (6) 0 In ft a o o ■ ti a H O (c) (d) a i ,ST 2° so o •a o a> - -. 73 03 «■§ o g O (e) 9-S^ K 79. H.E.S. 1905. Oct. 13-14. Oct. 14-15. 80. C.R.Y. Oct. 27.... Oct. 27-28. 7 a. 9 a. 11 a. lp. 3 p. 5 p. 7p. 9 p. lip. la. 3 a. 5 a. m. to m. m. m. m. m. ni. m. m. m. ni. m. 9 a. 11a. lp. 3 p. 5 p. 7 p. 9p. lip. la. 3 a. 5 a. 7 a. m. m. m m m. m. m. m. m. m. m. in. Total. 7 a. 9 a. 11a. lp. 3 p. op. 7 p. 9 p. lip. la. 3 a. 5 a. m. to m. ra. m. m. m. m. m. m. m. m. m. 9 a. 11 a. lp. 3 p. 5 p. 7 p. 9 p. lip. la. 3 a. 5 a. 7 a. m. m. m. m. m. m. m. m. m. m. m. m. Cals. 173.1 164.2 188.9 155.2 174.1 142.7 148.8 167.2 176.6 158.2 147.7 154.7 Grams. 67.3 42.7 49. 7! 48.5! 47.9 49. 6j 42.0 52.0 41.8 51.31 36.7 46. 41 38.9 26.0 26.3 31.2 27.5 34.7 28.2 31.1 23.7 32.4 24.9 30.0 1951.4 575.9! 29.5 Grams. 67.8 54.2 55 57 50 51 50 51 50.9 52.0 43.4 47.2 632.0 39.2 33.0 29.2 37.0 29.1 36.2 33.9 30.6 28.9 32.9 29.4 30.5 Liters. 34.5 27.6 28.1 29.2 25.8 26.3 25.7 26.1 25.9 26.5 22.1 24.0 Liters. 47.1 29.9 34.8 33.9 33.6 34.7 29.4 36.4 29.2 35.9 25.7 32.5 0.73 .92 .81 .86 .77 .76 .87 .72 .89 .74 .86 .74 32.4 321.8 403.10.80 187.5 163.1 180.7 179.1 158.9 187.0 178.7 136.2 189.1 174.7 150.5 161.5 Total. 11 p. m. to 1 a. m. 3 a. m. 5 a. m. in. m 2047.0 66. 4j 45.2 53.2 55.0 51.5 46.9 52.4 47.8 51.8 48.8 43.4 42.9 605.3 35.4 27.7 29.4 30.7 32.4 25.1 29.3 35.1 27.4 27.9 28.8 26.6 65.7 54.9 54. li 56.3 51.3 54.6 56.8 45.3 51.3 52.1 45.5 47.3 35.1 33.6 29.9 31.5 32.3 29.2 31.8 33.3 27.1 29.8 30.2 29.3 29.6635.2 31.0 33.5 27.9 27.5 28.7 26.1 27.8 28.9 23.1 26.1 26.5 23.1 24.1 323.3 46.5 31.6 37.2 38.5 36.0 32.8 36.7 33.5 36.3 34.2 30.4 30.0 423.7 0.76 0.72 .88 .74 .74 .73 .85 .79 .69 .72 .78 .76 .80 152.9 148.9 la. 3 a. 5 a. m.1 149.9 7a. m.!1 112.0 Total, 8 hours....'1 563. 7 32.2 21.1 61.6 40.3 31.4 22.6 40.4 27.1 47.5 31.9 24.2 28.2 39.1 26.1 50.2 33.5 25.5 27.4 33.0 29.5 52.4 46.8 26.7 23.1 144.7 25.7 211.7 37.6 107.8 101.3 1.39 .86 .93 1.16 1.06 7 a. 9 a. 11a. lp. 3 p. 5 p. 7 p. 9 p. lip. 1 a. 3 a. 5 a. m. m. m. m. m. m. m. m. m. m. m. m. to 9 a. 11a. lp. 3 p. 5 p. 7 p. 9 p. lip. la. 3 a. 5 a. 7 a. m. m. m. m. m. m. m. m. m. m. m. m. Total. 214.3 171.5 142.1 133.4 181.5 134.3 147.0 153.9 192.6 155.8 161.4 166.3 1954.1 70. 81 43.9 46.3 39.2 50.8 44.0 42.2 40.3 55.8 43.7 51.8 47.4 576.2 33.0 25.6 32.6 29.4 28.0 32.7 28.7 26.2 29.0 28.0 32.1 28.5 29.5 73.6 58.7| 55.2 49.0 51.3 46.2 43.3 43.6 58.7, 47.0 50.7 50.0 627.3 34.3 37.5 34.2 29.9 38.9 28.1 36.8 25.0 28.2 26.1 34.4 23.5 29.5 22.0 28.4 22.2 30.5 29.9 30.2 23.9 31.4 25.8 30.1 25.5 32.1 319.4 49.50.76 30.8 .97 32.4 27.5 35.6 30.8 29.6 28.2 39.0 30.6 36.2 33.2 403.4 .87 .91 .73 .76 .75 .79 .77 .78 .71 .77 0.79 1 See p. 122. 270 Influence oe Inanition on Metabolism. Table 174. — Oxygen and carbon dioxide thermal quotients and respiratory quotients — Continued. (a) (6) (c) (d) (e) (/) , ! i S0.C.K.Y.(cout.) 1905. Cals. Grams. Grams. Liters. Liters. Oct. 28-29. 7 a. m. to 9 a. m. 280.8 68.7 24.5! 64.3 22.9 32.7 48.1 0.68 9 a. m. 11 a. m. 171.6 48.1 28.0 55.0 32.0 28.0 33.7 .83 11 a. m. lp. m. 139.9 46.6 33.3 49.5 35.4 25.2 32.6 .77 1 p. m. 3 p. m. 169.8 53.2 31.41 52.0 30.6 26.5 37.3 .71 3 p. m. 5 p. m. 175.1 49.6 28.3 48.3 27.6 24.6 34.7 .71 5 p. m. 7 p. m. 186.5 59.5 31.9 56.5 30.3 28.8 41.7 .69 7 p. m. 9 p. m. 154.7 48.8 31.5 51.6 33.3 26.3 34.1 .77 9 p. m. 11 p. m. 170.3 44.1 25.91 49.3 28.9 25.1 30.9 .81 11 p. m. 1 a. m. 161.0 59.0 36.7 57.0 35.4 29.0 41.3 .70 1 a. m. 3 a. m. 153.9 43.7 28.4 47.7 31.0 24.2 30.6 .79 3 a. m. 5 a. m. 160.0 53.7 33.6! 52.5 32.9 26.7 37.6 .71 81. A.H.M. 5 a. m. 7 a. m. Total 175.9 53.5 30.4 56.6 32.2 28.8 37.4 .77 2099.5 628.5 29.9 640.3 30 . 5 325.9 440.0 0.74 Preliminary: Nov. 21... 1 a. m. to 3 a. m. 1 105.2 28.1 26.7 34.7 33.0 17.7 19.7 0.90 3 a. m. 5 a. m. 1 108.2 30.9 28.6 37.9 35.0 19.3 21.6 .89 Nov.21-22- 5 a. m. 7 a. m. Total, 6 hours... . 7 a. m. to 9 a. m. 1 107.5 32.2 30.0 39.9 37.1 20.3 22.6 .90 1 320.9 91.2 28.4 112.5 35.1 57.3 63.9 0.90 182.2 52.6 28.9 61.6 33.8 31.4 36.8 0.85 9 a. m. 11 a. m. 149.7 52.7 35.2 53.1 35.5 27.0 36.9 .73 11a. m. lp. m. 154.0 41.6 27.0 45.7 29.7 23.3 29.1 .80 1 p. m. 3 p. m. 187.1 42.4 22.7 42.4 22.7 21.6 29.7 .73 3 p. m. 5 p. m. 140.1 46.2 33.0 45.8 32.7 23.3 32.4 .72 5 p. m. 7 p. m. 164.5 45.5 27.7 44.0 26.7 22.4 31.9 .70 7 p. m. 9 p. m. 140.3 44.9 32.0 44.3 31.6 22.5 31.4 .72 9p. m. 11 p. m. 133.5 41.0 30.7 44.4! 33.3 22.6 28.7 .79 11 p. m. 1 a. m. 122.0 36.6 30.0 37.9 31.0 19.3 25.6 .75 1a.m. 3 a. m. 102.5 38.9 38.0 35.5 34.6 18.1 27.2 .66 3 a. m. 5 a. m. 101.7 35.4 34.8 40.1J 39.4 20.4 24.8 .82 Nov. 22-23. 5 a. m. 7 a. m. Total 150.9 38.9 25.7 39.9 26.5 20.3 27.2 .75 1728.5 516.7 29.9 534.7 30.9 272.2 361.7 0.75 7 a. m. to 9 a. m. 178.6 62.3 34.9 59. 71 33.4 30.4 43.6l0.70 9 a. m. 11 a. m. 184.7 50.5 27.3 51.7, 28.0 26.3 35.3 .75 11 a. m. 1 p. m. 176.8 46.7 26.4 49.2, 27.9 25.1 32.7 .77 1 p. m. 3 p. m. 111.6 35.1 31.4 38.3' 34.3 19.5 24.6 .79 3 p. m. 5 p. m. 121.4 33.3 27.4 37.3 30.7 19.0 23.3 .81 5 p. m. 7 p. m. 179.3 52.8 29.4 44.8; 25.0 22.8 36.9 .62 7 p. m. 9 p. m. 116.5 43.1 37.0 46.9 40.3 23.9 30.2 .79 9 p. m. 11 p. m. 157.7 45.9 29.1 39.6 25.1 20.2 32.1 .63 11 p. m. 1 a. m. 153.5 40.8 26.6 41.6 27.1 21.2 28.5 .74 1 a. m. 3 a. m. 132.8 32.1 24.2 33.9 25.5 17.2 22.5 .77 3 a. m. 5 a. m. 121.1 38.8 32.1 36.3 29.9 18.4 27.2 .68 5 a. m. 7 a. m. Total 146.6 45.7 31.2 45. 0! 30.7 22.9 32.0 .72 1780.6 527.1 29.6 524.3 2Q.fi 266.9 368.9 0.72 1 See p. 122. Metabolism Experiments Nos. 79-83, 85, and 89. 271 Table 174. — Oxygen and carbon dioxide thermal quotients and respiratory quotients — Continued. (a) (6) (c) (d) (e) (1) (o) (A) Experiment number, subject, and date. Period. o P. "5 g a> o _, o 03 3 %* i a o «-* a a § O Oxygen thermal quotient (1006 -so). Carbon dioxide eliminated. Carbon dioxide thermal quotient (lOOcN-a). I Volume of car- bon dioxide eliminated WX0.5091). Volume of oxy- gen consumed (6X0.7). Respiratory quotient. 82. H.C.K. 1905. Nov.23-24. Preliminary : 7 p. m. to 9 p. m. Cals. 1 206.6 Grams. 55.3 26.8 Grams. 69.9 33.8 Liters. 35.6 Liters. 38.7 0.92 9 p. m. 11 p. m. 1 207.6 50.8 24.5 62.1 29.9 31.6 35.5 .70 11 p. m. 1 a. m. 1 169.6 47.0 27.7 60.2 35.5 30.7 32.9 .93 1 a. m 3 a. m. 1 121.5 33.9 27.9 46.5 38.2 23.7 23.7 .99 3 a. m. 5 a. m. 1 138.3 46.7 33.8 51.5 37.2 26.2 32.7 .80 Nov.24-25. 5 a. m. 7 a. m. Total, 12 hours... 7 a. m. to 9 a. m. 1 156.4 52.0 33.2 26.4 1 1000.0 .... 342.2| 34.2 174.2 .... ... 250.2 80.2 32.1 82.7 33.1 42.1 56.2 0.75 9 a. m. 11 a. m. 241.4 65.9 27.3 72.1 29.9 36.7 46.2 .80 11 a. m. 1 p. m. 150.5 54.3 36.1 65.8 43.7 33.5 38.0 .88 1 p. m. 3 p. m. 215.8 63.6 29.5 71.3 33.0 36.3 44.5 .82 3 p. m. 5 p. m. 146.1 47.2 32.3 56.0 38.3 28.5 33.0 .86 5 p. m. 7 p. m. 165.9 47.1 28.4 56.7 34.2 28.9 33.0 .88 7 p. m. 9 p. m. 224.9 69.2 30.8 70.4 31.3 35.8 48.4 .74 9 p. m. 11 p. m. 199.6 58.4 29.3 62.9 31.5 32.0 40.9 .78 11 p. m. 1 a. m. 160.2 47.3 29.5 55.8 34.8 28.4 33.1 .86 1 a. m. 3 a. m. 145.3 44.6 30.7 47.7 32.8 24.3 31.2 .78 3 a. m. 5 a. m. 164.2 41.5 25.3 48.3 29.4 24.6 29.1 .85 Nov.25-26. 5 a. m. 7 a. m. Total 157.5 43.9 27.9 51.2 32.5 26.1 30.7 .85 2221.6 663.2 29.9 740.9 33.4 377 9. 464.3 0.81 7 a. m. to 9 a. m. 270.8 85.6 31.6 88.8 32.8 45.2 59.9 0.75 9 a. m. 11 a. m. 208.3 59.4 28.5 63.0 30.3 32.1 41.6 .77 11 a. m. lp. m. 230.0 70.9 30.8 72.0 31.3 36.7 49.6 .74 1 p. m. 3 p. m. 190.8 59.3 30.1 62.8 31.9 31.9 41.5 .77 3 p. m. 5 p. m. 188.5 51.8 27.5 64.2 34.0 32.6 36.3 .90 5 p. m. 7 p. m. 222.1 57.1 25.7 56.7 25.5 28.9 40.0 .72 7 p. m. 9 p. m. 256.7 75.3 29.4 71.1 27.7 36.2 52.7 .69 9 p. m. 11 p. m. 225.3 68.9 30.6 71.3 31.6 36.3 48.2 .75 11 p. m. 1 a. m. 152.7 50.4 33.0 55.1 36.1 28.1 35.3 .80 1 a. m. 3 a. m. 162.9 44.8 27.5 50.5 31.0 25.7 31.4 .82 3 a. m. 5 a. m. 174.2 52.8 30.3 52.7 30.3 26.8 37.0 .73 83. H.R.D. Dec. 4-5. . . 5 a. m. 7 a. m. Total 188.9 57.4 30.4 59.1 31.3 30.1 40.2 .75 2477.2 733.7 20 6 767 3 31.0 300 6 513 7 0.76 Preliminary: 11 p. m. to 1 a. m. '137.0 45.8 33.4 49.3 36.1 25.1 32.10.78 1 a. m. 3 a. m. 1 128.1 32.4 25.3 36.6 28.6 18.6 22.7; .82 3 a. m. 5 a. m. 1 123.3 41.9 34.0 40.2 32.6 20.5 29.3 .70 5 a. m. 7 a. m. Total, 8 hours... . '118.7 32.6 27.5 42.6 35.9 21.7 22.8 .95 '507.1 152.7 30.1 168.7 33.3 85.9 106.9 o.so 1 See p. 122. ■!r! Influence of Inanition on Metabolism. Table 174. — Oxygen and carbon dioxide thermal quotients and respiratory quotients— Continued. (a) (.b) (c) (d) (e) (/) (g) '< (« Experiment o ** - ■ a "3 a . lis is fa °§^' b number, subject, and Period. o3 a V 0 a v ■5 » T '■V g d'd •■3 8.2 + d oj oo o o «m date. - — MB £3 lis -9- oja 30 t. — ^ 3 a d.gx 8 w a -o O a H o o o O > > M « 83. H.B.D.(cont.) 1905. Cats. Grams. Grams. Liters. Liters. Dec. 5-6. . . 7 a. m. to 9 a. m. 209.5 76.9 36.7 67.8 32.4 34.5 53.8 0.64 9 a. m. 11 a. m. 171.7: 48.1 28.0 56.1 32.7 28.6 33.7 .85 11 a. m. 1 p. m. 184.6 58.8 31.9 58.4 31.6 29.7 41.2 .72 1p.m. 3 p. m. 127.7 35.5 27.8 44.8 35.1 22.8 24.8 .92 3 p. m. 5 p. m. 146.5! 51.8 35.3 51.1 34.9 26.0 36.2 .72 5 p. m. 7 p. m. 112.6 41.4 36.8 45.0 40.0 22.9 29.0 .79 7 p. m. 9 p. m. 222.9 61.6 27.7 54.4 24.4 27.7 43.1 .64 9 p. m. 11 p. m. 150.2 38.4 25.6 50.5 33.6 25.7 26.9 .96 11 p. m. 1 a. m. 163.4 49.0 30.0 48.4 29.6 24.7 34.3 .72 1a.m. 3 a. m. 134.1 36.1 26.9 41.3 30.8 21.0 25.2 .83 3 a.m. 5 a. m. 136.9 45.5 33.2 42.9 31.4 21.9 31.9 .69 Dec. 6-7... 5 a.m. 7 a. m. Total 153.5 42.1 27.5 46.0 30.0 23.4 29.5 .79 1913.6 H85 9 30.6 fiflfi 7 31.7 308.9 409.6 n 7K 7 a. m. to 9 a. m. 220.7 74.2 33.61 67.6 30.7 34.4 51.9'0.66 9 a. m. 11 a. m. 129.9 32.8 25.3 46.4 35.7 23.6 23.0 1.03 11 a. m. 1 p. m. 235.3 59.3 25. 2| 55.3 23.5 28.2 41.6! .68 1 p. m. 3 p. m. 113.5 32.0 28.2 44.6 39.3 22.7 22.4 1.01 3 p. m. 5 p. m. 168.4 55.6 33.0 51.6 30.7 26.3 38.9 .68 5 p. m. 7 p. m. 128.5 38.9 30.3 42.2 32.9 21.5 27.2 .79 7 p. m. 9 p. m. 202.4 65.5 32.3 56.3 27.8 28.7 45.8 .63 9 p. m. 11 p. m. 112.5 29.8 26.5 45.2 40.2 23.0 20.9 1.10 11 p. m. 1 a. m. 163.9 36.1 22.1 42.6 26.0 21.7 25.3 .86 1 a. m. 3 a. m. 141.2 43.9 31.1 38.9 27.5 19.8 30.7 .65 3 a. m. 5 a.m. 127.7 39.0 30.5 41.5 32.5 21.1 27.3 .77 85. N.M.P. 5 a. m. 7 a. m. Total Preliminary: 162.8 47.3 29.0 47.0 28.9 23.9 33.1 .72 1906.8 554.4 29.1 579.2 ^30.4 294.9 388.10.76 1 Dec. 8-9... 11 p. m. to 1 a. m. 1 171.3 53.0 30.9 58.3 34.0 29.7 37.1!0.80 1 a. m. 3 a. m. UeS.l 41.7 25.3 50.2 30.4 25.6 29. 2| .88 3 a. m. 5 a. m. 1141.9| 33.1 23.3 47.2 33.3 24.0 23.2 1.04 Dec. 9-10.. 5 a. m. 7 a. m. Total, 8 hours... . 7 a. m. to 9 a. m. 1 126.9 47.7 37.6 51.9 40.9 26.4 33.4 .79 '605.2 175.5 29.0 207.6 34.3 105.7 122.9 0.86 217.7 69.7 32.0 78.1 35.9 -39.8 48.8 0.82 9 a. m. 11 a. m. 196.5 55.8 28.4 62.4 31.7 31.8 39.1 .81 11 a. m. 1 p. m. 180.2 57.0 31.6 70.1 38.9 35.7 39.9 .90 1 p. m. 3 p. m. 152.8 47.8 31.3 55.5 36.3 28.2 33.4 .85 3 p. m. 5 p. m. 192.6 52.1 27.1 55.4 28.8 '28.2 36.5 .77 5 p. m. 7 p. m. 160.0 49.4 30.9 57.2 35.7 29.1 34.6 .84 7 p. m. 9 p. m. 226.7 67.1 29.6 65.4 28.9 33.3 47.0 .71 9 p. m. 11 p. m. 162.0 51.1 31.5 54.7 33.8 27.9 35.8 .78 11 p. m. 1 a. nu 145.7 39.9 27.4 52.0 35.7 26.5 27.9 .95 1 a. m. 3 a. m. 157.5 46.2 29.3 46.9 29.8 23.8 32.3 .74 3 a. m. 5 a. m. 157.8 46.0 29.2 49.9 31.6 25.4 32.2 .79 5 a. m. 7 a. m. Total 159.3 45.4 28.5 49.0 30.8 24.9 31.8 .79 2108.8 627.5 29.8 696.6 33.0 354.6 439.3 0.81 1 See p. 122. Metabolism Experiments Nos. 79-83, 85, and 89. 273 Table 174. — Oxygen and carbon dioxide thermal quotients and respiratory quotients — Continued. (a) (6) (O (d) (e) (/) fa) (ft) Experiment number, subject, and date. Period. o t-. p. cs a S3 «5 a o o . 5® &a >>3 ygen thermal quotient ■a §1 .0.9 rbon dioxide thermal quotient (100d-^a). lume of car- bon dioxide eliminated (dxO.5091). lume of oxy- en consumed (6X0.7). 5.8? M-w •• o-a y, °° M 3® a o O 60 a> o1 H O O o O > > 85. N.M.P. cont. 1905. Cals. Grams. Grams. Liter 8. Liters., Dec. 10-11. 7 a. m. to 9 a. m. 231.0 67.3 29.2 77.0 33.3 39.2 47.10.83 9 a. m. 11 a. m. 200.2 50.8 25.4 62. 2, 31.0 31.7 35.6 .89 11a.m. 1p.m. 207.2 61.9 29.9 61.8 29.9 31.5 43.3 .73 1 p. m. 3 p. m. 198.8 61.9 31.1 61.9 31.1 31.5 43.3 .73 3 p. m. 5 p. m. 166.5 50.2 30.1 56.6 34.0 28.8 35.1 .82 5 p. m. 7 p. m. 194.6 56.9 29.2 55.7 28.6 28.4 39.8 .71 7 p. m. 9 p. m. 230.9 67.1 29.1 67.5 29.2 34.3 47.0 .73 9 p. m. 11 p. m. 184.2 59.3 32.2 61.5! 33.4 31.3 41.5 .75 11 p. m. 1 a. m. 177.5 51.2 28.8. 56. 2i 31.7 28.6 35.8 .80 1 a. m. 3 a. m. 173.1 48.7 28.2 50.7! 29.3 25.8 34.1 .76 3 a. m. 5 a. m. 162.8 50.5 31. 0: 52.9 32.5 26.9 35.4 .76 89. D. W. 5 a. m. 7 a. m. Total 177.9 49.8 28.0 55.3 31.1 28.2 34. 9i .81 2304.7 675.6 29.3719.3 31.2 366.2 472.90.77 | 1906. Preliminary: Jan. 9-10.. 11 p. m. to 1 a. m. 1 158.1 54.8 34.7 66.2 41.9 33.7 38.30 1 a. m. 3 a. m. 1 177.0 45.9 25.9 51.6 29.1 26.2 32.2 .82 3 a. m. 5 a. m. U57.1 45.0 28.6 55.2 35.1 28.1 31.5 .89 Jan. 10-11. 5 a. m. 7 a. m. Total, 8 hours... . 7 a. m. to 9 a. m. 1 151.4 50.9 33.6 52.6 34.8 26.8 35.6 .75 1 643.6 196.6 30.5 225.6 35.1 114.8 137.6 0.84 231.2 62.4 27. 0| 75.1 32.5 38.2 43. 710.88 9 a. m. 11 a. m. 110.1 37.4 34.0 57.3 52.0 29.2 26.21.11 11 a. m. 1 p. m. 139.2 46.8 33.6 53.3 38.3 27.1 32.8 .83 1 p. m. 3 p. m. 256.0 67.1 26.2 65.6 25.6 33.4 47.0 .71 3 p. m. 5 p. m. 108.6 40.0 36.8 54.4 50.1 27.7 28.0 .99 5 p. m. 7 p. m. 241.6 68.4 28.3 63.5 26.3 32.3 47.9 .68 7 p. m. 9 p. m. 182.2 56.3 30.9 69.4 38.1 35.3 39.4 .90 9 p. m. lip. m. 236.9 79.5 33.5 74.6 31.5 38.0 55.6 .68 11 p. m. 1 a. m. 152.2 38.9 25.6 50.3 33.0 25.6 27.2 .94 1 a. m. 3 a. m. 154.0 46.0 29.9 48.2 31.3 24.6 32.2 .76 3 a. m. 5 a. m. 175.1 47.7 27.2 56.7 32.4 28.9 33.4 .87 Jan. 11-12. 5 a. m. 7 a. m. Total 163.0 54.9 33.7 54.0 33.1 27.5 38.4 .72 2150.1 645.4 30.0 722.4 33.6 367.8 451.80.81 7 a. m. to 9 a. m. 245.1 67.2 27.4 65.7 26.8 33.4 47.1 0.71 9 a. m. 11 a. m. 119.1 44.4 37.3 54.5 45.7 27.7 31.1 .89 11 a. m. 1 p. m. 166.2 52.6 31.6 58.6 35.3 29.8 36.8 .81 1 p. m. 3 p. m. 264.4 74.7 28.2 66.0 25.0 33.6 52.3 .64 3 p. m. 5 p. m. 111.3 49.7 44.6 50.8 45.7 25.9 34.8 .74 5 p. m. 7 p. m. 254.7 74.6 29.3 66.9 26.3 34.1 52.2 .65 - 7 p. m. 9 p. m. 200.4 65.7 32.8 75.1 37.5 38.2 46.0 .83 9 p. m. 11 p. m. 234.5 76.1 32.5 67.7 28.9 34.5 53.3 .65 11 p. m. 1 a. m. 191.3 40.4 21.1 53.0 27.7 27.0 28.2 .96 1 a. m. 3 a. m. 133.6 49.8 37. 31 49.5 37.1 25.2 34.9 .72 3 a. m. 5 a. m. 168.2 48.2 28.7 50.3 29.9 25.6 33.7 .76 5 a. m. 7 a. m. Total 165.5 37.9 22.9, 47.4 28.7 24.2 26.5 .91 2254.3 681.3 30. 2705. 5! 31.3 i 359.2 476.9 0.75 18 1 See p. 122. 274 Influence of Inanition on Metabolism. Nitrogen Metabolism Experiments. Second only in importance to the study of the effect of inanition on meta- bolism is the study of the recovery of material lost during fasting. In the original plan under which the fasting experiments were projected a study of the recovery after fasting was contemplated. It was soon seen that such experiments were impracticable because as a rule, the subjects were unable to consume large amounts of food on the day immediately following a fast. It became necessary to modify materially the original plan and continue the food experiments in the respiration chamber for only a few days and with a small ration (hardly more than maintenance). None of the food experiments reported above were therefore of more than 3 days' duration. But although it was impossible to continue the experiments in the respiration calorimeter and thus obtain the complete balance of income and outgo, the plan was adopted of weighing and sampling all the food of the subject and noting the amounts of nitrogen, phosphorus, and sulphur in the food, feces, and urine of each day. This was done in the hope of obtaining much new information on the recoup- ment after prolonged inanition, since the recorded observations on the effect on metabolism of the ingestion of food after a period of inanition are extremely few. Arrangements were made therefore to weigh and analyze all the food eaten by the subject S. A. B. from March 14, the first day after metabolism experi- ment No. 76, until the beginning of metabolism experiment No. 77 and from the first day after experiment No. 77, i. e., April 12, until April 25, when he left Middletown, and hence was beyond our control. During these periods all the food and the urine and feces were sampled and analyzed. The data thus obtained constitute a study of the complete intake aud output of nitrogenous material for a period of 53 days, i. e., from the first day of fasting experiment No. 75 until April 25. The intake and output of phosphorus were likewise studied and hence the gains or losses of this element can be determined. In a measure the same is true of the sulphur balance. Furthermore, since the heat of combustion of the food eaten was determined each day, much valuable information regarding the energy con- sumption was secured. Diet. — With the exception of experiment No. 77, each fast made by the subject S. A. B. was followed by an experiment in the respiration chamber in which food was given (experiments Nos. 72, 74, and 76). The change from fasting to food metabolism, therefore, was studied while the subject was in the chamber. Usually the quantities of food that the subject wished to take on the first day after the fasting period were small, barely more than enough for maintenance. Nitrogen Metabolism Experiments. 275 At the conclusion of the food experiment inside the chamber, which usually lasted 3 days, the subject came out of the respiration apparatus, and resumed his customary dietetic habits. After metabolism experiment No. 76, the subject began the experiment outside the chamber, the results of which are here recorded under the convenient heading of Nitrogen Metabolism Experi- ment No. 1. The second nitrogen metabolism experiment immediately followed metabolism experiment No. 77. The peculiar dietetic habits of the subject of these experiments resulted in his selecting a very extended list of food materials. He was left perfectly free to choose whatever he wished both as to kinds and amounts of food. In general, meat was partaken of very sparingly. Milk, fruit, cereals, and eggs, together with peanut butter, were the chief articles of diet. The wide variety of food selected may be seen from table 175, which gives the percentage composition and the different kinds of food used in both nitrogen metabolism experiments. The values here recorded are used in subsequent computations to test the accuracy of the method of sampling the food employed in these experiments. The percentage composition of the foods given in this table is taken for the most part from the compilation of analyses of American food materials published by Atwater and Bryant.83 In many instances the analyses taken from the bulletin referred to, were revised by averaging with new unpublished data. Sampling and Analysis. It was impracticable to sample and analyze each kind of food consumed each day. The following plan was therefore adopted: The amount of each kind of food eaten in a given 24 hours was determined, and a tenth of each kind of food was placed in a dish. These different samples formed, then, a com- posite representing one-tenth of all the food eaten and including each kind of food. This composite sample was dried in the water oven, then weighed, mixed and ground preparatory to its analysis. Determinations were made on the sample for each day, of nitrogen, phosphorus, sulphur, and the heat of combustion. Calculated amounts of protein and energy of food. — The method of sampling and analysis outlined above is open to the objection that one or more articles of so varied a diet might be overlooked in preparing the daily composite samples. As a partial check on the accuracy of this method of sampling, the amounts of protein and energy in the food consumed per day have been computed from the weights of the different foods eaten and the percentage composition given in table 175. This table is furthermore of interest in showing the kinds and amounts of food eaten each day. 33 U. S. Dept. Agr., Office of Expt. Sta. Bui. 28 (1898). 27 0 Influence of Inanition on Metabolism. Table 175. — Proportion of nutrients assumed l for and calculated heat of combus- tion of food eaten — Nitrogen metabolism experiments Nos. 1 and 2. Kind of food. (a) Protein (2^X6.25). Fat. (c) Carbo- hydrates. (d) Heat of combustion per gram. Per cent. 23.5 28.6 19.0 21.7 21.8 14.8 14.0 1.0 18.7 3.5 3.3 16.1 14.3 10.9 10.6 9.9 9.2 9.7 9.8 5.9 6.9 1.1 1.1 1.2 1.6 3.6 .9 1.1 .3 1.3 1.0 .8 2.1 4.3 2.1 1.5 2.3 .8 21.0 5.7 .4 29.3 11.0 16.6 .4 12.9 2.4 1.1 Per cent. 30.4 11.6 34.1 12.1 10.5 12.0 85.0 27.4 18.5 4.0 7.4 3.1 2.0 1.4 'i:3 .9 9.1 10.9 2.5 .4 .1 .3 .3 :! 27.6 .3 .6 .7 .2 2.8 .3 3.0 54.9 50.6 1.5 46.5 71.2 63.4 48.7 100.0 Per cent. .... i!s 4.5 5.0 66.5 73.3 75.4 78.1 53.1 49.7 73.1 64.1 19.6 9.3 3.3 2.9 9.9 9.8 3.9 11.6 10.8 22.0 8.5 11.6 78.4 74.2 73.3 52.3 68.5 39.6 17.3 27.9 4.6 17.1 13.3 16.1 100.0 81.2 30.3 69.3 Calories. 3 266 Chicken, canned * 2 718 4 313 4 070 2.381 1 849 Eggs, boiled 1 945 Butter 7 919 3 650 2 028 Milk 751 4 415 4 191 Wheat breakfast food, rolled * 3 985 Wheat breakfast food, shredded4 4.025 Breakfast food, crisped and malted4 3.822 2.885 2.734 4 . 485 4.007 1.407 .483 .203 Lettuce .210 .524 .635 .246 3.088 .476 1 .003 .457 .524 3 . 506 3.219 3.041 2.170 3.139 1.626 7.010 6.186 .353 6.690 7.795 7.502 3.960 3.207 6.515 2 . 873 .310 9.300 Lemons Sugar Honev Molasses candy8 Olive oil . . 1 O. S. Dept. Agr., O. E. S., Bui. 28, ex- cept as noted. 2 Composition assumed from unpublished data. 8 Protein and energy determined. U. S. Dept. Agr., O. E. S., Bui. 152. * Conn. (Storrs) Report, 1904. 6 Composition assumed as fruit cake. 8 Composition calculated from U. S. Dept. Agr., O. E. S., Bui. 150. by means of known percentage of water added. T Assumed as chocolate. 8 Assumed as molasses. Nitrogen Metabolism Experiments. 277 Table 176. — Calculated protein and energy in fresh food — Nitrogen metabolism experiments Nos. 1 and 2. NITROGEN METABOLISM EXPERIMENT No. 1. Date. Kind of food. (a) Weight of food. (6) Protein. (c) Energy. 1905. Mar. 14. Mar. 15. Mar. 16. Eggs, boiled Butter Cheese, cream Cream Milk Wheat, rolled, raw Bread, white Bread, wheat, whole Onions, raw Apples Bananas Oranges Dates Figs Raisins Peanut butter Sugar Cocoa, as beverage Total for day Eggs, raw Butter Cheese, cream Cream Milk Wheat, rolled, raw Bread, white Bread, whole wheat Onions, raw Apples Bananas Oranges Dates Prunes Raisins Peanut butter Total for day Eggs, boiled and scrambled Butter Cream Milk Bread, white Apples Bananas Oranges Figs Peanut butter Total for day Grams. 101.6 38.1 16.5 235.9 1057.0 118.8 69.6 264.4 75.3 612.1 302, 145, 95, 95, 56. 35, 6, 199, 3525 . 6 79.4 39.9 102.5 368.8 1186.4 95.1 222.9 190.7 75.7 352.8 435.8 624.9 109.5 181.1 55.1 61.7 4182.3 71.1 27.6 84.1 1537.0 375.1 373.8 177.7 636.9 71.9 95.4 Grams. 14.22 .38 3.09 5.90 34.88 12.95 6.40 25.65 1.20 1.84 3.93 1.17 2.01 4.10 1.30 10.31 2! i6 131.52 11.75 .40 19.17 9.23 39.15 10.37 20.51 18.50 1.21 1.06 5.67 5.00 2.30 3.80 1.27 18.08 167.47 9.95 .28 2.10 35.61 34.51 1.12 2.31 5.09 3.09 27.96 3450.6 122.02 Calories. 198 302 60 478 794 473 201 723 39 275 303 76 336 307 177 235 25 62 5064 147 316 374 748 891 379 743 521 40 168 437 327 384 551 173 413 6612 138 218 171 1155 1082 179 179 334 231 638 4325 278 Influence of Inanition on Metabolism. Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 1 (Cont'd). Date. Kind of food. (a) Weight of food. (6) Protein. (O Energy. 1905. Mar. 17 Eggs, boiled and scrambled. . Butter Grams. 81.5 38.8 46.3 748.5 178.0 111.3 301.8 189.9 310.6 177.1 220.5 106.7 27.0 Grams. 11.41 .38 8.66 24.70 16.38 10.80 .91 2.47 2.48 7.62 3.31 .85 7.91 Calories. 159 307 169 562 514 304 144 190 163 570 478 173 181 Mar 18 Milk Bread, white Bread, whole wheat Apples Bananas Oranges Fisrs Peanut butter 2538.0 97.88 3914 84.3 11.0 35.3 359.3 1593.7 63.9 316.1 241.5 440.1 342.5 246.0 86.0 94.3 208.8 17.8 88.4 22.4 12.48 .11 6.60 8.99 52.60 6.97 30.66 .72 5.72 2.74 5.16 3.70 3.L3 3.74 25.90 3.72 156 87 131 729 1197 255 864 115 441 179 863 277 *453 125 591 168 Mar. 19 Butter Cheese, cream Cream Milk Wheat, rolled, raw Bread, whole wheat Apples Bananas Oranges Dates Figs Prune juice Prunes, soaked in water Almonds Peanut butter Walnuts, English 4251.4 172.94 6631 Eggs, boiled and scrambled. . Butter 121.3 20.7 46.5 279.6 1338.0 70.5 106.9 48.2 34.4 354.5 308.8 310.2 141.2 52.8 52.8 16.98 .21 8.70 6.99 44.16 7.68 10.37 4.72 .55 1.06 4.01 2.48 2.97 1.21 .21 236 164 170 567 1005 281 292 216 18 169 310 163 495 166 169 Cheese, cream Cream Milk Wheat, rolled, raw Bread, whole wheat Crackers Onions Apples Bananas Oranges Dates Raisins Total for day 3286.4 112.30 4421 Nitrogen Metabolism Experiments. 279 Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 1 (Cont'd). Date. Kind of food. (a) Weight of food. (6) Protein. (e) Energy. 1905. Mar. 20. Mar. 21. Mar. 22. Eggs, boiled and scrambled Butter Cheese, cream Cream Milk Wheat, rolled, raw Bread, whole wheat Cake, raisin Bananas Oranges Dates Figs Raisins Almonds Honey Walnuts, English Total for day Eggs, boiled and scrambled Butter Milk Bread, whole wheat Cake, raisin Beans, baked, canned Onions Apples Bananas Oranges Figs. Raisins Peanut butter Total for day Eggs, boiled and scrambled Butter Cream Milk Oats, rolled, raw Bread, whole wheat Crackers, soda Onions Peas, canned Apples Bananas Oranges Dates Figs Raisins Walnuts, English Total for day Grams. 90.6 23.5 74.6 310.0 1361.1 82.6 148.7 44.1 449.7 148.1 123.3 140.0 72.9 64.0 56.1 52.2 3241.5 .3 .5 .7 .3 .5 69 37 1240, 316, 64. 310.2 45.8 330.4 457.4 642.6 134.3 65.8 57.7 3772.5 102.8 86.6 324.5 731.7 70.2 337.3 18.3 11.2 361.6 283.1 506.1 303.5 198.1 110.2 45.3 43.2 3533.7 Grams. 12.68 .24 13.95 7.76 44.92 9.00 14.42 2.60 5.85 1.18 2.59 6.02 1.68 13.44 .22 8.67 145.22 9.70 .38 40.94 30.69 3.81 21.40 .73 .99 5.94 5.14 5.77 1.51 16.91 143.91 14.39 .87 8.11 24.14 11.30 32.72 1.79 .18 13.02 .85 6.58 2.43 4.16 4.74 1.04 7.17 133.49 Calories. 176 226 272 628 1022 329 407 177 552 78 432 451 229 448 180 392 5999 135 307 932 865 258 435 24 157 459 337 432 207 386 4934 200 686 659 549 310 922 82 6 230 136 507 159 695 355 142 324 5962 280 Influence of Inanition on Metabolism. Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 1 (Cont'd). Date. Kind of food. (a) Weight of food. (b) Protein. (c) Energy. 1905. Mar. 23. Mar. 24. Mar. 25. Ham, pressed and deviled Butter Cream Milk Wheat, rolled, raw Bread, whole wheat Apples Bananas Oranges Dates Prunes Raisins Almonds Total for day Beefsteak Butter Cream Milk Wheat, rolled, raw Bread, whole wheat Onions Bananas Oranges Dates Figs Raisins Molasses candy Total for day Eggs, boiled and scrambled Butter Cheese, cream Cream Milk Wheat, rolled, raw Bread, whole wheat Tomatoes Apples Bananas Oranges Dates Figs Raisins Pecans Walnuts, English Honey Total for day Grams. 64.4 60.0 205.9 484.3 82.6 177.8 282.9 570.9 323.5 114.5 334.3 91.7 30.3 2823 . 1 153.6 56.7 205.7 1263.3 71.2 197.4 45.6 166.5 433.0 158.9 31.5 72.0 37.9 2893.3 119.4 51.5 48.8 248.0 1125.2 48.3 174.1 88.4 343.8 402.5 340.0 119.4 210.2 53.6 58.5 51.7 102.6 3586.0 Grams. 12.24 .60 5.15 15.98 9.00 17.25 .85 7.42 2.59 2.40 7.02 2.11 6.36 88.97 36.10 .57 5.14 41.69 7.76 19.15 .73 2.16 3.46 3.34 1.35 1.66 .91 124.02 16.72 .52 9.13 6.20 37.14 5.26 16.89 .80 1.03 5.23 2.72 2.51 9.04 1.23 6.44 8.58 .41 129.85 Calories. 278 475 418 364 329 486 135 573 170 410 1017 288 212 5155 502 449 417 948 282 540 24 167 227 557 101 226 109 4549 232 408 179 503 846 192 476 22 164 404 178 419 677 168 456 388 329 6041 Nitrogen Metabolism Experiments. 281 Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 1 (Cont'd). Date. Kind of food. (a) Weight of food. (b) Protein. (c) Energy. 1905. Mar. 26. Mar. 27. Mar. 28. Salmon Butter Cream Milk Wheat, rolled, raw Bread, whole wheat Carrots Onions Peas, canned Tomatoes Bananas Lemons Dates Figs Raisins Cocoanut Cocoanut milk Peanut butter Olive oil Total for day Beefsteak Lamb chops Eggs, boiled and scrambled Butter Cheese, cream Cream Milk Bread, whole wheat Lettuce Tomatoes Bananas Oranges Figs Almonds Peanut butter Pecans Walnuts, English Chocolate, milk Total for day Eggs, scrambled Butter Milk Bread, whole wheat Peas, canned Bananas Oranges Dates Prunes Raisins Honey Total for day Grams. 272.1 24.7 234.1 1058.0 50.2 126.5 39.5 59.4 204.0 100.8 229.0 35.7 118.8 73.9 65.6 74.4 62.6 14.7 15.0 2859.0 35.4 46.6 108.9 88.8 61.6 146.4 1117.0 409.1 81.0 125.8 421.1 753.0 48.4 36.0 44.0 47.0 46.2 12.0 3628.3 160.5 41.4 1447.7 219.6 189.2 225.9 188.0 39.3 298.9 65.8 47.8 2924 . 1 Grams- 59.32 .25 5.85 34.92 5.47 12.27 .43 .95 7.34 .91 2.98 .36 .49 ,18 1.51 4.24 .25 4.31 2. 3. 147.03 8.32 10.11 15.25 .90 11.52 3.66 36.86 39.68 .97 1.13 5.47 6.03 2.08 7.56 12.89 5.17 7.67 1.55 176.82 22.47 .41 47.77 21.31 6.81 2.94 1.50 .83 6.28 1.51 .19 112.02 Calories. 648 196 475 795 200 346 19 31 133 25 230 16 417 238 206 460 22 98 140 4695 116 190 212 703 225 297 840 1119 17 31 422 395 156 252 294 366 347 78 6060 312 328 1087 600 120 227 99 138 909 207 153 4180 •28-3 Influence of Inanition on Metabolism. Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 1 (Cont'd). Date. Kind of food. (a) Weight of food. (&) Protein. (c) Energy. 1905. Mar. 29 Eggs, boiled and scrambled . . Butter Grams. 126.1 34.8 83.0 836.9 128.4 35.4 54.3 49.4 96.7 211.8 264.1 21.8 179.2 120.2 53.1 53.7 45.2 42.3 12.8 Grams. 17.65 .35 2.08 27.62 12.45 .39 .65 .79 .87 .64 3.43 .22 1.43 2.52 1.22 11.28 4.97 7.02 Calories. 245 276 168 628 351 17 11 26 24 101 265 10 94 421 167 376 352 317 119 Mar. 30 Cream Milk Bread, whole wheat Carrots Lettuce Onions Tomatoes Apples Bananas Lemons Oranges Dates Raisins Almonds Pecans Walnuts, English Olive oil Total for day 2449.2 95 . 58 3968 Butter 45.1 175.9 506.8 37.3 139.6 392.5 98.4 97.4 139.3 232.0 180.4 21.5 129.1 78.5 87.0 46.3 .45 4.40 16.72 4.07 13.54 27.08 1.18 1.56 1.25 .70 2.35 .22 1.03 1.65 2.00 .19 357 357 380 149 382 550 21 51 34 110 181 10 68 275 273 148 Mar. 31 Cream Milk Wheat, rolled, raw Bread, whole wheat Lettuce Onions Tomatoes Apples Bananas Lemons Oranges Dates Raisins Honey Total for day 2407.1 78.39 3346 Salmon, canned 250.3 36.6 191.8 1456.2 26.4 42.2 144.5 179.8 213.6 193.9 117.2 54.8 69.2 30.1 32.8 54.57 .37 4.80 48.06 3.75 4.60 14.02 1.62 .64 2.52 .94 1.15 1.59 3.31 5.44 596 290 389 1094 111 168 395 44 102 194 61 192 217 235 246 Butter Cream Milk Wheat breakfast food, raw . . Wheat, rolled, raw Bread, whole wheat Tomatoes Apples Bananas Oranges Raisins Pecans Walnuts, English Total for day 3039.4 147.38 4334 Nitrogen Metabolism Experiments. 283 Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 1 (Cont'd). Date. Kind of food. (a) Weight of food. (&) Protein. (c) Energy. 1905. Apr. 1. Apr. 2. Apr. 3. Salmon, canned Eggs, boiled and scrambled Butter Cheese, cream Cream Milk Bread, whole wheat Peas, canned Tomatoes Bananas Oranges Prunes Peanut butter Total for day Beefsteak Butter Cream Milk Wheat, rolled, raw Bread, whole wheat Lettuce Onions Tomatoes Apples Bananas Oranges Dates Figs. Raisins Total for day Eggs, boiled Butter Cheese, cream Cream Milk Wheat breakfast food, shredded Bread, whole wheat Apples Bananas Oranges , Raisins , Walnuts, English ". , Total for day Grams. 61.0 88.6 65.2 55.2 168.4 1610.7 211.7 209.7 133.3 147.5 293.2 148.8 23.5 3216.8 166.9 33.5 152.6 1259.1 48.4 108.8 142.5 81.3 222.8 355.8 209.1 259.7 63.5 39.3 40.9 3184.2 82.3 29.6 66.6 321.9 861.0 58.2 104.5 231.2 375.9 260.1 18.8 56.4 2466.5 Grams. 13.30 12.40 .65 10.32 4.21 53.15 20.54 7.55 1.20 1.92 2.35 3.12 6.89 137.60 39.22 .34 3.82 41.55 5.28 10.55 1.71 1.30 2.01 1.07 2.72 2.08 1.33 1.69 .94 115.61 11.52 .30 12.45 8.05 28.42 6.17 10.14 .69 4.89 2.08 .43 9.36 94.50 Calories. 145 172 516 201 342 1210 579 133 33 148 154 453 157 4243 545 265 309 946 193 297 30 43 55 169 210 136 223 127 128 3676 160 234 243 753 646 234 286 110 377 136 59 423 3661 284 [npldence of Inanition on Metabolism. Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 1 (Cont'd). Date. Kind of food. (a) Weight of food. (b) Protein. Energy. 1906. Apr. 4. Apr. 5. Apr. 6. Eggs Butter Cream Milk Wheat, rolled, raw Bread, whole wheat. . . . Tomatoes Apples Bananas Raisins Peanut butter Total for day Beefsteak Butter Cream Milk Wheat, rolled Bread, white Onions Tomatoes Apples Bananas Dates Figs Raisins Total for day Butter Cream Milk Wheat breakfast food, shredded Bread, white Beans, baked, canned. . Lettuce Apples Bananas Lemons Oranges Almonds Pecans Total for day Grams. 136.3 30.3 296.5 868.4 34.5 108.1 219.7 203.8 198.0 71.0 46.4 2213.0 165.8 35.1 337.8 859.8 36.2 106.8 75.9 132.3 260.8 357.8 76.3 320.8 41.2 2806.6 36.5 227.7 1395.5 59.0 106.7 355.3 115.7 150.1 157.3 14.1 146.6 55.3 49.7 2869.5 Grams. 19.08 .30 7.41 28.66 3.76 10.49 1.98 0.61 2.57 1.63 13.60 90.09 38.96 .35 8.44 28.38 3.95 10.36 1.21 1.19 .78 4.65 1.60 13.79 .95 114.61 .37 5.69 46.05 6.25 10.35 24.52 1.39 .45 2.04 .14 1.17 11.61 5.47 115.50 Calories. 265 240 601 653 137 276 54 97 199 223 310 3055 542 278 685 646 144 292 40 33 124 359 268 1033 129 4573 289 462 1048 237 292 498 24 71 158 6 77 388 387 3937 Nitrogen Metabolism Experiments. 285 Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 1 (Cont'd). Date. Kind of food. (a) Weight of food. (6) Protein. (O Energy. 1905. Apr. 7 Eggs, boiled and scrambled . . Grams. 100.9 37.0 30.9 783.4 64.7 91.9 225.6 129.0 76.9 128.2 Grams. 14.13 .37 5.78 25.85 6.28 8.91 15.57 1.03 3.31 2.95 Calories. 196 293 113 589 177 251 316 68 248 402 Milk Bread, whole wheat Beans, baked, canned Oranges Fiers Raisins Total for day 1668.5 84.18 2653 NITROGEN METABOLISM EXPERIMENT No . 2. 1905. Apr. 12 Milk 665.9 166.6 124.9 158.4 433.8 21.97 15.33 1.12 1.58 3.47 500 481 31 72 227 Apr. 13 Bread Tomatoes Lemons Oranges Total for day 1549.6 43.47 1311 Beefsteak 123.7 26.7 47.7 50.5 761.0 269.4 88.5 46.3 262.2 79.9 206.5 330.8 29.07 3.95 .48 9.44 25.11 24.78 1.06 .74 2.36 1.04 2.07 2.65 404 49 378 184 572 777 19 24 65 80 94 173 Apr. 14 Eire, raw Butter Cheese Milk Bread Lettuce Onions Tomatoes Bananas Lemons Oranges 2293.2 102.75 2819 Salmon, canned Eee:. raw 164.9 20.3 21.6 131.3 931.0 27.8 146.6 139.5 178.5 136.0 17.8 160.4 81.7 236.6 24.3 45.8 35.95 3.00 .22 3.28 30.72 3.95 13.49 5.02 1.61 1.77 .18 1.28 1.72 3.55 .56 13.42 393 38 171 266 699 117 423 89 44 136 8 84 286 513 76 306 oo' Butter Cream Milk Wheat germ food Bread Peas l. Tomatoes Bananas Lemons Oranges Dates Prunes, soaked in water Raisins Peanut butter Total for day 2464 . 1 119.72 3649 1 Composition assumed as canned peas. 286 Influence of Inanition on Metabolism. Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 2 (Cont'd). Date. Kind of food. (a) Weight of food. 00 Protein. (c) Energy. 1905. Apr. 15. Apr. 16. Apr. 17. Apr. 18. Chicken, canned Salmon Butter Milk Bread Celery Peas1 Tomatoes Apples Bananas Oranges Total for day Beefsteak Eggs, raw Cream Cheese Milk Bread, whole wheat Celery Tomatoes Olives Apples Oranges Total for day Eggs, boiled and scrambled Cream Milk Bread Beans, baked, canned Lettuce Tomatoes Bananas Oranges Dates Peanut butter Pecans Walnuts, English Total for day Beefsteak Cream Milk Wheat, rolled Bread Bananas Oranges Dates Prunes, soaked in water. . . Raisins Almonds Total for day Grams. 131.0 114.7 26.3 1481.1 260.7 61.8 179.3 156.1 142.1 307.0 114.9 2975.0 177.1 114.4 146.4 54.2 1322.5 206.1 81.0 135.0 39.9 332.9 160.4 2769.9 139.4 186.0 1657.3 281.5 396.4 128.4 166.3 313.2 170.0 119.4 79.2 61.4 56.7 3755.2 192.2 175.0 1142.9 25.4 118.2 129.7 176.4 139.9 267.0 28.4 68.9 2464.0 Grams. 37.47 25.00 .26 48.88 23.98 .68 6.45 1.40 .43 3.99 .92 149.46 41.62 16.93 3.66 10.14 43.64 19.99 .89 1.22 .44 1.00 1.28 140.81 19.52 4.65 54.69 25.90 27.35 1.54 1.50 4.07 1.36 2.51 23.21 6.75 9.41 182.46 45.17 4.38 37.72 2.77 10.87 1.69 1.41 2.94 4.01 .65 14.47 126.08 Calories. 356 273 208 1112 752 13 114 38 68 308 60 3302 578 212 297 198 993 563 16 33 123 158 84 3255 271 377 1245 812 555 27 41 314 89 419 530 479 425 5584 628 355 858 101 341 130 92 490 579 89 483 4146 1 Composition assumed as canned peas. Nitrogen Metabolism Experiments. 287 Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 2 (Cont'd). Date. 1905. Apr. 19. Apr. 20. Apr. 21. Apr. 22. Kind of food. (a) Weight of food. Milk Bread Beans Celery Lettuce Tomatoes Bananas Oranges Dates Peanut butter Total for day Beefsteak Eggs, boiled and scrambled Cheese Milk Bread Celery Onions Tomatoes Apples Bananas Oranges Total for day Salmon, canned Butter Cream Milk Wheat, raw Bread Celery Onions, raw Apples Bananas Oranges Dates Raisins Total for day Chicken, canned Butter Cream Milk Bread Celery Tomatoes Bananas Oranges Dates Raisins Total for day Grams. 1313.1 197.1 388.6 31.5 18.9 243.0 193.8 190.8 93.1 61.3 2731.2 168.1 147.9 69.3 1291.3 178.5 48.0 51.9 148.4 279.7 134.3 210.6 2728.0 234.9 31.6 184.6 1111.3 35.3 120.6 51.1 64.1 370.1 252.6 129.8 153.1 43.1 2782.2 95.0 22.3 198.4 1208.3 193.2 76.1 155.0 155.2 372.4 46.0 45.3 2567.2 (W Protein. Grams. 43.33 18.13 26.81 .35 .23 2.19 2.52 1.53 1.96 17.96 115.01 39.50 20.71 12.96 42.61 16.42 .53 .83 1.34 .84 1.75 1.68 139.17 51.21 .32 4.62 36.67 3.85 11.10 .56 1.03 1.11 3.28 1.04 3.22 .99 119.00 27.17 .22 4.96 39.87 17.77 .84 1.40 2.02 2.98 .97 1.04 99.24 (O Energy. Calories. 986 569 544 6 4 60 194 100 326 410 3199 549 288 253 970 515 10 27 37 133 135 110 3027 559 250 374 835 141 348 10 34 176 253 68 537 135 3720 258 177 402 907 557 15 38 156 195 161 142 3008 2SS Influence of Inanition on Metabolism. Table 176. — Calculated protein and energy in fresh food — Continued. NITROGEN METABOLISM EXPERIMENT No. 2 (Cont'd). Date. Kind of food. (a) Weight of food. (0) Protein. (O Energy. 1905. Apr. 23 Beefsteak Orams. 182.4 23.8 181.2 1186.0 50.7 118.8 211.3 272.6 168.8 89.6 49.9 80.4 61.6 Orams. 42.86 .24 4.53 39.14 5.53 10.93 1.90 3.54 1.35 1.88 .75 8.84 10.23 Calories. 596 188 367 891 202 343 52 273 88 314 108 627 462 Apr. 24 Butter Cream Milk Wheat, raw Bread Tomatoes Bananas Oranges Dates Prunes Pecans Walnuts, English Total for day 2677.1 131.72 4511 Salmon, canned. . . , 168.0 78.7 38.2 1375.5 151.0 51.2 289.8 46.6 188.6 298.0 36.62 11.65 7.14 45.39 13.89 .56 2.61 .51 2.45 2.38 400 146 139 1033 436 10 71 144 189 156 Apr. 25 Cheese Milk Bread Celery Tomatoes Olives Bananas Oranges Total for day 2685.6 123.20 2724 Lamb chops 101.7 186.1 1497.4 43.5 157.6 119.7 324.2 200.5 232.9 106.3 22.07 4.65 49.41 4.31 14.50 1.44 2.92 2.61 1.86 2.23 414 377 1125 166 455 25 80 201 122 373 Cream Milk Wheat breakfast food Bread Lettuce Tomatoes Bananas Oranges Dates Total for day 2969.9 106.00 3338 1 Composition assumed to be the same as for raw. Determinations of nitrogen, phosphorus, sulphur, and heat of combustion of food. — While for the approximations of the ordinary dietary study the use of average figures for the percentage composition of food materials is fully justified, in an accurate balance experiment in which it is desired to know the income and outgo of material and energy, accurate analyses alone can be used. Such analyses were made of the composite sample of food for each day. The results are given in table 177, which includes also the total amounts of each element determined and the total energy of the food for each week and the average per day. Nitrogen Metabolism Experiments. 289 Table 177. — Amounts of determined nitrogen, protein, phosphorus, sulphur, and energy in food — Nitrogen metabolism experiments Nos. 1 and 2. Labor- atory num- ber. Experiment number and date. (a) tf» (c) (d) Nitro- gen. Pro- tein (JVX 6.25). Phos- phorus. Phos- phoric acid (P«05). (e) Sul- phur. (f) (0) Sul- phur Heat of triox- combus- ide tion. (S03). 3864 3865 3866 3867 3868 3869 3870 3871 3872 3873 3874 3875 3876 3877 3878 3879 3880 3881 3882 3883 3SS4 3885 3886 3887 3888 3893 3894 3895 3896 3897 3898 3899 Experiment No. 1. First week: Mar. 14-15, 1905 Mar. 15-16, 1905 Mar. 16-17, 1905 Mar. 17-18, 1905 Mar. 18-19, 1905 Mar. 19-20, 1905 Mar. 20-21, 1905.... Total for 1st week. Average per day. . . Second week : Mar. 21-22, 1905 Mar. 22-23, 1905 Mar. 23-24, 1905 Mar. 24-25, 1905 Mar. 25-26, 1905 Mar. 26-27, 1905 Mar. 27-28, 1905 Total for 2d week. . Average per day. . . Third week: Mar. 28-29, 1905 Mar. 29-30, 1905 Mar. 30-31, 1905 Mar. 31- April 1,1905. Apr. 1-2,1905 Apr. 2-3,1905 Apr. 3-4,1905 Total for 3d week.. Average per day. . . Fourth week: Apr. 4-5,1905 Apr. 5-6, 1905 Apr. 6-7,1905 Apr. 7-8,1905 Total for 4 days Average per day. . . Experiment No. 2. First week: Apr. 12-13,1905 Apr. 13-14,1905 Apr. 14-15, 1905 Apr. 15-16, 1905 Apr. 16-17,1905 Apr. 17-18,1905 Apr. 18-19, 1905 Total for 1st week. . Average per day. . . 0 ins. Gins. 20.55 128.44 25.09 21.65 14.90 26.34 17.10 22.14 147.77 21.11 156.81 135.31 93 . 13 164.62 106.88 138.38 923 . 57 131.94 Gins. 4.479 6.853 5.162 4.228 3.895 3.420 4.703 32 . 740 4.677 Gins. 10.257 15.696 11.823 9.685 8.922 7.833 10.770 Gins. 1.822 2.038 1.142 .335 1.613 1.135 3.066 Gins. 4.547 5.086 2.851 .836 4.025 2.834 7.652 74.986 11.151 10.712 1.593 27.831 3.976 Cals. 5,592 6,771 4,857 3,943 6,527 4,810 6,200 38,700 5,529 22. 14! 138. 38 19.90 124.38 15.01 21.55 20.80 25.55 27.64 93.81 134.69 130.00 159.69 172.75 3.461 4.000 2.260 3.716 4.534 4.006 4.610 152.59 953.70 26.587 21.80136.24; 3.798 7.926 9.161 5.175 8.510 10.385 9.176 10.558 60.891 8.699 1.538 1.577 1.095 1.569 1.764 1.922 2.061 11 1 526 647 3.839 3.938 2.733 3.916 4.403 4.797 5.143 28 . 769 4.110 5,611 6,092 4,598 4,847 6,786 4,752 6,507 39,193 5,599 16 12 11 24 22 19, 13. 62 103.88 83 80.19 74 73.38 69 154.31 65 141.56 14 119.63 99 87.44 2.755 2.053 1.922 3.188 3.220 2.586 2.306 6.311 4.701 4.401 7.301 7.376 5.922 5.281 121.66 760.39 18.030|41.293 17.38 108.63 2.576' 5.899 .444 .094 036 .677, ,609 419 972 3 . 606 2.730 2.586 4.187 4.015 3.541 2.426 9.25123.091 1.322 3.299 3,874 3,440 3,446 4,357 4,356 3,798 3,847 27,118 3,874 14.03 17.91 18.01 12.73 62.68 15.67 87.69, 2.194 111.94 2.467 112.56 79.56 391.75 97.94 3.171 1.927 5.025 5.651 7.263 4.413 9.759 22.352 2.440! 5.588 1.070 1.430 1.211 .990 4.701 1.175 2.671 3.569 3.020 2.471 11.731 2.933 3,395 4,640 4,208 2,752 14,995 3,749 7.10 44.38 16.23 101.44 20.04 125.25 23.81 148.81 21.67 135.44 28.98 181.13 19.45 121.56 137.28 858.01 19.61 122.57 1.185 1.958 2.575 2.759 2.635 4.302 2.586 18.000 2.571 2.713 4.484 5.897 6.320 6.035 9.854 5.923 .610 1.275 1.452 1.727 1.525 2.037 1.264 1.522 3.183 3.624 4.312 3.805 5.084 3.155 41.226 9. 890124. 685 5.889 1.413 3.526 1,334 2,806 3,573 3,300 3,362 5,621 3,768 23,764 3,395 19 •2\ tO Influence of Inanition on Metabolism. Table 177. — Amounts of determined, nitrogen, protein, phosphorus, sulphur, and energy in food — Continued. Labor- atory num- ber. Experiment number and date. (a) Nitro- gen. 0>) Pro- tein (Nx 6.26). (c) Phos- phorus. m Phos- phoric acid (Ps06). (e) Sul- phur. (/) Sul- phur triox- ide (SO,). (ff) Heat of combus- tion. 3900 3901 3902 3903 3904 3905 3906 Experiment No. 2 — Cont'd. Second week: Apr. 19-20, 1905 Apr. 20-21, 1905 Apr. 21-22, 1905 Apr. 22-23, 1905 Apr. 23-24, 1905 Apr. 24-25, 1905 Apr. 25-26, 1905 Oms. 19.05 22.72 20.22 14.93 21.57 20.28 17.12 Oms. 119.06 142.00 126.38 93.31 134.81 126.75 107.00 Oms. 2.674 2.592 2.613 2.181 2.896 2.519 2.606 Gms. 6.124 5.936 5.984 4.996 6.633 5.770 5.968 Oms. 1.243 1.582 1.509 1.170 1.538 1.387 1.171 Oms. 3.102 3.949 3.766 2.919 3.840 3.462 2.922 CaU. 3,086 3,146 3,752 2,890 4,411 2,865 3,295 135.89 849.31 19.41 121.33 18.081 2.583 41.411 5.916 9.600 1.372 23.960 3.423 23,445 3,349 Comparison of calculated and determined amounts of protein and energy of food. — For reasons which will appear later, it seemed desirable to obtain some check upon the analytical work connected with the determination of nitrogen, and also upon the heats of combustion determined by the bomb calorimeter. The calculated protein and energy shown in table 176 were obtained primarily for this purpose. In table 178 the comparison of the calculated with the determined results is shown. It should be stated, however, that the unusual care taken in the preparation and analysis of these samples renders the comparison a test of the accuracy of the average figures rather than a check on the analyses. The comparison is nevertheless not without interest and it is accordingly here presented. Table 178. — Comparison of determined with calculated protein and energy of food — Nitrogen metabolism experiments Nos. 1 and 2. Protein (2VX6.26). Energy. (a) (&) (c) (d) (e) (/) Amount Amount determined determined Date. Deter- Calcu- greater(+) Deter- Calcu- greater(+) mined. lated. or less(— ) than calculated (a-b). mined. lated. or less(— ) than calculated (d-e). Experiment No. t. First week: Orams. Orams. Orams. Calories. Calories. Calories. Mar 14-15, 1905 128.44 131.52 - 3.08 5,592 5,064 + 528 Mar. 15-16, 1905 156.81 167.47 -10.66 6,771 6,612 + 159 Mar. 16-17, 1905 135.31 122.02 + 13.29 4,857 4,325 + 532 Mar. 17-18, 1905 93.13 97.88 - 4.75 3,943 3,914 + 29 Mar. 18-19, 1905 164.62 172.94 - 8.32 6,527 6,631 - 104 Mar. 19-20, 1905 106.88 112.30 - 5.42 4,810 4,421 + 389 Mar. 20-21, 1905 Total for 1st week.. . . 138.38 145.22 - 6.84 6,200 5,999 + 201 923.57 949.35 -25.78 38,700 36,966 + 1734 131.94 135.62 - 3.68 5,529 5,281 + 248 Nitrogen Metabolism Experiments. 291 Table 178. — Comparison of determined with calculated protein and energy of food — Continued. Protein (.ZVX6.25). Energy. (a) (6) (c) id) (e) (ft Amount Amount Date. determined determined Deter- Calcu- greater(+) Deter- Calcu- greater(+) mined. lated. or less(— ) than calculated (a-b). mined. lated. or less (— ) than calculated (d-e). Experiment No. ./ — Cont'd. Second week: Grams. Grams. Grams. Calories. Calories. Calories. Mar. 21-22, 1905 138.38 143.91 - 5.53 5,611 4,934 + 677 Mar. 22-23, 1905 124.38 133.49 - 9.11 6,092 5,962 + 130 Mar. 23-24, 1905 93.81 88.97 + 4.84 4,598 5,155 - 557 Mar. 24-25, 1905 134.69 124 . 02 + 10.67 4,847 4,549 + 298 Mar. 25-26, 1905 130.00 129.85 + .15 6,786 6,041 + 745 Mar. 26-27, 1905 159.69 147.03 + 12.66 4,752 4,695 + 57 Mar. 27-28, 1905 .... Total for 2d week. . . . 172.75 176.82 - 4.07 6,507 6,060 + 447 953.70 944.09 + 9.61 39,193 37,396 + 1797 Average per day Third week: 136.24 134.87 + 1.37 5,599 5,342 + 257 Mar. 28-29, 1905 103.8 ? 112.02 - 8.14 3,874 4,180 - 306 Mar. 29-30, 1905 80.19 95.58 -15.39 3,440 3,968 - 528 Mar. 30-31, 1905 73.38 78.39 - 5.01 3,446 3,346 + 100 Mar.31-Apr. 1,1905... 154.31 147.38 + 6.93 4,357 4,334 + 23 Apr. 1-2, 1905 141.56 137.60 + 3.96 4,356 4,243 + 113 Apr. 2-3, 1905 119.63 115.61 + 4.02 3,798 3,676 + 122 Apr. 3-4, 1905 Total for 3d week 87.44 94.50 - 7.06 3,847 3,661 + 186 760.39 781.08 -20.69 27,118 27,408 - 290 Average per day Fourth week: 108.63 111.58 - 2.96 3,874 3,915 - 41 Apr. 4-5, 1905 87.69 90.09 - 2.40 3,395 3,055 + 340 Apr. 5-6, 1905 111.94 114.61 - 2.67 4,640 4,573 + 67 Apr. 6-7, 1905 112.56 115.50 - 2.94 4,208 3,937 + 271 Apr. 7-8, 1905 Total for 4 days 79.56 84.18 - 4.62 2,752 2,653 + 99 391.75 404.38 -12.63 14,995 14,218 + 777 Average per day Experiment No. 2. 97.94 101.10 - 3.16 3,749 3,555 + 194 First week: Apr. 12-13, 1905 44.38 43.47 + .91 1,334 1,311 + 23 Apr. 13-14, 1905 101.44 102.75 - 1.31 2,806 2,819 - 13 Apr. 14-15, 1905 125.25 119.72 + 5.53 3,573 3,649 - 76 Apr. 15-16, 1905 148.81 149.46 - .65 3,300 3,302 - 2 Apr. 16-17, 1905 135.44 140.81 - 5.37 3,362 3,255 + 107 Apr. 17-18, 1905 181.13 182.46 - 1.33 5,621 5,584 + 37 Apr. 18-19, 1905 Total for 1st week.. . . 121.56 126.08 - 4.52 3,768 4,146 - 378 858.01 864.75 - 6.74 23,764 24,066 - 302 Average per day Second week: 122 . 57 123.54 - .-97 3,395 3,438 - 43 Apr. 19-20, 1905 119.06 115.01 + 4.05 3,086 3,199 - 113 Apr. 20-21, 1905 142.00 139.17 + 2.83 3,146 3,027 + 119 Apr. 21-22, 1905 126.38 119.00 + 7.38 3,752 3,720 + 32 Apr. 22-23, 1905 93.31 99.24 - 5.93 2,890 3,008 - 118 Apr. 23-24, 1905 134.81 131.72 + 3.09 4,411 4,511 - 100 Apr. 24-25, 1905 126.75 123.20 + 3.55 2,865 2,724 + 141 Apr. 25-26, 1905 Total for 2d week. . . 107.00 106.00 + 1.00 3,295 3,338 - 43 849.31 833.34 + 15.97 23,445 23,527 - 82 121.33 119.05 + 2.28 3,349 3,361 - 12 292 Influence of Inanition on Metabolism. A comparison of the computed and determiued amounts of protein shows that considerable differences appear on individual days, ranging in nitrogen metabolism experiment No. 1 from -j-13.29 grams on March 16-17 to — 15.39 grams on March 29-30. The differences are less in the second experiment, +7.38 to —5.93 grams. The average daily results for the different weeks show very much smaller discrepancies. The grand average for experiment No. 1 shows about 2 grams more per day by calculation than by actual determination, a difference of approximately 2 per cent. The agreement in the two methods of obtaining the amounts of protein in the second experiment is all that could be desired. The average amounts per day show discrepancies for 2 weeks of the experiment of less than 1 per cent. The general uniformity observed in the calculated amounts of protein when compared to the determined amounts is unfortunately not recognized so readily in the energy determinations. The variations in experiment No. 1 in the amounts of energy run from — 557 calories on March 23-24 to -j-745 on March 25-26. On the average the energy as calculated is less than that actually determined by about 160 calories per day. It is to be borne in mind, however, that the quantities of energy are, in general, very large averaging about 4500 calories per day. Thus the average discrepancy is about 4 per cent. It is interesting to note that in the majority of instances no uniformity exists between the discrepancies appearing in the protein and the energy, though on March 29-30 the large minus discrepancy (in terms of the amount calculated) in the protein corresponds with a very large minus difference in the energy. On March 16-17 also, when large discrepancies are found in both protein and energy, both results in terms of the amount calculated are positive. In the second experiment the agreement between the calculated and determined amounts of protein and energy is very satisfactory. On the whole the agreement between the amounts calculated and those determined is sufficiently close to indicate that there was no material error, either in sampling or in analysis. Feces. — The time of defecation, the weights of fresh feces, and air-dry material 84 are recorded in table 179. The copious defecations are especially noticeable. For example, on March 19 there were 349.4 grams of fresh feces passed at 9h 15m a. m., and at 5 p. m. of the same day 262.4 grams were excreted, i. e., 611.8 grams of feces were passed within 8 hours. Indeed, the next morning at 8.30 there was a defecation amounting to 312.4 grams. Within 24 hours, therefore, this subject excreted 924.2 grams of fresh feces. 84 Dried in a water oven and allowed to stand in the laboratory for 36 hours before weighing. Nitrogen Metabolism Experiments. 293 Table 179. — Feces passed — Nitrogen metabolism experiments Nos. 1 and 2. Date. Time. (a) Weight of fresh sub- stance. (b) Weight of par- tially dried sub- stance. Date. Time. (a) Weight of fresh sub- stance. (b) Weight of par- tially dried sub- stance. NITROGEN METABOLISM EXP'T No. 1. NITROGEN METABOLISM EXP'T No. 1. First week : Fourth week:'2 1905. 8h00m a.m. Grams. 221.7 Grams. 42.8 1905^ 9h40m a.m. Grams. 198.8 Grams. 43.4 Mar. 16 8 00 a.m. 108.4 23.6 5 50 p.m. 63.9 19.5 7 00 p.m. 176.6 33.5 Apr. 6 7 55 a.m. 217.7 49.0 Mar. 17 8 00 a.m. 122.9 23.3 6 00 p.m. 122.4 27.6 Mar. 18 2 30 p.m. 43.6 9.7 7 30 a.m. 179.1 41.1 Mar. 19 9 15 a.m. 349.4 64.1 Apr. 8 7 36 a.m. 141.0 33.2 5 00 p.m. 262.4 50.1 Apr. 9 3 9 30 p.m. 188.3 10.2 Mar. 20 8 30 a.m. 312.4 48.0 K 15 a.m. 20.4 10.4 Mar. 21 10 00 p.m. 8 30 a.m. 7 15 p.m. 113.5 193.7 i 94.2 27.1 46.0 26.5 Total, 9 1131.6 234.4 NITROGEN MKTARnr.TSM uyp'T isrn 9. Total, Is Second week: t week 1998.8 394.7 First week : Mar. 21 ... 7''15m p.m. 61.4 14.2 Apr. 14. . . . 8h05m a.m. 183.9 46.5 Mar. 22 8 30 a.m. 291.7 66.9 1 30 p.m. 139.7 23.3 7 30 p.m. 210 . 6 48.1 Apr. 15. . . . 8 15 a.m. 163.3 46.2 Mar. 23 8 30 a.m. 139.3 32.6 12 35 p.m. 37.8 12.1 2 00 p.m. 211.2 52.7 6 45 p.m. 66.2 14.7 7 10 p.m. 246.5 39.0 Apr. 16. . . . 9 25 a.m. 46.8 14.3 Mar. 24 8 30 a.m. 126.0 30.8 5 55 p.m. 45.1 14.1 7 00 p.m. 190.1 44.9 Apr. 17 7 20 a.m. 301.8 51.5 Mar. 25 8 30 a.m. 81.7 25.1 6 15 p.m. 49.8 11.2 6 15 p.m. 230.7 44.3 Apr. 18 8 30 a.m. 60.4 14.4 Mar. 26 9 30 a.m. 305.6 64.2 5 20 p.m. 102 . 5 29.3 7 00 p.m. 195.4 46.9 Apr. 19 8 10 a.m. 196.8 48.9 Mar. 27 8 00 a.m. 229.8 53.3 5 45 p.m. 67.9 22.0 Mar. 28 8 30 p.m. 8 30 a.m. 7 30 p.m. 74.3 262.3 100.6 20.3 52.7 28.4 Total, Is Second week : 1462.0 348.5 Total, 2d Third week : week 2957.2 664.4 Apr. 19 Apr. 20 51145m p m 7 50 a.m. 5 15 p.m. 4 31.2 165.0 96.5 9.8 33.7 25.2 Mar. 29 8h00m a.m. 160.6 35.0 Apr. 21 6 30 a.m. 92.5 20.3 7 00 p.m. 138.9 30.6 5 25 p.m. 90.2 23.5 Mar. 30 8 15 a.m. 182.9 44.5 Apr. 22 7 35 a.m. 159.4 32.7 6 00 p.m. 55.1 20.0 5 35 p.m. 64.3 19.2 Mar. 31 8 30 a.m. 39.0 12.0 Apr. 23 9 30 a.m. 118.1 30.2 2 00 p.m. 298.2 55.5 5 15 p.m. 70.6 15.5 5 30 p.m. 43.5 11.5 11 30 p.m. 26.2 6.9 8 00 a.m. 149.7 37.7 Apr. 24 7 10 a.m. 256.1 53.5 5 30 p.m. 104.5 37.4 4 55 p.m. 19.6 5.9 Apr. 2 8 00 a.m. 150.1 38.5 Apr. 25 5 30 a.m. 53.2 11.1 5 00 p.m. 83.5 20.6 10 45 a.m. 118.9 24.3 8 00 a.m. 205.9 39.1 5 25 p.m. 38.5 8.4 2 30 p.m. 22.1 10.2 Apr. 26 8 00 a.m. 162.6 37.4 Total, 3d w 8 05 a.m. 2 45 p.m. eek 98.8 77.9 29.5 21.1 Total, 2d y 'eek 1562.9 357.6 1810.7 443.2 1 Separation between this and the following amount. 2 See note 2, table 180. 3 After enema. 4 Showed lampblack from capsule taken with breakfast April 19, included with feces for week beginning April 20. 294 Influence of Inanition on Metabolism. The "dried" feces were not absolutely anhydrous, but the comparison between the weights of the fresh and dried material indicates considerable variations in the percentages of water in the feces from day to day. Thus in the 246.5 grams of fresh feces passed March 23 at 7h 10m p. m. there were but 39.0 grams of air-dry material. Allowing 5 per cent of water in the air-dry material, the water-free substance would constitute only 15 per cent of the whole amount, an unusually low percentage. There was no indication that the subject had any diarrhoea. The 20.4 grams of feces passed on April 13, at 8h 15m a. m. (second day after a 4-day fast) on the contrary, contained 10.4 grams of air-dry matter or about 50 per cent of the total. Fluctuations of similar character, though not so marked, can be observed in the water content of the feces passed during the second nitrogen metabolism experiment. Analysis of feces. — The air-dried feces for each week were ground, sampled, and analyzed. The amounts of nitrogen, sulphur, phosphorus, and the heat of combustion per gram are recorded in table 180. Table 180. — Determined weight of nitrogen, sulphur, and phosphorus, and heat of combustion of partially dried feces — Nitrogen metabolism experiments Nos. 1 and 2. Labor- atory num- ber. Date. (a) Nitro- gen.1 (&) Phos- phor- us. (c) Phos- phoric acid (P206). (d) Sul- phur. (e) Sul- phur triox- ide (SO,). (/) Heat of combus- tion per gram 3889 3890 3891 3892 3907 3908 Experiment No. 1: First week, Mar. 14 to 21 Second week, Mar. 21 to 28. . Third week, Mar. 28 to Apr. 4 Fourth week, Apr. 4 to 13. . . Experiment No. 2: First week, Apr. 122 to 19. . . . Second week, Apr. 19 to 26. . Grams. 16.42 27.17 19.50 10.14 17.22 16.52 Grams, 6.16 8.52 6.06 3.26 6.83 6.08 Grams. Grams. Grams. 14.11 1.50 3.75 19.51 2.49 6.21 13.88 1.79 4.47 7.46 .98 2.44 15.65 1.57 3.93 13.91 1.46 3.63 Calories, 5.094 5.146 5.185 5.059 4.779 4.785 1 The per cents of nitrogen in feces for the different weeks of the experiments were as fol- lows : Experiment No. 1, first week, 4.16 per cent ; second week, 4.09 per cent ; third week, 4.40 per cent ; fourth week, 4.32 per cent. For experiment No. 2, the corresponding per cents were 4.94 and 4.62. 2 The separation of feces from nitrogen metabolism experiment No. 1 was not obtained until April 13. The second experiment, however, began on April 12, when food was ingested. No attempt was made to determine the water in the feces, as the subject was not within the respiration chamber and consequently the water balance could not be determined. The analyses of the feces were accordingly made on the air-dry material. Amounts of fecal nitrogen excreted per day. — Although it is obviously impossible to separate the feces for each experimental day, it is of advantage Nitrogen Metabolism Experiments. 295 to estimate as nearly as possible the quantity of fecal nitrogen excreted each day in order to compute the daily gains or losses of nitrogen to the body. Separation of the feces was made each week. In the computations of the amount of daily fecal nitrogen recorded in table 181 it is assumed that the Table 181. — Calculated daily excretion of fecal nitrogen — Nitrogen metabolism experiments Nos. 1 and 2. Date. (a) (6) Dai|y | Calcu- propor- V t;, tionof £Ji?J total ^y nit™- ' ™ r eTef -tro- week. Sen- Date. (a) Daily propor- tion of total nitro- gen in- gested per week. (&) Calcu- lated daily excre- tion of fecal nitro- gen. Experiment No. 1 . First week: Mar. 14-15 Per cent. 13.9 17.0 14.6 10.1 17.8 11.6 15.0 Grams. 2.28 2.79 2.40 1.66 2.92 1.91 2.46 Fourth week: ADr. 4-5 Per cent. 22.4 28.6 28.7 20.3 Grams. 2.27 2.90 2.91 2.06 Apr. 5—6 Mar. 15-16 Apr. 6-7 Mar. 16-17 Apr. 7-8 Mar 17-18 Average per day .... Experiment No. 2. First week: Apr. 12-13 Apr. 13-14 Apr. 14-15 Mar. 18-19 100.0 10.14 2 54 Mar. 19-20 Total, 1st week Average per day. . . . Second week: Mar. 21-22 5.2 11.8 14.6 17.3 15.8 21.1 14.2 0.90 2.03 2.51 2.98 2.72 3.63 2.45 100.0 16.42 | 2.35 14.5 13.0 9.9 14.1 13.6 16.8 18.1 3.94 3.53 2.69 3.83 3.70 4.56 4.92 Apr. 15-16 Mar. 22-23 ADr. 16-17. . Mar. 23-24 Apr. 17-18. . Mar. 24-25 Apr. 18-19 Mar. 25-26 Total, 1st week Average per day. . . . Second week: Apr. 19-20 Mar. 26-27 100.0 17.22 9 4fi Mar. 27-28 Total, 2d week Average per day Third week : Mar. 28-29 100.0 27.17 3.88 14.0 16.7 14.9 11.0 15.9 14.9 12.6 2.31 2.76 2.46 1.82 2.63 2.46 2.08 Apr. 20-21 13.7 10.5 9.7 20.3 18.6 15.7 11.5 2.67 2.05 1.89 3.96 3.63 3.06 2.24 Apr. 21-22 Apr. 22 23 Mar. 29-30 Apr. 23-24. . Mar. 30-31 Apr. 24-25. . Mar.31-Apr. 1 Apr. 1-2 Apr. 25-26 Total, 2d week. Average per day. . . . Apr. 2-3 100.0 16.52 2.36 Apr. 3-4 Total, 3d week Average per day 100.0 19.50 2.79 fecal nitrogen was proportional to the nitrogen ingested each day. The total nitrogen ingested each week was taken as 100 per cent and the proportion of this amount ingested each day calculated. These values are recorded in the first column of the table. The total nitrogen of the feces for the week was then 296 Influence of Inanition on Metabolism. © A < s Date. (a) i . ©^ -a .5 o a O S © 1— 1 (W © © a h- 1 In urine (cal- culated) "o (2VX9.0O). "" (d) s ©* cs nY Experiment No. 1. First week: Mar. 14-15. .. . Mar. 15-16... . Mar. 16-17... . Mar. 17-18.. . . Mar. 18-19.. . . Mar. 19-20... . Mar. 20-21.... Total, 1st week Second week : Mar. 2 1-22.... Mar. 23-24 Mar. 25-26. . . . Mar. 26-27.... Mar. 27-28. . . . Total, 2d week Third week : Mar. 28-29 .... Mar. 29-30 .... Mar. 30-31 .... Mar.31-Apr.2 [ Apr. 2-3 Apr. 3-4 Cals. 5,592 6,771 4,857 3,943 6,527 4,810 6,200 Ccds. 218 291 119 49 582 383 369 Cals. 2113 119 100 57 3 86 107 86 Cals. 5,261 6,361 4,638 3.837 5,859 4,320 5,745 Fourth week : Apr. 4-5 Apr. 5-6 Apr. 6-7 Apr. 7-8 Total, 4 days. . Experiment No. 2- First week: Apr. 12-13... . Apr. 13-14... . Apr. 14-15. . . . Apr. 15-16. . . . Apr. 16-17.... Apr. 17-18.... Apr. 18-19.... Total, 1st week Second week: Apr. 19-20.... Apr. 20-21.... Apr. 21-22.... Apr. 22-23.... Apr. 23-24.... Apr. 24-25. . . . Apr. 25-26. . . . Total, 2d week Cals. 3,395 4,640 4,208 2,752 Cals. 318 388 208 272 Cals. 114 104 6107 102 Cals. 2,963 4,148 3,893 2,378 14,995 1186 427 13,382 1,334 .... 2,806 334 3,573 349 3,300 136 3,362 300 5,621 , 209 3,768 J 339 38,700 2011 668 36,021 7100 105 1,234 2,367 5,611 6,092 4,598 4,847 6,786 4,752 6,507 665 640 390 357 572 379 417 92 84 72 78 4 108 101 131 4,854 5,368 4,136 4,412 6,106 4,272 5,959 136 125 149 161 114 3,088 3,039 2,913 5,251 3,315 23,764 1667 890*21,207 3,086 329 3,146 | 210 124 128 109 100 96 120 110 2,633 2,808 3,395 2,538 4,030 2,535 3,006 39,193 3420 666 35,107 3,874 3,440 3,446 4,357 4,356 3,798 3,847 340 334 410 389 1 306 f 256' 262 113 83 81 5235 115 91 3,421 3,023 2,955 7,783 3,427 3,494 3,752i 2,890 4,411 2,865 3,295 248 252 285 210 179 23,445 1713 787 20,945 Total, 3d week 27,118,2297 1 718 24,103 It has been assumed that the feces for a given day result from the food of the previous day ; the feces from which the energy here given was calculated were excreted on the day following the one to which they are here assigned. Compare table 170, p. 293. 'Determined, 111 calories. 4 Determined. 115 calories. " Determined, 99 calories. Determined, 105 calories. 6 Determined, 244 calories. 7 Determined, 109 calories. 298 Influence of Inanition on Metabolism. Table 183. — Determinations in urine — Nitrogen metabolism experiments Nos. 1 and 2. Date. (a) Weight of urine. Spe- cific gravity (0 (d) Vol- ume (a+b). Nitro- gen. (e) Phos- phorus3 Phosphoric acid (P206>. , By fusion. titration Experiment No. 1. Mar. 14-152 Mar. 15-163 Mar. 16-174 Mar. 17-18 Mar. 18-19 Mar. 19-20 Mar. 20-21 Grams. 807.1 769.8 1.0183 1.0242 1,460.8 1.0168 1,072.2 1,818.5 1.0162 1.0153 1,669.9 1.0172 1,667.7 1.0167 Total, 1st week. . Average per day. 9,266.0 1,323.7 c.c. 793 752 1,437 1,055 1,791 1,642 1,640 Grams. 12.61 13.18 11.07 6.31 9.55 11.89 9.55 9,110 74.16 l,30li 10.59 Grams. 0.642 .952 .744 .397 .867 .987 .895 Grams. 5.484 .783 Mar. 21-22 s. Mar. 22-23 \ Mar. 23-24.. Mar. 24-25. . Mar. 25-26. . Mar. 26-27. . Mar. 27-28.. 1,312.31 1,515.9' 1,243.1 1,349.4 1,133.6 913.5 1,229.5 Total, 2d week. . . Average per day. Mar. 28-29 Mar. 29-30 Mar. 30-31 Mar.31-Apr.2 Apr. 2-3 Apr. 3-4 8,697.3 1,242.5 .0213 1.0212 1.0210; 1.0178 1.0245 1.0283 J 1.0261 1,285 1,484 1,218 1,326 1,106 888 1,198 8,505 1,215 10.17 9.35 8.02 8.63 11.95 11.26 14.53 73.91 10.56 1.149 1.084 .993 1.072 1.166 1.295 1.412 171 167 1,880.0 1,600.2 1,464.1 3,534.9 1,910.7 1,596.7 Total, 3d week. . . Average per day. 11,986.6 1,712.4 1.0142 1.0144 1.0144 1.0168 1.0144 1.0145 l,854i 1,577 1,443 3,477 1,884| 1,574 12.54 1.559 9.27jl.l00 9.001.150 26.14 2.908 12.77 10.11 11,809' 1,687 79.83 11.40 1.510 1.240 9.467 1.352 Apr. 4-5 1,622.4 Apr. 5-6 1,433.1 Apr. 6-7. Apr. 7-8. Total, 4 days Average per day. 1,780.8 1,134.3 5,970.6 1,492.7 1.0126 1,602 1.0152 1,412 1.0142 1,756 1.0200 1,112 12.65| 1.280 11.561.350 11.87 1.251 11.36 .944 5,882 47.44 4.825 l,47ll 11.86 1.206 470 180 704 908 987 261 050 Grams. 1.656 2.196 1.268 1.086 1.978 2.322 2.118 561 12.624 794 ! 1.803 2.631 2.482 2.274 2.456 2.670 2.966 3.234 118.713 I 2.673 3.571 2.513 2.635 6.660 3.459 2.840 121.678 3.097 866 2.934 3.093 2.992 2.162 11.181 2.795 Experiment No. 2. Apr. 12-13 7 89 1 . 0 1 . 0160 Apr. 13-14 8 1,006.8 1 .0173 Apr. 14-15 1 1,011 . 7 1 . 0203 Apr. 15-16 1,426.2 1 .0136 Apr. 16-17 2,296.2 1 .0121 Apr. Apr. 17-18.. 18-19" Total, 1st week. . Average per day. 2,440.5 1.0153 1,834.9:1.0129 10,907.3 1,558.2 877 990 992 1,407 2,269 2,404 1,812 10,751 1,536 11.14 0.652 1.493 11.65 .499 1.142 15.08 .971 2.225 13.89 .956 2.189 16.53 1.307 2.994 17.93 1.432 3.280 12.72:1.079 2.472 98.94!6.896 15.795 14.13 .985 2.256 1.671 1.375 2.458 2.067 2.731 3.752 2.799 16.853 2.408 1 Phosphorus from results by titration when fusion not available. 2 Total creatinine, 1.566 grams ; preformed creatinine, 1.493 grams ; creatine, 0.073 gram. 8 Preformed creatinine, 1.433 grams. ° Preformed creatinine, 1.487 grams. 4 Preformed creatinine, 1.251 grams. 8 Preformed creatinine, 1.523 grams. 'Total creatinine, 1.450 grams; preformed creatinine, 1.407 grams; creatine, 0.043 gram. * Total creatinine, 1.420 grams ; preformed creatinine, 1.384 grams ; creatine, 0.036 gram. Total creatinine, 1.334 grams ; preformed creatinine, 1.300 grams ; creatine, 0.034 gram. Nitrogen Metabolism Experiments. 299 Table 183. — Determinations in urine — Continued. Date. Experiment No. #— Cont'd Apr. 19-20 Apr. 20-21 Apr. 21-22 Apr. 22-23 Apr. 23-24 Apr. 24-25 Apr. 25-26 Total, 2d week Average per day (o) Weight of urine. (6) (c) Specific Volume gravity, (a-z-b). I Grams. 1,840.0 1,937.0 1,896.4 2,417.9 1,497.1 2,189.2 1,674.2 13,451.8 1,921.7 1.0148 1.0142 1.0131 1.0112 1.0143 1.0136 1.0155 c.c. 1,813 1,910 1,872 2,391 1,476 2,160 1,649 (d) Nitro- gen. M Phos- phorus1 13,271 1,896 Grams. 13.80 14.181 12.09 11.13 10.66 13.30 12.19 Grams. 1.141 1.100 1.303 1.464 .994 1.629 87.35 8 7.631 12. 48 8 1.272 Phosphoric acid (P2Os). (f) to) By By ision. titration Grams. 2.612 2.521 2.984 3.354 2.277 3.732 Grams. 2.946 2.972 3.249 3.202 2.460 3.372 3.743 17.48021.944 2.9131 3.135 Date. (ft) Sulphur. Sulphur trioxide ■ (S03). (*) Total. t 0'} • Inorganic and ethereal. (*) Neutral. (J) Chlorine. Cm) Sodium chloride. Experiment N"o. 1. Mar. 14-152 Mar. 15-163 Mar. 16-174 Mar. 17-18 Mar. 18-19 Mar. 19-20 Mar. 20-21 Total, 1st week Average per day. . . Apr. 6-7 Apr. 7-8 Experiment No. 2. Apr. 12-13 5 Apr. 13-14 6 Apr. 14-15 Apr. 15-16 Apr. 16-17 Apr. 17-18 Apr. 18-19' Total, 1st week. . Average per day. Apr. 19-20. Apr. 20-21. Apr. 21-22. Apr. 22-23. Apr. 23-24. Apr. 24-25. Apr. 25-26. Total, 2d week. . , Average per day. Grams. 0.977 .872 .650 .451 .736 .752 .709 Grams. 2.440 2.176 1.622 1.126 1.838 1.876 1.769 5.147 .735 12.847 1.835 Grams. 2.174 1.854 1.411 .867 1.496 1.634 1.508 Grams. 0.266 .322 .211 .259 .342 .242 .261 Grams. 1.181 1.804 7.980 5.010 6.225 8.664 7.570 10.944 1.563 1.903 .272 38.434 5.491 Grams. 1.948 2.977 13.171 8.272 10.275 14.296 12.490 63.429 9.061 0.764 1.906 1.647 1.565 0.259 0.812 .967 1.010 .826 1.016 1.023 .814 6.468 .924 2.026 2.412 2.524 2.063 2.536 2.555 2.033 1.803 2.179 2.158 1.853 2.384 2.313 1.877 16.149 2.307 14.567 2.081 0.224 1.725 .233 4.075 .366 2.313 .209 2.117 .152 4.807 .242 10.062 .156 4.562 1.582 29.661 .226 4.237 2.847 6.724 3.816 3.494 7.932 16.605 7.529 48.947 6.992 .772 1.067 .788 .790 .681 .947 8 5.045 8 .841 1.927 2.662 1.968 1.973 1.700 2.364 1.753 2.339 1.751 1.760 1.639 2.276 1.997 812.594l 13.515 8 2.099' 1.931 0.174 4.025 .323 4.538 .217 .... .213 .... .061 .... .088 .... .... .... 8 1.076 8 .179 .... 6.642 7.487 1 Phosphorus from results by titration when fusion not available. 2 Total creatinine, 1.566 grams ; preformed creatinine, 1.493 grams ; creatine, 0.073 gram. 8 Preformed creatinine, 1.433 grams. * Preformed creatinine, 1.251 grams. 6 Total creatinine, 1.450 grams ; preformed creatinine, 1.407 grams ; creatine, 0.043 gram. 6 Total creatinine, 1.420 grams ; preformed creatinine, 1.384 grams ; creatine, 0.036 gram. 7 Total creatinine, 1.334 grams ; preformed creatinine, 1.300 grams ; creatine, 0.034 gram. 8 Six days only. 300 Influence of Inanition on Metabolism. PART 3. DISCUSSION OF RESULTS. The data accumulated in the experiments made on fasting men in this laboratory furnish material for a general discussion of the subject of the influence of inanition on metabolism. Of the numerous questions in physiology upon which these results throw light, it will obviously be impossible in this volume to discuss any except those which have more specifically to do with the general subject of metabolism. .Numerous important subjects in metabolism, such as the protein requirement of the body and the relation of the active mass of protoplasmic tissue to the heat production, must be deferred for subsequent discussion. Certain aspects of the influence of inanition on metabolism, however, may with propriety be dealt with now and the following discussion is designed to present all the information thus far obtained regarding the various phases of physiological activity upon which the data throw light. Literature.1 — Where reference to the earlier literature on experiments with fasting men is made, the articles given in the following list will be indicated by the number preceding each title, as for example, Luciani (4). (1) On the body-weight and urea in a case of starvation. David Nicholson. Brit. Med. Journ. (1870), 1, p. 4. (2) Metabolism in fasting. A. Sadovyen. Russian Society of General Hygiene (1887-1888), 12, pp. 13-76. (3) Observations on the metabolism of man during starvation. D. Noel Paton and Ralph Stockman. Proceedings of the Royal Society of Edinburgh (1888- 1889), 16, p. 121. (4) Das Hungern. Luigi Luciani (1890). Leipzig. (5) Die Eiweisszersetzung beim Menschen wahrend der ersten Hungertage. W. Prausnitz. Zeitschrift f. Biologie (1892), 29, p. 151. (6) II ricambio materiale e la tossicita dell' urina nell' inanizione dell' uomo. G. Ajello & A. Solaro. La Riforma Medica (1893), Anno ix, 2, p. 542. (7) Untersuchungen an zwei hungernden Menschen. Curt Lehmann, Friedrich Mueller, Immanuel Munk, H. Senator, N. Zuntz. Archiv f. pathologische Anatomie u. Physiologie u. f. klinische Medicin (1893), 131, Supplementheft. (8) Metabolism during fasting in hypnotic sleep. C. F. Hoover and Torald Soil- man. Journ. Exper. Med. (1897), 2, p. 403. (9) Beitrage zur Kenntniss des Stoffwechsels beim hungernden Menschen. J. E. Johansson, E. Landergren, Klas Sonden, und Robert Tigerstedt. Skand. Archiv f. Physiologie (1897), 7, p. 29. (10) Beitrage zum Stoffwechsel im Hungerzustande. E. Freund und O. Freund. Wiener klinische Rundschau (1901), 15, pp. 69-71 and 91-93. (11) Beobachtungen iiber die Kreatininausscheidung beim Menschen. C. J. C. Van Hoogenhuyze and H. Verploegh. Zeitschrift fur physiologische Chemie (1905), 46, p. 415. (12) Eiweisszerfall und Acidosis im extremen Hunger mit besonderer Beriick- sichtigung der Stickstoffvertheilung im Harn (nach Untersuchungen an dem Hungerkiinstler Succi). Theodor Brugsch. Zeitschrift f. expt. Pathologie u. Therapie (1905), 1, p. 419. 1 The excellent article by C. von Noorden on Fasting and Chronic Starvation (Metabolism and Practical Medicine, Vol. II, Pathology, pp. 1-61; Keener & Co., Chicago, 1907) contains much of interest regarding metabolism during inanition. Body-Weight. 301 Experiments During Inanition, body-weight. Aside from the general observations on fasting individuals, such as the visible loss of flesh, strength, etc., perhaps the earliest measurement was that of the loss in body-weight. Since in fasting the body subsists upon its own tissues, it persistently loses in weight. It is to be regretted that in the majority of the earlier experiments, more careful consideration was not given to the weighings, accuracy of the scales used, uniformity in the amount of clothing worn, etc. The records of loss in body-weight from day to day are therefore in most instances very unsatisfactory, although in a few cases they were seemingly made with care. The extreme accuracy of the weighing apparatus used in this laboratory tends to make the weighings of more than ordinary value. Moreover, verification of the changes in body-weight was secured in the experiments reported in the preceding sections by comparing them with the balance of income and outgo. It is therefore reasonable to suppose that the records of body-weight and changes of body-weight here given are much more accurate than any earlier records. In instances of so-called complete fasting, i. e., where no drinking-water is consumed, the loss in body-weight might be expected to be larger than in those experiments in which water was taken, although the drinking of large quantities of water is almost immediately compensated by the voiding of large quantities of urine. In the complete fasting experiments, the only intake is the oxygen of the air. This intake varies from 500 to 600 grams per day, according to the muscular activity and size of the subject. Since this oxygen combines with carbon and hydrogen in the body, and with the disintegration of the body tissue there is a consequent loss of water, it is to be expected that the loss in body-weight during complete fasting will be not far from 700 to 1000 grams per day. Experiments on animals have shown that with mammals, birds, amphibia, and fishes, the loss reached 40 per cent of the original weight before death ensued. Although numerous cases of fasting insane and pathological instances of long fasting are recorded, no satisfactory evidence has been accumulated to show the extent of the loss which the body can undergo before death ensues in the case of men. Apparently authentic instances are recorded, however, of professional f asters who succeeded in fasting from 30 to 45 (?) days and again recovered their usual health. During these latter tests of endurance, water was taken in every instance. Possibly the most accurate of the earlier studies of loss of body-weight durino- complete fasting is that of Laun ' in an experiment on himself lasting 21^ hours. From the completion of the evening meal at 10 o'clock February 19 1 Untersuchungen zur Naturlehre des Menschen und der Thiere. Herausgegeben von J. Moleschott. Frankfurt a. M. (1857), p. 278. 302 Influence of Inanition on Metabolism. until 5h 30ra a. m., February 21, neither food nor drink was taken. He weighed himself at 7h 30m a. m., February 20, and again at 5 a. m., February 21. During this period the body-weight was reduced from 77.64 kg. to 75.77 kg., that is, in 21£ hours there was a loss in weight of 1.87 kg. The author calculates that for 24 hours the total loss in weight would have amounted to 2.09 kg. and that if he had weighed himself immediately after the evening meal on the 19th, the loss for the entire period would have been 2.34 kg. The balance used was accurate to 45 grams. In the three 24-hour fasting experiments reported by Pettenkofer and Voit * the losses in body-weight were 930, 660, and 680 grams, respectively. In the last experiment work was performed. Eanke 8 lost 1.130 grams in weight in a 24-hour fast with no water. Obser- vations began 24 hours after the last meal, and consequently corresponded to those on the second fasting day. Nicholson (1) made observations on a fasting prisoner whose body-weight at the beginning was 107£ pounds and 100^ pounds on the sixth day, after a small portion of food had been taken. According to the statement of the author, there was absolute fast from the commencement up to the fifth day, during which period there was a loss of 7 pounds (3.2 kg.). At the end of the third day the loss was 5 pounds or an average loss of 1.7 pounds per day. On the fourth day the subject lost 1 pound and during the fifth day the author estimates that he lost 1 pound although weighings were not made. Assuming a loss of 1 pound on the fifth day, there was an average daily loss of 1.4 pounds during the period of starvation, the greatest loss appearing during the first part of the experiment. Miiller4 reports a case of fasting caused by oesophagus stenosis, in which the body-weight on the sixth of July was 34.5 kilos, and on the ninth of July was 33 kilos. These 4 days, which represented the fifth to eighth day of complete inanition, resulted, therefore, in the loss to the body of 1.5 kg. The body-weight of the subject, however, was much below the normal. Schaefer * reports the body weights of a number of insane persons who fasted. One woman 51 years old, weighing about 68 kg., fasted from March 3 to March 8, but drank 500 cc. of water per day. On March 6, the body-weight was 63.3 kg. and on March 7 it was 63.3 kg. A second case was that of a woman 57 years old, weighing 56 kg., who fasted from March 19, drinking only a little water and clear cold coffee. On March 23, the body-weight was 49.3 kg., on the 24th, 48.8 kg., and on the 25th, 48 kg. Another instance was that of a woman weighing 41.2 kg., who fasted from April 28 until May 4. No water nor food was consumed. On the second of May the body-weight was 38.7 kg., and on May 3, 37.85 kg. 2Zeit. f. Biol. (1866), 2, p. 479. 8 Die Ernahrung des Menschen (1876), p. 211. 69.3 K. A. H. M. Experi- ment 81.3 62.0 K. H. C. K. Experi- ment 82.3 71.5 K. H. R. D. Experi- ment 83.3 55.6 K. N. M. P. Experi- ment 85.3 67.6 K. D. W. Experi- ment 89/ 79.1 K. First day. . . Second day. Per ct. Per ct. Per ct. Per ct. Per ct. Per ct. 1.90 2.49 1.56 0.85 0.97 2.22 4.11 5.28 3.56 2.98 2.52 3.44 Per ct. 1.90 3.14 1 Beginning weight assumed from weigh- ing of later date, includes underclothing. 3 Weights include underclothing. 8 Weights without clothes. 4 Weights include all clothing. 5 Weights without clothes. water of respiration and perspiration, and carbon dioxide, but also by the water ingested. Furthermore, it is highly probable that temperature, muscular activity, state of bodily condition and other more or less obscure factors all contribute toward the balance between income and outgo in such a manner as to cause the marked fluctuations in body-weight appearing on different days. Hence it is obvious that losses in body-weight in experiments of but few days duration are wholly without significance. With regard to the total cumulative loss as the experiment progresses, it appears that in the long experiments of Succi the loss bears in general a direct ratio to the length of the experiment. Beitrage z. Physiol., Hyg., Pharm. u. Toxikol. (1875). Body Temperature. 311 BODY TEMPERATURE. Since muscular activity has a very pronounced and immediate effect on the body temperature, experiments during fasting when the internal muscular activity due to peristalsis is at a minimum are of especial interest. In con- nection with the earlier fasting experiments made in this laboratory/ obser- vations regarding body temperature were made and in some instances curves were drawn showing the body temperature for 24 hours. These curves together with a large number of other curves of body temperature were published accompanied by a special discussion of the fluctuations in body temperature.10 The most noticeable effect of fasting on the normal body temperature curve seems to be a flattening of the curve during the waking hours, although in the few experiments made, the evening fall especially after retiring and the early morning rise were, if anything, more marked than the normal temperature fluctuations at the same hours. The night curves for a number of fast days were also studied, and the general conclusion was that fasting results in a slight lowering of the average body temperature, amounting, however, to but a few tenths of a degree. This is in harmony with the observations of Jurgensen n who, not only finds the average body temperature somewhat lower during fast, but also notes a smaller amplitude of the daily curve. Although no temperature observations were recorded by Nicholson (1) he states that there was no fever during the fast of the prisoner under his care. In the fast made by Tanner,12 no fall in temperature was noticed. The tem- perature taken in the mouth was 36.9° on the 25th day and 37.1° on the 30th day. As has been pointed out previously, this fast was very imperfectly reported and its authenticity is somewhat in doubt. Noyes 13 observed a temperature of 34.4° in an insane man who had consumed no food for 45 days. Pembrey " points out in commenting on this, that this subject was paralyzed in the lower limbs. In the report of the observations made by Schaefer 15 on the fasting insane it is stated that the body temperature was normal in all cases. Paton & Stockmann (3) record that during the fast of Jacques his temper- ature was invariably subnormal ranging from 35.6° to 34.1° C. During a slight attack of gout, on the 16th day, the temperature ranged from 36.0° to 36.6°. During the fasting experiment with Cetti (7) in Berlin, the temperature observations were made in the axilla twice a day. One observation is recorded on the day preceding the fast and one on the following food day. The average body temperature for the 10 fasting days is 36.7°, which is, however, somewhat higher than the observations made before and after the experiment. »U. S. Dept. of Agric, Office of Expt. Sta. Bui. 136 (1903). "Benedict & Snell. Archiv f. d. ges. Physiol. (1902), 90, p. 33. u Die Korperwarme des Gesunden Menschen, Leipsic, 1873. 12 British Medical Journ., London (1880), 2, p. 171. "British Medical Journal, London (1880), 2, p. 557. "Schaefer's Physiology, 1, p. 810. "Allgem. Zeit. f. Psychiatrie (1897), 53, p. 525. 312 Influence or Inanition on Metabolism. With Breithaupt, the second subject of the Berlin experiments (7) the temperature was taken, presumably in the axilla, morning and evening. Xo records were shown for the day with food preceding the fast, but temperature records were given for the first food day after the fast. The tendency for the temperature to decrease slowly as the fast progresses is noted, but after the resumption of the ingestion of food it was not overcome. The body temperature of the subject J. A. (9) was taken every 2 hours during the waking period. This is true for both the fasting and the food periods. The average temperature for the 3 days with food was 37.39°, while the average temperature of 5 days fasting was 37.23°. The average daily temperatures of the 5-day fast were 37.20°, 37.27°, 37.22°, 37.13°, and 37.23°, respectively. Body temperature observations were made by Hoover and Sollman (8) on their hypnotic subject during the 8 days of the experiment. It was not stated how the observations were made. A slight tendency for the average temper- atures to fall as the fast progressed is noticeable and in general the evening fall and morning rise are readily recognized on the temperature curve. The average temperatures for the succeeding days of the fast were: 36.6°, 36.8°, 36.7°, 36.6°, 36.3°, 36.4°, 36.3°, 36.2°. In only one of the longer fasts made by Succi were the temperature fluctua- tions accurately observed and recorded. Luciani (4), in the Florence fast, made temperature observations in the left axilla throughout the fast. The highest temperature recorded was on the 23d day, 37.25°. The lowest temper- ature was on the seventh day, 36.2°. No tendency for the temperature to fall as the fast progressed was observed. The newspaper report of the medical bulletin issued by the attending physicians on the 45th clay of the uncertain New York fast states that the temperature was normal. No temperature records of this fast have been published. In considering many of the earlier temperature observations on fasting subjects it is extremely difficult to give suitable value to the different obser- vations owing to the fact that little regularity is exhibited by different observers as regards the method of taking body temperatures. It is of very great importance to know whether the temperature was taken in the mouth, axilla, or rectum. Unfortunately, in many of the cases recorded, the normal body temperatures of the subjects, i. e., when subsisting on an ordinary diet, were not recorded and hence the data are missing for an accurate study of the question as to whether fasting actually lowers the average body temperature. The body temperatures recorded in the Middletowh experiments were ordi- narily taken in the rectum by means of an electrical resistance thermometer." 16Archiv f. d. ges. Physiologie (1901), 88, p. 492. Body Temperature. 313 In a few instances the temperature observations were taken by clinical thermometers, in which case they are noted in table 190, and in no cases are they included in the averages. The temperature measurements are based upon the absolute international hydrogen thermometer scale. In the majority of experiments here reported, temperature observations by means of the rectal thermometer were continuously obtained. Contrary to the experience of practically all the fasting subjects, S. A. B., the subject of the longer experiments, was extremely uncomfortable when wearing the rectal thermometer. Unfortunately, therefore, the records during the longer fasts were but imperfectly taken. For the specific purpose of determining the total heat production, the body temperature at the end of each experimental period is of especial importance. Eeserving for discussion in another publication the temperature fluctuations from hour to hour and from minute to minute, only the actual temperature observations at the end of each experimental period are recorded in this place. The results are presented in table 190, and for purposes of comparison the few observations made during experiments with food are also given. Under normal conditions the maximum temperature occurs late in the afternoon. The temperature then falls rapidly during the evening and reaches the minimum between 3 and 5 o'clock in the morning, after which it rises rapidly for 2 or 3 hours." The results obtained during the fasting experiments here reported show that even during fasting the body temperature generally follows the normal course. While the difference between the minimum and maximum temperature for the day with normal individuals with food is about 1° C, in the majority of the fasting experiments here reported this amplitude is very much less.18 The average difference for all Middletown fasting experi- ments is not far from 0.5 degree. This is wholly in accord with the obser- vations noted previously in this laboratory. The general effect of fasting on the absolute body temperature can best be studied in the longer fasting experiments, for if the contention is made that as the fast progresses there is a tendency for the body temperature to fall, this should be especially notice- able in fasts of long duration. An examination of the average temperatures for the day shows that there is no such tendency exhibited during the experi- ments here reported, although the verification of the statement is not easy owing to deficient data. It is clear, however, that in experiments of 7 days or less there is no indication that fasting lowers body temperature. This is in agreement with the observations made by Paton and Stockman (3) on 17 Benedict & Snell. Loc. cit. 18 On the 6th and 29th days of Succi's Florence fast, Luciani made temperature observations in the axilla from hour to hour during the day and night. The fluctuations thus measured are not far from the normal. As Luciani states, how- ever, Succi's sleep was much disturbed by the observers. 314 Influence of Inanition on Metabolism. Table 190. — Body temperature by rectal thermometer — Metabolism experiments with and without food. Body Tempeeatdre. 315 Table 190. — Body temperature by rectal thermometer — Continued. Experiment number, subject, and date. 7 a.m. 9 a.m. 11 a.m. 1 p.m. 3 p.m. 6 p.m. 7 p.m. 9 p.m. 11 p.m. 1 a.m. 3 a.m. 5 a.m. Aver- age. 79. H. E. S. Oct. 14-15, 1905 Average per day. . 80. C. R. Y. Oct. 37-38, 1905 Oct. 38-29, 1905 Average per day . . 81. A. H. M. Nov. 21-22, 1905 Nov. 22-23, 1905 Average per day.. 83. H. C. K. Nov. 24-25, 1905 Nov. 25 26,1905 Average per day. . 83. H. R. D. Dec 5-6, 1905 °C. 36.55 37.17 °C. 36.67 36.95 °C. 36.79 36.92 °C. 37.11 37.13 36.90 37.26 °C. 37.33 37.38 °C. 37.35 37.46 °C. 37.10 37.53 37.08 37.11 °G. 37.13 37.30 36.85 37.24 37.08 36.90 °C. 36.99 37.19 36.86 36.81 36.86 37.12 37.08 1 37.31 37.36 37.32 37.10 37.22 37.05 36.99 37.09 36.86 37.31 37.03 37.28 36.98 37.17 36.78 37.18 36.83 37.33 37.00 36.95 37.58 37.65 36.93^ 37.00 37^ 37.46 I 37.02; 37.03 37.74 37.54 37.16 37.56 36.96 37.43 37.09 37.16 37.08 36.98 37.08 37.39 37.30 37.17j 37.23 37.38 37.29 37.36 37.30 36.39 36.70 36.59 36.70 36.18 36.65 35.97' 36.80 36.53 36.76 36.45 36.90 36.60 37.18 36.29 36.64 36.02 36.82 36.02 36.14 36.81 36.82 36.27 36.78 36.31 36.77 36.55 36.65 36.42 36.25' 36.78 1 36.68 36.89 36.47 36.42 36.42 36.48 36.63 36.65 1637.20 37.30 36.96, 37.02 36.81 1636.91 37.06 36.94 37.03 36.69 36.94 36.80 37.04 36.88: 36.68 37.36 37.00 36.41 36.66 36.49 36.92 36.70 37.16 36.82 37.00 37.08, 37.16 36.81 36.99 36.99 36.83 36.93 37.12 36.84 36.53 36.71 36.93 36.91 36.58 36.92 36.83 36.86 36.99 36.65 36.71 36.81 36.46' 36.50 36.71 36.93 36.20 36.64 36.51; 36.33 36.5i; 36.19 36.37 36.35 36.31 36.19 36.49 36.43 36.53 36.60 Dec. 6-7, 1905 Average per day.. 85. N. M. P. Dec. 10-11, 1905 Average per day. . 89. D. W. Jan. 10-11, 1906 Jan. 11-12, 1906 Average per day. . Average of all fast- ing experiments.. Average of all food 36.75 36.85 36.82 36.76 i 36.59 36.72 36.43 36.51 36.26 36.36 36.25 36.46 36.56 36.33 36.65 36.80! .... 36.57 "36.61 36.45 36.74 36.50 36.93 36.87 36.71 37.00] 37.33 36.84 37.32 36.47 36.96 36.20 37.00 36.13 37.00 36.45 37.17 36.53 36.96 36.56 36.65 36.59 36.60 36.61 | 36.97 37.05 37.08 36.72 36.60 36.57 36.81 36.73 35.99 36.33 36.41 35.87 36.16 35.88 36.41' 36.18 36.45 36.33 36.27 36.13 36.45 36.10| 36.37 36.19 36.09 35.99 35.85 35.77 35.66 35.96 1835.62 36.13 | 36.09 35.99 36.37 36.02 36.01 36.43 36.14 36.41 36.26 36.18 35.92 35.72 35.78 36.10 36.70 36.73 36.76 37.00 36.67 37.14 36.70 37.32 36.77 37.53 36.79 37.63 36.88 37.63 36.74 37.49 36.49 37.44 36.43 37.08 36.40 36.98 36.56 36.91 36.66 37.24 1 Reading by clinical thermometer in the mouth. Not included in average. 2 Record at 7" 30m a. m. 8 Record at 7h 12m p. m. * Record at 7" 15m a. m. 5 Record with clinical thermometer 97°. 2 F. in the mouth = 36.22 C + .47 = 36.69 by rectal. 8 The only obtainable records for experiment 74 were those taken in the mouth with the clinical thermometer at 5 p. m. each day of the experiment. These were respectively as follows : Februarv 2-3, '05, 36.56 ; February 3-4, '05, 36.56 ; February 4-5, 37.00. T Record at 9h 16m a. m. 8 Record at l" 12m a. m. 9 Record at 9h 20m a. m. 10 Record at 9h 24™ p. m. "Record at 9* 36™ p. m. 12 Record at 6h 30™ and 7" 30™ p. m., with clinical thermometer in the mouth. Not included in average. 13 Record at 8 a. m., with clinical thermom- eter in the mouth. Not included in average. 14 Record at 5h 25™ p. m., with clinical thermometer in the mouth. Not included in average. 15 Record at 7h 30™ a. m. 18 Record at lh 44™ p. m. 17 Record at llh 12™ a. m. 18 Record at 5h 28™ a. m. 316 Influence of Inanition on Metabolism. Jacques, and Luciani (4) and Daiber 19 on Succi, but contrary to the conclusion drawn by Hoover & Sollman (8), Senator & Mueller (7) studying Breithaupt, Jurgensen, and Benedict & Snell."0 Upon the ingestion of food after fasting, the body temperature tends to rise. This rise is evident in all the experiments with food here reported, but is particularly noticeable in experiment No. 70, where the rise is continuous for the three days of the experiment. It is obvious, however, that the average temperature for the third day (38.22° C.) indicates a febrile condition and hence the temperature observation for this day can not be considered normal. In experiment No. 76, the average tempera- tures are likewise slightly above those of experiment No. 75, although it is evident that the defective method of averaging in both series of experiments might readily account for this difference. The observations given in table 190 indicate that during short fasts no noticeable disturbances between thermogenesis and thermolysis can be observed. They are furthermore of specific value in furnishing data for computing the total heat production according to the formula discussed on page 49. PULSE RATE. As an index of the variations in the degree of muscular activity the pulse rate is of great value. Even in experiments where the external muscular activity is reduced to a minimum the pulse rate may be of greatest service in furnishing a clue to the degree of internal muscular activity, muscular tonus, etc. The heat production, carbon dioxide elimination, and oxygen consumption vary within wide limits from experiment to experiment, and indeed from day to day of the same experiment. It therefore becomes very important to secure all possible data regarding the intensity of metabolism for the fullest under- standing of the phenomena of fasting. It is for this reason probably that in some of the earlier fasting experiments on record we find more or less complete observations regarding the pulse rate. The oldest observation on the pulse rate during fasting is that of Nicholson (1), who, without giving any exact data, states that the heart sounds of the fasting prisoner under his care became rather faint and the pulse soft and slow. Luciani (4) took Succi's pulse twice each day. The subject was resting quietly in bed, save on one occasion when he had returned from an evening visit with friends. The pulse remained, in practically all cases, between the limits of 54 and 70 beats per minute. On the occasion referred to above the pulse had risen to 85. No decrease was noted as the fast progressed. Senator & Mueller (7) made observations on Cetti and Breithaupt twice daily. Cetti's pulse when resting varied from 68 to 92, while Breithaupt's "Loc. cit. : Loc. cit. Pulse Bate. 317 varied from 47 to 66. Other observations made on these two f asters in con- nection with the research showed an unusual irritability of the heart. On one occasion Cetti's pulse rose from 80 to 104 when he rose from his couch and sat up. Breithaupt showed likewise an unusual irritability of the heart while doing small amounts of muscular work. These observations are in accord with the one observation made by Luciani (4), in which the increase in pulse rate after a short run was much greater with Succi than with several well-fed assistants with the same exercise. No increase of Cetti's pulse was observed with resumed feeding, but a marked increase occurred in the case of Breithaupt on the first day after the fast. The subject "J. A." (9) in the Stockholm laboratory took his pulse rate every two hours while he was awake. The lowest recorded rate was 62 and the highest 92. The average rates for the five days of the fast were 70.1, 70.3, 75.6, 71.1, and 70.4. On the 2 days with food following the fast the average pulse rate was 80.3 and 78.8, noticeably higher than during the fast. Hoover & Sollman (8) have reported the most elaborate series of obser- vations on the pulse rate during fasting thus far recorded. The pulse was taken every 2 hours during an 8-day fast in hypnotic sleep. The pulse was highest at the time when the subject was awakened from his sleep, when a rate of 79 was observed. The nearest approach to this maximum was found on an occasion when the subject awoke, arose from the bed, and went to a water tank and drank several cups of water. At that time the pulse rose to 72. On no other occasion did the pulse rate rise above 68, and a minimum of 36 was reached. The average pulse for the 8 days was 62, 56, 48, 46, 47, 44, 43, and 46. The authors, however, point out that the pulse rate of their subject can hardly be considered as indicative of the normal since a low rate was maintained by command of the hypnotist. Paton & Stockman (3) state that the pulse of their subject Jacques averaged from 50 to 60 beats per minute. No data regarding the variations in the average rate as the fast progressed are given. In the first experiments made in this laboratory, the pulse rates were taken by the subjects themselves. The number of beats was counted for 2 minutes, and the average accepted as the true count. In connection with the statistics of each experiment in which the pulse was counted, records of pulse rate as taken by the subject are given. In the later experiments, the pulse rates were obtained by means of the Pitz pneumograph, and recorded by an assistant outside the respiration chamber. By this means it was possible to secure data regarding the pulse at any time of the day or night. Marked variations in the pulse rate during different fasting days may be noted in many of the experiments. With the subject B. F. D. in experiment No. 59, the pulse rate rose gradually throughout the 3 days of the fast. In 318 Influence of Inanition on Metabolism. experiment No. 68, no records of the pulse were made. But two records per day were made in experiment No. 69, and these showed general uniformity of the pulse rate from day to day, although the rate was somewhat higher on the last day than on the first. In the food experiment No. 70, immediately follow- ing fasting experiment No. 69, only two observations were made, both on the second day. The two records show an increase of about 25 beats per minute over those of experiment No. 69. The pulse rates were taken more frequently in the series of experiments with S. A. B. than in Nos. 69 and 70. In experi- ment No. 71, the rate steadily diminished during the four days of the fast. During the following day (experiment No. 72) food was ingested and there was a noticeable increase over the rate of the previous day. In experiment No. 73 the pulse rate continually decreased on each of the 5 days of the fast. After the ingestion of food (experiment No. 74) the pulse rate increased materially on the first day, fell off slightly on the second, and on the third day regained the rate of the first. In the 7-day experiment (No. 75) the pulse increased on the second day, but continually diminished on the succeeding 6 days of the experiment. After the ingestion of food (experiment No. 76) the pulse increased materially and Avas highest on the third day. Contrary to previous experience with this subject, in experiment No. 77, the pulse rate steadily increased for the 4 days of the fast, being highest on the last day. In the series of 2-day experiments with different subjects, in all except experiment No. 89 the pulse rate increased on the second day. The rate during this last experiment remained about the same on both days. The pulse rate was usually lower during the night than during the day. Excepting the observations during the day itself, and comparing only those during the night it is seen that the pulse rate steadily increased during the 3 nights21 of the experiments. In a few instances the records for pulse rate are defective during the preliminary nights and hence these can not be compared. But the increase of pulse on the 3 nights is strikingly evident in experiments Nos. 80, 82, 83, and 85, though it remained practically the same on all 3 nights of experiment No. 89. An examination of the records for each experiment shows marked, and frequently, sudden changes in the pulse rate. In the large majority of instances the sudden increases in pulse can be accounted for by some body movement immediately preceding them, such as going to the food aperture, taking the strength tests, or changing from the reclining to the sitting or standing positions. In a number of instances, however, the marked fluctuations are not so readily explainable, and it seems not at all impossible that we have 21 It will be remembered that the subject entered the chamber on the evening before the experiment proper began. Hence there are 3 nights and 2 days in each experiment. Pulse Eate. 319 here some examples of that increasing irritability of the heart observed in Sucei (4) and Cetti (7) during their longer fasts. In all cases sudden changes in muscular activity, even though slight, noticeably affected the rate. It should be noted that there was an increase in the pulse rate in all experi- ments in which food was ingested. The relation between pulse rate and internal muscular work is more fully discussed in a succeeding paragraph/2 where it is pointed out that the pulse rate is a remarkably good index of the degree of internal muscular activity. This is especially true when comparing different days of the same experiment with the same subject. It still remains a fact, however, that some subjects, notably A. H. M. (experiment No. 81), have an unusually low pulse rate, while others, as for example, S. A. B. (experiment No. 77), have a very much higher pulse, although it should be borne in mind that all the subjects were living under conditions assuring approximately the same muscular activity. The study of the pulse rate, especially with the Fitz pneumograph, shows marked variations when the sleeping and waking hours are compared. But as this study of the pulse rate was only secondary to the securing of the heat measurements and of chemical data, the records are not sufficiently complete to permit careful consideration of the fluctuations in pulse rate from hour to hour and day to day. In subsequent experiments it is hoped that observations may be taken at sufficiently frequent intervals to show the relationship between the pulse rate, respiration rate, and body temperature, and thus permit of a strict comparison of the three factors. An examination of the changes in the pulse rate as the experiment progresses shows that in the longer experiments, especially those with S. A. B., the pulse rate almost invariably decreased as the fast progressed. In the majority of the short experiments on the contrary, the pulse rate was greatest on the second day of the experiment. From the observations during the 3 nights the deduction may be made that there was a marked tendency for the rate to increase from day to day. The pulse rates, then, are of specific value in this discussion only as indicating the varying degrees of muscular activity on the different days of the same experiment. There was no opportunity to study the character of the pulse in the experi- ments made in this laboratory other than the occasional observations of the attending physician. 22 See p. 487. 320 Influence of Inanition on Metabolism. RATE OF RESPIRATION. The well-known effect of muscular work in increasing the rate of respiration makes records of this factor of importance in fasting experiments as a further index to the degree of internal muscular work. In several of the earlier fasts the respiration rate was observed. Luciani (4) found with Succi a respiration rate of from 15 to 25 per minute. The average rate for the last week of the fast was 18 per minute. In connection with the numerous experiments on Cetti and Breithaupt to determine the respiratory exchange, Lehmann & Zuntz (7) had occasion to record the rate of respiration. It should be said, however, that the subject was in all cases breathing through the mouthpiece of the Zuntz-Geppert respiration apparatus, and hence the rate may not have been normal. The average rate per minute for Cetti observed duriug the respiration experiments on the ten fasting days was 13.1, 14.1, 13.3, 14.6, 15.4, 16.2, 17.3, 17.6, 16.2, 15.7. On the second food day the average rate was 13.05 per minute. Fifteen days after the end of the fast the average rate was 12.1 per minute. Observations were made with Breithaupt during both work and rest. The data were obtained as with Cetti during the progress of a respiration experi- ment with the Zuntz-Geppert respiration apparatus. The rates of respiration when at rest for the 6 days of the fast were: 19.2, 19.0, 20.4, 22.8, 26.3, 22.2 per minute, respectively. The average for the 2 days with food before the fast was 17.9 per minute, for the fasting period 21.5 per minute, and for the 2 days with food following the fast 23.6 per minute. Similar fluctuations were observed in the respiration rate during muscular work. Paton and Stockman (3) state that during his 30-day fast Jacques's res- piration rate varied from 23 to 30 per minute. In all the experiments cited above, the respiration rate was recorded only during the waking hours. In the 8-day experiment with Sohn, Hoover & Sollman (8) recorded the respiration every hour. The average rate per minute for each day was found to be 18, 26, 20, 19, 19, 18, 17, 20, respectively. The lowest rate was 13, the highest 45 per minute. In general it was lowest during the night, although the subject was in a hypnotic sleep during the whole time of the study. For all the Middletown experiments prior to No. 79, no data are available regarding the respiration rate. Several futile attempts were made to secure respiration rates from time to time, but it was not until the Fitz pneumograph was used that any satisfactory data could be obtained. In experiment No. 79 the average number of respirations per minute during the preliminary night was 12. During the first day, from 7 a. m. to 11 p. m., while the subject was awake the rate was not far from 18. This high rate continued throughout the night and the next forenoon, increasing during the second day and decreasing only very slightly during the last night. On the Eespiration Kate. 321 whole, then, there was a marked tendency for the respiration rate to increase as the experiment continued. The usual rate during the preliminary night of experiment No. 80 was 14 per minute rising somewhat (17) in the fore- noon of the next day. In the afternoon the respiration was irregular (12 to 20) and during the night of October 27-28 it was noticeably higher than during the preceding night. On the second day of the experiment the rate was slightly higher than on the first day, while on the last night the rate was practically the same as during the preceding night. The respiration rate therefore showed a distinct tendency to increase as the fast progressed. In experiment No. 81 the respiration rate during the preliminary night was about 13. On awaking it increased to 17, but did not materially quicken during the day. While the subject was asleep during the day the rate was even less than that of the night. On the second night in the chamber (first night of the experiment proper) the rate was about 14 per minute. It was perceptibly higher on the whole throughout the second day and higher also during the last night. Here also there was a tendency for the rate to increase as the experiment progressed. The respiration rate on the first preliminary night of experiment No. 83 was 17 per minute. During the day it increased to an average of about 20, falling off in the evening and remaining at about 16 to 19 during the night. On the second day of the experiment proper the average respiration rate was about 20, falling off somewhat during the evening, although rising again during the last 2 hours of the experiment to 20. There was therefore a slight tendency for the respiration rate to increase on the second day of the experiment, and there was a noticeable rise in the morning just before awaking. In experiment No. 85 the respiration rate during the last part of the first night averaged 12. It remained about the same during the next day, but increased slightly in the latter part of the afternoon. During the entire remainder of the experiment the respiration rate remained singularly constant and the differences usually observed between night and day almost disappeared. In experiment No. 89 the rate during the preliminary night was 16. During the following day it rose to 19, falling off in the evening to 16. During the second day the rate was unusually low for this subject during the forenoon, but rose to 21 at 2 o'clock, and during the 2 hours, from 6h 30m p. m. to 8h 30m p. m., remained at about 21 or 22. During the last night the rate remained practically uniform at 16 per minute. While recorded with considerable frequency and hence of great value as supplementing the data furnished by the pulse rate for estimating the amount of internal muscular work, nevertheless the respiration rate was not studied with sufficient accuracy to enable special conclusions to be drawn from it. 21 ;;j-.) Influence of Inanition on Metabolism. It is further of interest to compare the ratios between the respiration rate and pulse rate in the series of 2-day fasting experiments. While with normal individuals the pulse rate is usually 4.5 times that of the respiration, here very wide variations in this ratio occur from day to day and indeed from hour to hour. Inspection of the data confirms the belief that at least during the first 2 days of fast the pulse rate is much more liable to fluctuations than the respiration rate. BLOOD. In many of the earlier fasting experiments, both with animals and with men, examinations of the blood, chemical and microscopical, were included in the routine of the investigation. Unfortunately almost all of the chemical work, when viewed from the standpoint of the present knowledge of the chemistry of the blood, is extremely unsatisfactory, and it is greatly to be regretted that the microscopical examination of the blood was not carried out in all cases by methods that are free from criticism. Although micro- scopic examination is a phase of investigation distinctly outside the sphere of the ordinary chemical laboratory, it was possible in some of the experiments here reported to make such examinations of the blood, although with by no means the completeness that could be desired. Such information as was obtained is accordingly here presented and the question of the influence of inanition upon the composition of the blood discussed. Luciani, in 1890, made an examination of the blood of Succi every third day during the 30-day fast at Florence. The erythrocytes and leucocytes were counted and the relative amount of hemoglobin determined. The results are expressed in the form of a curve.23 The erythrocytes showed fluctuations during the fast, with a tendency for the number per cubic millimeter to increase as the fast progressed. Although convinced of the relative increase in the number of erythrocytes, Luciani contends that there is no evidence that the absolute number in the blood of the body is increased. These fluctuations are explained by Luciani as the result of changes in the water content of the blood. The results of the hemoglobin tests showed a loss of hemoglobin unaccom- panied by a loss of erythrocytes. The leucocytes were very considerably dimin- ished (from 14,536 on the first day of the fast to 861 on the seventh fasting day). Both Luciani and Daiber * discuss in considerable detail the relations of the blood to metabolism. Senator & Mueller (7), in reporting the results of their examinations of the blood of Cetti and Breithaupt, note an increase in the red blood corpuscles with both subjects. In the case of Cetti it was very considerable, amounting to more than 1,000,000 per cubic millimeter. The blood of Breithaupt, on the contrary, showed only a slight increase. 23 Das Hungern, Leipzig (1890), Tafel i. * Loc. cit. Blood Examination. 323 Observations regarding the number of the leucocytes were taken on only 3 fasting days, i. e., the fourth and ninth days of the fast with Cetti and on the last day with Breithaupt. In all instances the number was considerably less than the normal. This agrees with the observations of Luciani. The observations regarding the hemoglobin in Cetti's blood showed a continued diminution, while Breithaupt's showed a slight increase. At the end of the sixth fasting day it was one-fifth greater than on the last food day, in spite of the fact that the red blood corpuscles had not increased. Senator" also found a slight increase in the hemoglobin content of the blood in a case of inanition with a woman 54 years of age. In a later examination of Succi's blood by Tauszk M the conclusions reached were : ( 1 ) that after a short period of diminution in the number of red blood corpuscles there is a slight increase; (2) the number of white blood corpuscles decreases as the fast progresses; (3) the number of the mononuclear corpuscles decreases; (4) the number of the eosinophiles and polynuclear cells increases and finally that the alkalescence of the blood diminishes. The samples of the blood examined in connection with the fasting experi- ments reported here were taken according to the method described on page 12. It frequently happened that it was extremely difficult to get a sufficient supply of blood from the finger to enable the counts to be made of the red and white corpuscles on each day of the fast, but fortunately blood smears were always obtained which afterwards were subjected to the differential count. The regular Thoma-Zeiss hemocytometer was used in counting the erythro- cytes and leucocytes. The determinations of hemoglobin were in some instances made by the Tahlquist method. In other cases a von Fleischl hemometer was used. The same difficulties and criticisms that pertain to blood examinations in previous fasting experiments undoubtedly to a certain extent obtain here with regard to the taking of the blood sample, the enumeration of the blood corpuscles, and the hemoglobin tests. As is seen from a record of drinking water, there were marked differences in the amount of water taken on the different days of the experiments, although on the majority of days the samples of blood were taken at about the same hour. In experiment 3SJ~o. 75 the samples for blood were taken almost invariably at 9 a. m. On the 2 days when two samples were taken, the second was taken at 3 p. m. In experiment No. 77 the samples were taken as a rule from 3 p. m. to 4 p. m. Although the irregularity in the amounts of drinking-water consumed and 24 Charite-Annalen (1887), 12, p. 327. ^Orvosi hetilap. Budapest (1894), p. 512; abstract in Jahresbericht f. Tier- chemie von Maly (1894), 24, p. 147. 324 Influence of Inanition on Metabolism. in the hours of taking the samples is to be regretted, it is important to note that in the absence of digestion and muscular activity, those physiological factors that tend most to cause variation are reduced to a minimum. The differential counts are believed to be as accurate as it is possible to make, and we are now aware of no criticism that can properly be applied to these measurements. They apparently represent the true condition of the blood of these subjects. The results of all the blood examinations are expressed in the form of curves given on the two figures herewith. MAR. APRIL WAY 3I\4 o.750.00o\ 5 6 7 8 9 10 ill I'2 13 14 16 20 3 7 1 8 9 10 ii |l2 20 25 26 C 1 !H &500.0OQ 1 1 ! l\ :/ ER fT HR( }C1 'TE .s Ij M00.0001 r ^ / f* 1 i 5,7SC.OO0| / / / 1 ! 5300,000! I f\ 1 ! 5,260,0001 1 ! Moaooo, s 1 1 ! 4,750.000 \ :\ 1 ! 4,500,000! M \ / 1 !«- 4.250.0C 0| NA NIT 10 M F >ER IOC M u NANI1 PERK ior DD" H \ / 4000.0001 V f PERCE 100 NTl 95 H AEI /IOC 5LC BIN 90 85 80 \y / V \ 75 i i Fig. 1. The blood examinations covered experiments Nos. 75 and 77, with a few observations on the intermediate period and the period after the last fasting day. The full interpretation of these curves is given essentially as in the formal report submitted by Dr. A. E. Diefendorf, Pathologist of the Connecti- cut Hospital for the Insane, Middletown, Conn. The red blood corpuscles show only a slight diminution in number during each fasting period with a relatively rapid rise in the days immediately follow- ing the end of the fast. The same is true of the hemoglobin curve, which seems to have been more profoundly affected by the first and longer fast. The rise in the hemoglobin immediately after the end of the fast corresponds with the rise in the number of erythrocytes. MAR. A °f?IL i /IAY ?nm\ \4 5 6 7 8 9 10 i " 12 13 14 15 20 3 7 |8 g 10 M |l2 20 25 26 5 15.000 LE :uc OC YT ES IQ000 80 -5( ool PERC 75 :nt -\ PC )LY /f RP HOI nIUC lIe AR LE .uc dc YT ES 70 ^ ^ \ A '\ 65 ■ !— ' It r v / | v_ ^ A \ / V fin \ i 55 k J \ 50 \ 45 42.5 1^ NA NIT IO nI f >ER IOC -»! U MAT "PE vl 1 1 RIC Lj j / 40.0 / 37.5 / 35.0 / 32.5 30.0 27.5 it |C/D ;S MA LL u 'MF >HC CY TE s! 25.0 '5 22.5 / \ 20.0 IO 1+ / \ 17.5 ' / \ \ J f \ / 15.0 ! / \ J V / > 12.5 1/ 17+10 V 15 LA RG L I .YN 4R1- OC YT ES c A 13 V v.! A V II J s. J \ 9 A 7 l-J ?t r\ 1 /-■ \ 5 ii V \ 2.4KJ 1 2.0 EO sir JOF 'NI _E5 1.6 \ ,.2L ovy NOR mAl A /"A A" 0.8 \/ \ i y ' \ w \ i I 0.4 V V V \ \ \ I.0S1 ).o| V \ 0.9 \ / 0.8 MA ST CE LL! 0.7 0.6 \ 0.5 \ / A h V ! | H 0.4 IGH \J0R MAI \l; MIT \ \l \ / 0.3 :\ 0.2 \ 0.1 Fig. 2. 326 Influence of Inanition on Metabolism. As regards the leucocytes, it was apparent at the outset that the subject normally presented a condition usually regarded as a leucocytosis. During the long fast there was a progressive diminution until the morning of the end of the fast when there was a comparatively slight rise. The second fasting period in which the observations were made produced very little effect upon the number of the leucocytes, there being a gradual rise, amounting, however, to only 2000 leucocytes. Only in the two weeks following the end of the second fast did the number of leucocytes approach the normal. In the differential count of the white blood corpuscles it was found that the polymorphonuclear leucocytes averaged high during the first fast, except during the second and seventh days when their number fell to low normal range, 63 and 64 per cent. During the second fast the percentage again rose above the normal level, but in neither instance did the percentage reach a distinctly pathological level. It fell below normal twice in the interval between the fasts and three times during the two weeks following the last fast. The small lymphocytes were below normal range with the exception of two instances, namely, upon the second and seventh days during the long fast. The same average low percentage of small lymphocytes prevailed in the second fast with the exception of the last day of the fast. As regards the large lymphocytes, it was found that the percentage rose above the normal during the last two days of the first fast, and remained wholly above the normal range during the second fast. The eosinophiles averaged low throughout the first experiment and below the normal range except on the second and fourth days. The same applies to the second fast during which the percentage rose above the low normal range upon only one day, the third. The mast cells during the first test averaged high normal and above normal, and the same is true of the second and shorter fast In conclusion, the most striking results of blcod examinations in the continued fasts seem to be : (a) The progressive average fall in the number of erythrocytes with the recuperation following. (b) A corresponding diminution in the percentage of hemoglobin. (c) A relative progressive fall in the percentage of leucocytes in the prolonged fast, but no remarkable effect of fasting to be seen in the relative percentages of the various types of leucocytes. (d) A high percentage of polymorphonuclear leucocytes during the fasts, explained by the relative leucocytosis, which in turn explains the relatively low percentage of the small lymphocytes and the high percentage of large lymphocytes. Strength Tests. 327 STRENGTH TESTS. The popular belief that strength diminishes rapidly as a fast progresses has been based upon the innumerable personal impressions of individuals who have from time to time gone without meals and observed an apparent lack of strength. These personal impressions have been scientifically substantiated by tests with the ergograph as made by Maggiora * in which a rapid diminution in strength immediately follows the beginning of a fast. The tests made by Luciani (4) on Succi in which a dynamometer was used to measure the strength of the right and left hands showed results seemingly at variance with the popular impression. Thus, on the 21st day of the fast, Succi was able to register on the dynamometer a stronger grip than when the fast began. From the 20th to the 30th day of the fast, however, his strength decreased, being less at the end than at the beginning of the fast. In discussing these results, Luciani pointed out the fact that Succi believed that he gained in strength as the fast progressed and hence probably did not exert the greatest power at the beginning of the experiment. Considering the question of the influence of inanition on the onset of fatigue, Luciani states that the fatigue curve obtained from Succi on the 29th fast day was similar to that obtained with an individual under normal conditions. The interesting observations made by the subject of Johansson's experiments, J. A. (9), in which he noted the length of time he could suspend himself on his arm, showed a marked falling off in strength as the fast progressed. This strength was rapidly regained on the subsequent ingestion of food. As has frequently been pointed out, tests with the dynamometer and tests such as were employed by J. A. are, however, not necessarily true indices of the actual strength but a resultant of will and strength.27 Therefore strength tests with the hand dynamometer are of questionable value as accurate records of the strength of muscles at different periods of the fast. In the experiments on Cetti (7) the dynamometer was not used, but the measurement of the vital capacity was taken as an index of the strength. This measurement tested the tension of the respiratory muscles. From the 5th to the 10th day there is no noticeable diminution in the vital capacity, although on taking food after the fast there was an increase of several hundred cubic centimeters above that of the fasting period. The results with Breithaupt indicate an actual increase in the capacity of the muscles to perform work as the fast progresses. In the series of 2-day experiments here reported, tests were made with a Tiemann hand dynamometer to secure data regarding any apparent influence of inanition on strength. In the following table the results are recorded in ie Arch. f. Anatomie und Physiologie, 1890, p. 227. 27 For a more complete discussion of this point see Luciani (4), p. 56, and Leh- mann & Zuntz (7), p. 184. 328 Influence of Inanition on Metabolism. pounds, 1 pound being equivalent to 453.6 grams. An attempt was made to obtain data regarding the rapidity of the onset of fatigue by having the subject test the strength of the hand muscles with the dynamometer in a series of 5 to 7 tests, first with the right hand, and then with the left. In all the experiments except No. 79 and No. 80 the tests were taken in succession, finishing the set on one hand before beginning the tests on the other. At certain times only 3 and 4 consecutive observations were taken, while at other times 6 and even 7 tests were made, and hence the averages are not strictly comparable. An examination of the data in table 191 shows that as a rule the strength decreases rapidly at the beginning of the fast and is almost immediately, in part at least, regained with the ingestion of food. In experiment No. 89 there was a marked increase in strength on the first day of fasting, followed by a slight loss on the second day. The effect of the subsequent ingestion of food in increasing the strength is not as noticeable in this experiment as in others. This is doubtless due to the small loss observed in the fasting period. The records were not made at sufficiently frequent intervals to draw any particular conclusions regarding the fluctuations in the strength for different periods of the day. In general, the observations accord with the conclusions of Mosso and Maggiora with the ergograph, namely, that as the fast progresses there is a noticeable falling off in the strength as measured by the hand dyna- mometer. After renewed ingestion of food, the strength is rapidly regained. The apparent onset of fatigue seems in general to be somewhat more rapid as the fast progresses, but the data are not sufficient to warrant any positive state- ments on this point. Strength Tests. 329 Table 191. — Strength tests with hand dynamometer, preliminary to, during, and following metabolism experiments. METABOLISM EXPERIMENT No. 79. Date. Hand. Record of individual tests. Av«r- Total. a&e. Oct. 13, 9 a. m., fasting. Oct. 14, 9 a. m., fasting. Oct. 14, 8 p. m., fasting. Oct. 15, 7h 15'" a.m., fasting. Oct. 16, 3 p. m., food. Pounds. Right. 100 92 93 92 93 Left Right Left . Right Left . Right Left . Right Left . 95 85 81 80 80 88 75 85 70 84 70 82 70 80 70 88 80 85 70 78 65 80 60 78 65 95 84 70 93 76 80 70 91 80 85 95 87 90 75 Lbs. 94 84 84 71 82 68 86 75 92 80 Lbs. 178 155 150 161 172 METABOLISM EXPERIMENT No. 80. Oct. 26, 8h 45ma. m., food 7 40 p.m., food Oct. 27, 9h 00m a. m., fasting . 4 07 p.m., fasting. 11 00 p.m., fasting. Oct. 28, 9h 00ma. m., fasting. 5 00 p.m., fasting. 11 00 p.m., fasting. Oct. 29, 71' 00m a.m., fasting. 9 15 a. m., food . 11 30 a.m., food, 2 10 p.m., food. Right Left . , Right . Left . . Right . Left . . Right. Left . . Right . Left . . Right. Left . . Right . Left . . Right . Left . . Right . Left . . Right . Left . . Right . Left . . Right . Left . 105 106 108 103 85 100 106 86 84 94 115 111 108 96 107 95 94 85 84 94 94 90 94 95 90 84 90 84 106 105 99 95 95 89 95 90 94 90 93 88 82 81 90 90 99 99 86 81 88 81 88 95 92 88 84 80 74 84 92 90 95 95 88 86 87 78 102 97 101 98 93 85 86 80 105 107 110 109 108 95 93 95 95 89 107 117 110 110 109 95 96 95 92 105 125 128 121 110 116 105 108 120 103 100 101 94 107 90 93 87 101 92 92 84 95 84 91 81 93 85 100 86 108 93 111 97 120 107 195 197 180 193 176 179 172 178 186 201 208 227 330 Influence of Inanition on Metabolism. Table 191. — Strength tests with hand dynamometer, preliminary to, during, and following metabolism experiments — Continued. METABOLISM EXPERIMENT No. 80-fConf d.) Date. Hand. Record of individual tests. Aver- age. Total. Oct. 29, 7h 10m p. m., food . Oct. 30, 8h 45m a.m., food. Pounds. Right. 109 98 95 95 90 Left . . 90 80 90 90 80 Right. 104 101 119 106 101 Left..! 89 88 100 89 95 Lbs. 97 86 106 92 Lbs. 183 198 METABOLISM EXPERIMENT No. 81. Nov. 20, 8h30ma. m., food.. 12 10 p.m., food.. 3 40 p.m., food.. 7 40 p.m., food.. Nov. 21, 9" 00ma. m., fasting 3 00 p. m., fasting 9 00 p. m., fasting Nov. 22, 9U 00m a.m., fasting 7 00 p. m., fasting Nov. 23, 7h 00m a. m., fasting 9 45 a. m., food. . 1 45 p. m., food . . Nov. 24, 10h 40™ a. m., food . . Right Left . Right Left . . Right . Left . . Right , Left . , Right . Left . . Right Left . , Right. Left . . Right . Left . . Right. Left . . Right . Left . . Right Left . Right Left. Right Left. 82 83 81 79 86 83 81 82 82 82 78 69 84 76 71 81 85 84 82 82 83 81 83 82 71 78 75 78 75 76 80 78 83 81 78 83 80 79 82 81 84 86 75 86 86 84 75 81 87 86 87 83 79 81 84 86 75 67 78 75 79 80 78 71 73 75 73 78 80 68 80 79 81 80 79 73 78 70 86 79 77 81 75 72 75 75 84 80 80 78 80 80 77 83 85 84 82 82 80 78 87 86 82 85 84 82 81 83 83 86 79 84 79 80 71 73 71 78 74 69 81 80 75 65 75 75 80 84 74 63 81 85 78 76 84 74 79 85 78 78 82 82 82 82 82 78 86 88 86 83 80 73 87 84 81 85 80 .. 83 .. 86 84 80 81 80 81 79 78 70 72 71 71 82 82 78 81 80 80 75 83 74 80 90 87 82 82 81 80 80 75 75 75 80 78 79 80 73 81 77 77 80 80 79 81 81 84 82 78 87 84 164 161 155 150 158 159 154 154 160 160 165 160 171 Strength Tests. 331 Table 191. — Strength tests tcith hand dynamometer, preliminary to, during, and following metabolism experiments — Continued. METABOLISM EXPERIMENT No. 82. Date. Hand. Record of individual tests. Aver- age. Total. Nov. 23, 8h 15ma. m., food. Nov. 24, 9h 00ma. m., fasting. 3 00 p.m., fasting. 10 00 p.m., fasting. Nov. 25, llh 00ma. m., fasting. 3 40 p.m., fasting. 9 30 p. m., fasting. Nov. 26, 7h00ma. m., fasting. 9 30 a. m., food 12 30 p.m., food 3 10 p. m., food 6 40 p. m., food Nov. 27, 7h 10m a. m., food 8 16 a. m., food 4 50 p. m., food Dec. 4, 3h 00mp.m.,food Pounds. Right. 100 90 90 84 73 82 74 70 Left . . 88 88 81 88 75 73 78 68 Right. 100 84 74 80 80 70 72 Left . . 99 80 99 80 76 82 80 Right. 95 79 65 79 79 82 75 Left . . 82 88 80 90 85 84 80 Right. 84 75 85 94 71 74 85 Left.. 100 88 70 70 82 71 70 Right. 88 82 66 62 75 68 65 79 Left . . 75 70 61 54 53 65 56 Right. 86 69 78 74 68 70 71 Left.. 75 71 69 60 67 64 72 Right. 70 70 62 63 52 55 55 Left.. 69 66 55 60 54 55 55 Right. 74 65 65 58 52 65 66 Left . . 87 82 78 65 60 70 69 Right. 84 76 70 68 66 56 73 Left.. 85 70 72 64 63 56 55 Right. 90 80 85 76 85 84 Left . . 91 80 71 70 73 80 Right. 90 80 79 71 82 80 Left.. 89 77 75 73 70 74 Right. 89 74 62 65 80 81 Left.. 80 70 71 80 83 86 Right. 80 65 57 74 80 77 Left . . 62 61 60 75 70 70 Right. 88 81 80 88 84 86 76 78 Left.. 90 83 73 71 76 76 76 64 Right. 86 71 81 85 85 70 80 89 Left.. 91 81 79 78 76 74 70 Right. 93 80 74 74 75 79 Left . . 89 71 82 70 74 80 Lbs. 83 80 80 85 79 84 81 79 73 62 74 68 61 59 64 73 70 66 83 78 80 76 75 78 72 66 83 76 81 78 79 77 Lbs. 163 165 163 160 135 142 120 137 136 161 156 153 138 159 159 156 332 Influence of Inanition on Metabolism. Table 191. — Strength tests with hand dynamometer, preliminary to, during, and following metabolism experiments — Continued. METABOLISM EXPERIMENT No. 83. Date. Hand. Record of individual tests. Aver- age. Total. Dec. 4, 10h 05ma. m.,food. 3 15 p.m., food. Dec. 5, llh 00ma. m., fasting. . . 7 00 p. m., fasting. . Dec. 6, llh00ma. m., fasting. . 7 00 p.m., fasting. Dec. 7, 9h 00m a. m., food 7 30 p.m., food. Dec. 8, 7h 30m a. m., food . . 10 00 a.m., food. Right Left. Right Left . Right Left . Right Left . Right Left . Right Left . Right Left . Right Left . Right Left . Right Left . Pounds. 95 92 76 80 74 63 53 56 51 52 89 81 76 81 83 72 55 58 52 56 85 73 82 80 83 55 59 55 50 58 78 78 66 70 72 56 59 52 51 46 82 81 73 72 75 53 44 53 51 50 77 71 73 71 68 55 62 83 72 61 54 87 71 58 56 94 86 65 63 85 81 67 63 64 65 55 53 53 47 61 64 66 50 53 53 73 80 63 52 53 54 81 61 81 58 57 53 68 71 65 53 54 53 Lbs. 80 54 58 79 51 61 47 56 48 68 51 60 51 68 51 73 49 74 52 72 57 56 71 59 78 43 76 43 58 49 56 41 71 53 78 51 170 158 160 154 55 80 57 80 58 72 56 77 53 71 48 71 50 70 40 72 49 61 48 63 51 65 49 69 53 61 52 73 53 78 54 78 58 65 53 71 57 Lbs. 135 138 130 121 121 114 122 126 136 128 METABOLISM EXPERIMENT No. 85. Dec. 8, 9" 10m a.m., food. 3 20 p.m., food. Dec. 9, 9h 10ma. m., fasting.. 3 40 p. m., fasting . . . 9 05 p.m., fasting. . . Right Left. Right Left . Right Left . Right Left . Right Left . 64 54 59 54 70 63 61 56 63 53 82 76 70 68 70 73 68 61 63 63 56 59 62 63 44 53 48 47 40 34 52 51 42 49 51 48 36 42 43 42 53 49 45 44 46 44 48 44 48 52 69 56 68 53 59 56 43 53 47 41 36 39 38 44 34 48 43 36 62 58 49 41 44 45 59 64 68 58 54 49 67 60 45 50 70 64 48 46 54 46 120 134 100 90 89 'Additional records: 7.30 a. m., right, 78; left, 63. 10.00 a. m., right, 73; left, 59. Strength Tests. 33J Table 191. — Strength tests with hand dynamometer, preliminary to, during, and following metabolism experiments — Continued. METABOLISM EXPERIMENT No. 85-(Cont'd). Date. Hand. Record of individual tests. Aver- age. Total. Dec. 10. llh 30ma. m., fasting... 4 05 p.m., fasting. 9 20 p.m., fasting. Dec. 11, 11" 00m a.m., food 10 00 p.m., food Dec. 12, 3h 15m p.m., food Dec. 13, 3h 00mp. m., food Dec. 14, 12h 10m p.m., food Dec. 15, 4h 00m p. m., food Right Left. Right Left. Right Left . Right Left. Right Left. Right Left. Right Left . Right Left. Right Left. Pounds. 56 46 45 42 51 41 36 44 42 42 36 38 50 40 49 45 63 55 53 56 41 49 51 44 43 46 41 32 47 42 46 32 58 48 46 58 46 49 52 39 41 41 54 39 49 41 43 44 58 58 53 53 61 51 51 43 38 51 58 51 51 56 49 58 49 47 47 41 74 64 66 60 52 59 62 59 57 50 69 59 72 66 63 56 63 57 59 53 72 62 62 65 59 72 63 56 59 53 48 51 51 46 48 40 61 58 47 39 54 48 53 67 62 53 50 55 56 64 52 60 53 56 55 60 63 51 55 62 78 65 72 54 68 62 68 58 60 70 75 55 58 52 62 55 Lbs. 45 43 52 41 50 44 54 46 54 48 62 56 63 57 62 59 66 61 Lbs. 88 93 94 100 102 118 120 121 127 METABOLISM EXPERIMENT No. 89. Jan. 9, 8h 30m a.m., food. 12 noon, food. Jan. 10, l1' 40mp.m., fasting. 8 00 p.m., fasting. 10 00 p.m., fasting. Jan. 11, lh 42mp. m., fasting. Right . Left . . Right. Left . . Right . Left . . Right. Left . . Right . Left . , Right Left . 126 92 81 90 88 82 84 80 81 78 88 75 104 98 95 82 98 88 84 84 80 86 80 82 95 88 89 83 81 84 93 100 88 88 82 81 100 105 110 105 95 100 95 73 82 91 82 71 106 115 108 101 97 98 91 74 86 84 81 78 98 85 85 90 85 86 89 87 74 74 76 72 85 92 92 75 73 88 79 82 92 74 89 79 82 81 85 83 76 86 86 93 99 72 72 91 80 84 100 80 74 75 81 81 86 63 76 76 171 174 171 179 181 162 334 Influence of Inanition on Metabolism. Table 191. — Strength tests with hand dynamometer, preliminary to, during, and following metabolism experiments — Continued. METABOLISM EXPERIMENT No. 89-(Cont'd). Date. Hand. Record of individual tests. Aver- age. Total. Jan. 11, 8h OCT p.m., fasting. 10 00 p. m., fasting Jan. 12, 3h 00m p.m., food. . . 8 16 p. m., food. . . 10 20 p.m., food. .. Jan. 13, 2h 10mp. m.,food.. . 8 15 p. m., food. . . 10 00 p.m., food. .. Right Left Right Left Right Left Right Left Right Left Right Left Right Left Right Left Pounds. 95 95 92 86 86 72 72 79 94 91 88 71 81 81 79 88 91 91 95 95 84 91 78 79 90 91 72 78 75 73 78 75 95 91 92 S5 86 81 79 81 93 88 88 84 81 72 74 79 95 95 95 88 88 89 89 90 95 99 85 80 79 82 75 82 105 110 104 93 93 86 85 89 90 90 83 84 83 84 74 76 94 95 88 83 79 80 86 84 97 83 81 79 78 78 78 71 85 86 91 92 89 81 81 84 88 82 81 74 80 80 74 80 99 99 91 92 90 91 89 85 88 92 85 75 79 74 66 95 Lbs. 85 84 88 79 86 82 91 85 96 83 86 81 86 80 92 82 Lbs. 169 167 168 176 179 167 166 174 PHYSICIAN'S REPORT. Many of the subjects of the fasting experiments here reported were inex- perienced in experiments of this nature. To insure the use of normal, healthy persons, and to strengthen their confidence, arrangements were made with Dr. J. E. Loveland, a practicing physician, to examine each of the men before beginning the fasts. In certain of the longer experiments he was also requested to visit the laboratory each day, and to report the condition of the subjects as the fasts progressed. The results of his observations during the progress of three of the longer fasts are recorded in the reports given herewith. The following is my report as attending physician to S. A. B., the subject who fasted in the calorimeter from January 7-11, 1905. The observations were made by telephoning to and inspecting the subject through a double glass window, the subject being in a dimly lighted room. Such observations were limited to noting the subject's mental condition as shown by oral and written statements, the facial expression, the actions, movements, tone and strength of voice, color of skin, condition of mouth and tongue, and physical condition, including pulse rate and strength and body temperature as reported by the subject. During the 4 days of Physician's Eepoet. 335 the fast there was no perceptible change to my mind in any particular, save that a thin, white, moist coating of the tongue at the start developed in 4 days into a moderately thick, white, moist coating, and the blood pressure in the radials was lowered enough during the stay in the calorimeter to be perceived by the examining finger. Yours respectfully, John E. Loveland, M. D. The reports for experiments No. 75 and No. 77 follow : During the fasting experiment of March 4-10, 1905, I visited your subject at practically the same hour each afternoon and endeavored to judge of his condition by observing him through the window of the calorimeter, talking to him through the telephone, reading his written reports, and observing his pulse rate and force by the examining finger introduced through the rubber curtain. As far as could be made out by my observations, the history of the subject's condition during this fasting experiment was almost entirely uneventful. On the sixth day of the fast the subject expressed himself as feeling doubtful about his condition. On all other days he was more or less buoyant. On the seventh day of the fast and the first day of food he appeared to show emaciation in the face and the color of the skin appeared slightly dusky. On the same days the subject showed a slight sordes on the gums and the tongue was slightly swollen showing indentations by his teeth. The tongue on the first day of the fast showed a moderately thick white coating, which grew thinner during the fast. The pulse rate varied from 50 to 68 beats per minute. The rate, roughly speaking, tended to grow less as the fast progressed. On the first day with food it rose to 74. On the fourth day of the fast the force of the pulse appeared less than on previous days. On the fifth day there was an irregularity noted — the individual beats varying in force. On the sixth and seventh days of fasting, and the first day with food, the force appeared greater than on the other 5 days. At no time did the pulse rate and force appear to approach a dangerous condition. At the end of the fast the subject was in a condition that, in my opinion, would have warranted his con- tinuing the fast with impunity. During the fasting experiment of April 8 to 11, 1905, the subject's mental condi- tion, as shown by his statements, his facial expression, his carriage and actions, was one of depression and apprehension. His physical condition was apparently normal. On the first day there was a moderately thick, dirty, white coat on the tongue. This coating appeared less thick on the second day and did not change in character from that day. The pupils were moderately dilated on the fourth day. The pulse was always of good quality. John E. Loveland, M. D. SUBJECTIVE IMPRESSIONS AND OBSERVATIONS REGARDING GENERAL CONDITION. It is commonly believed that the withdrawal of food for one or two meals results in dizziness, a feeling of faintness, and, at times, in pains in and about the epigastrium. With fasting men the experiences are varied. The fast of Merlatti,1* which was said to have continued 50 da}Ts, was characterized by 28 Monin et Marechal; Stefano Merlatti, Histoire d'un jeune celebre. Paris (Mar- pon et Flammarion). 336 Influence of Inanition on Metabolism. extreme discomfort, pain, and sensation of coldness. During the 30-day fast of Jacques (3) the only marked discomfort noticed was a slight attack of gout which appeared on the 16th day. In the numerous fasts of Succi (4) no marked discomfort was observed. In fact during his fast at Florence his cheerfulness and apparent good health were the subject of much comment. It should be stated, however, that both Jacques and Succi took small amounts of narcotics from time to time throughout their fasts, though, as Prausnitz (5) has pointed out, this may have been as much to stimulate a popular interest in the concoctions as to dull the senses to any possible pain, except possibly during the early days of the fast. Cetti (7) experienced considerable dis- comfort during the first one and one-half days of his fast, but this suddenly ceased after a movement of the bowels. The condition during Breithaupt's (7) experiment was somewhat complicated by the fact that on the third day of the experiment he contracted a cold in the head which caused him discomfort with a slight temperature increase. It was the opinion of Senator and Mueller (7) that the disturbances observed in both cases were not marked enough to cause any material effect on the metabolism. The records of the subjective impressions of J. A. (9) in the experiments in the Stockholm laboratory show that on the first day of the fast he noticed no dizziness. On the second day, while his general condition was good, he observed unusual weakness follow- ing a slight muscular exertion. On the third day he was in not a little discomfort and was dizzy when climbing on a short ladder inside the respiration chamber. On the fourth day the pain in the stomach disappeared and no dizziness was noticed in the experiment on the ladder. On the fifth day the general condition was excellent, and there was no pain or discomfort in the stomach. His strength, too, was greater, although he noticed that if he arose suddenly from the bed there appeared to be black spots before the eyes. Kosemann,29 who fasted forty hours, records that on the first day there was no very noticeable feeling of hunger. On the second day he was hungry, especially at the regular meal hours. There was furthermore a feeling of weak- ness and a pain in the heart with palpitation. The effect of climbing a ladder was to cause dizziness. In the series of 2-day experiments made by Prausnitz (5) discomfort was experienced in one or two cases, but on the whole the subjects had no pain or other disagreeable results from the fast. There was in general a feeling of weakness on the second day. although all the subjects went about their usual daily occupations. In Prausnitz's opinion the feeling of discomfort attending hunger is, in many instances, a purely psychical condition. That the psychical condition has much to do with the feelings of discomfort during fasting is clear from observation of the notes made by the subjects of "Archiv f. die ges. Physiologie (1897), 65, p. 359. Feces. 337 the Middletown experiments. During a large majority of the fasts the only discomfort noticed was a slight headache or dizziness, and a number of the subjects were cheerful and contented. The attempt to read by means of the insufficient light in the chamber may well account for much of the pain in the eyes and perhaps also for the dizziness. The psychical condition of the subject S. A. B., who was distinctly of a hypochondriacal disposition, determined in a very large measure his subjective symptoms. For example, during the longest fast, 7 days in experiment No. 75, he reported himself in excellent condition and his attending physician substantiated his statement. In fact, at the conclusion of the 3-day food experiment following this fast, the subject remarked to the assistants in the laboratory that he could have fasted for three days more. On the contrary, in experiment No. 77, although the subject, had endured the 7-day fast inside the calorimeter with excellent results, he was extremely apprehensive and nervous. In the series of 2-day experiments a feeling of weakness and occasionally a sense of hunger were the only disagreeable sensations noted. It seems, therefore, that from the experiments made in this laboratory the conclusion can properly be drawn that fasting, per se, produces no marked symptoms of pain or weakness, at least during the first days of inanition. FECES. The excretion of feces has commonly been considered to be the rejection of undigested food material. From more careful examination of the nature and composition of feces, however, it is seen that undigested food may form but a small portion of the total fecal mass. Among the ingredients of normal feces may be mentioned residues of digestive juices and epithelial tissue. These are conveniently termed the " metabolic products." Since under ordinary conditions these are incident to the passage of food through the body, the practice of considering feces undigested food is not without some justification. For while the metabolic products are not a part of the food passing through the alimentary tract, yet they are present in the feces as a result of the ingestion and digestion of food, and hence they may properly be considered as material expended for the digestion of food and should be taken into account in discussing digestibility or the cost of digestion. A number of experimenters have in recent years attempted to study the formation of feces by ligaturing a loop of the intestine, analyzing the contents of the ligature and comparing the composition of this matter with that of normal feces. After the ligature was made and the wound healed the diet was resumed, and under these conditions there was a large amount of material thrown off from the intestinal wall into the ligatured section. This closely resembled fecal matter. It had a chemical composition not unlike that of feces 22 338 Influence of Inanition on Metabolism. resulting from a flesh diet, and many analyses of so-called " fasting feces " show a similar chemical composition. Indeed, Voir, on the basis of the quantity of material thus secured by a ligature of a section of the intestine, has computed the total amount that would be found in the whole intestine. This was found to correspond with the total quantity of feces excreted during normal feeding experiments. It is a well-known fact that fasting animals while passing feces with more or less regularity, reject much smaller amounts of fecal matter than do animals which are fed. If, therefore, the material formed in a ligatured intestine is a true index of the actual amount thrown off under normal conditions during fasting, there must be a marked subsequent absorption of such material as it passes farther along through the intestines. Hence it is not logical to conclude that because large amounts of epithelial debris are found in the ligatured intestine of a dog consuming food, proportional amounts may be formed and pass through and out of the alimentary tract during fasting. The withdrawal of food results in a cessation of the stimuli to peristalsis and thus affects the expulsion of feces, while the total mass of fecal matter becomes diminished as a result of the absence of undigested material, the diminished flow of digestive fluids, and the decreased mass of intestinal debris resulting from the quiescence of the alimentary tract. Fasting, therefore, affects first the amount and regularity of defecation. In considering the influence of fasting on the frequency of defecation, it is important to note that there is usually a normal amount of partially digested food in the alimentary tract at the beginning of a fast. Moreover, in many instances the subjects partake of an unusually large amount of food on the day, if not, indeed, the last meal of the day, immediately preceding the first day of the fast. As a result, this undigested and partially digested food in all probability undergoes the normal digestive processes, and gives rise to the production and flow of digestive juices, leaving according to its nature more or less unabsorbed material in the alimentary tract. Obviously, the fecal matter resulting from food thus ingested prior to the fast can not in any way be considered as fasting feces. The influence of the fecal matter thus formed on the regularity of defecation during fasting naturally varies with the amount and character of the diet before the fast, the length of time intervening between the last meal and the beginning of the fast, and the usual habits of defecation of the subject. In the large majority of the fasts recorded in the literature, no feces were passed during the fasting period. This was the case in Schaefer's so obser- vations on the fasting insane. Nicholson's fasting prisoner (1) did not defecate until the twelfth day of the experiment, though only the first 6 days were wholly without food. MLoc. cit. Feces. 339 During the 30-day fast reported by Paton and Stockman (3) there was no excretion of feces during the whole period of the fast except on the first day when a few pilular masses were passed. In the 10-day fast of Cetti (7) feces were passed only once, i. e., at the end of the seventh fast day when about 175 grams were collected. Breithaupt (7), during a 6-day fast, passed 37.5 grams of fresh feces on the first fasting day and 107 grams at the end of the third fasting day. No more feces were collected till after the period of inanition. Landergren * does not state that feces were passed during his experi- ment, but notes that it was impossible to separate the fasting feces. With the subject J. A. (9) no feces were passed between 7h 45m a. m. on the day before the fast and 7h 40m a. m. on the second day after it concluded. Sohn, the subject of Hoover and Sollman (8), did not defecate during the 8 days of an hypnotic sleep. It should be added, however, that suggestion was used to prevent defecation. Flora Tosca (11) during her 15-day fast defecated but once, i. e., the evening of the sixth day : — after taking a saline purgative. Baumstark and Moler f report that no feces were passed by the fasting woman (Schenk) between the second and fourteenth days of fasting. No weights of feces are given. Owing to long retention in the colon, fasting feces become hard, much dried and pilular, and frequently cause considerable uneasiness. Much difficulty is experienced in passing them, and at times they may cause considerable pain with slight hemorrhages. The use of an enema to remove fecal matter during inanition is qiiite common. This method was employed throughout the 30-day fast of Sued, reported by Luciani. Separation of feces. — In studying the feces resulting from different diets, it is common to separate that portion of the feces belonging to one diet from that belonging to another by means of some material which will either color the feces or will mingle with them in such other manner as to enable the separation to be sharply made. For the coloring material carmine, charcoal, and lampblack have most commonly been used, while the other type of separating agent has more commonly been berry and fruit seeds, silica, and other similar insoluble materials. While with subjects undergoing experi- ments in which a sufficient quantity of food is given, these separating agents serve fairly well to distinguish the feces resulting from the differ- ent foods, the difficulties incident to the separation of fasting feces are much greater. It may reasonably be questioned whether the presence of berry seeds or indeed even coarsely powdered charcoal may not result in a stimulation to * Loc. cit. f Zeit. f. experimentelle Pathologie und Therapie (1906), 3, p. 687. 340 Influence of Inanition on Metabolism. peristalsis, if not, indeed, to a flow of digestive juices which would compare with many normal diets, and hence the fecal products are not necessarily representative of an absolute fast. The use of an insoluble material requires the separation to be made in many instances with the aid of the microscope, and it is probably true that the most reliable information regarding the nature of feces is furnished us by this instrument. In the fasting feces of Cetti, studied by Mueller, numerous tobacco fibers resulting from the large number of cigarettes consumed by this subject, were found throughout the whole mass. Knowing as we do the irritating or stimulating action of undigested cellulose in inducing peristalsis and consequent abrasion of epithelial debris, it is to be questioned whether feces which have been produced under conditions favoring the presence of cellulose material in quantities even so slight as those necessar- ily resulting from the small amount of cigarette tobacco involuntarily swal- lowed are, strictly speaking, fasting feces. In a number of experiments the attempt has been made to separate the feces by means of high enemata. It is a well-known fact that enemata of this nature frequently mix the contents of the bowel in such a manner that an accurate separation is precluded. Furthermore, when the total possible amount of the fasting feces is taken into consideration, it is clear that any method of separation as gross as that involved in the use of enemata is without actual value. Quantitative factors. — In spite of the difficulties of isolating fasting feces, a number of investigators have attempted to separate feces that might be ascribed to the fasting period. It is of interest to observe the quantitative relations of these necessarily imperfectly separated amounts. Obviously the sharpest separation of fasting feces would be expected in the longest experi- ments. Of the prolonged fasts, that of Succi (4) in Florence furnishes the most satisfactory record of the collection of feces. During this fast feces were passed for the first time on the 13th day. An enema was used. Luciani attempted to secure a water content that would represent the normal and hence dried the feces and injection water until the mass was of a pasty consistency. In this form the feces weighed 117 grams. On the 16th day 2 grams of a pasty material were obtained as the result of an injection. On the 21st day a small quantity of glucose was given with the water used in the enema and the subject attempted to retain the water in the colon as long as possible and thus permit the absorption of the glucose. One and a half hours after the enema was taken a greenish yellow fluid with some particles of solid feces was passed. The solid material, weighed in the pasty form, was 19 grams. The last defecation during the 30-day experiment occurred on the 24th day. The feces had little form and weighed in the pasty condition but 12 grams. Thus during the 30 days the total weight of fecal matter estimated as of the Feces. 341 pasty consistency of normal feces was but 150 grams. It is much to be regretted that the actual amount of solid matter was not determined, but allowing the greatest possible error in the estimate of water content, the total quantity of dry matter for the whole experiment was probably somewhat less than that excreted daily by a man with an average diet. The percentage of water in normal feces is not far from 75 to 80 per cent. Consequently it may readily be computed that during the 30-day fast, assuming that the pasty consistency secured by Luciani in the feces would correspond to feces with a water content of 75 per cent, the actual amount of solid matter would be not far from 37.5 grams or a little over 1 gram per day. Since, however, the last feces were passed on the 24th day of the fast, and consequently the fasting feces for the remaining 6 clays were not included, the total amount of dry matter per day for 24 hours would be about 1.5 grams. During the 10-day fasting experiment with Cetti (7) the feces were separated from those of the food eaten previously by means of the softer consistency and more yellowish-brown color. This separation was further confirmed by the microscope. The fasting feces were passed in two portions, the first of 27 grams, collected at the end of the 7th fasting day, and the second after the experiment ended. The dried portion of the feces amounted to 34.147 grams, or 3.4 grams per day, but the writer states that the separation was unsatisfactory. With Breithaupt (7) 28 grams of so-called "fasting feces" were collected at the end of the third fasting day, and after the period of inanition 29 grams more were passed. The total weight of dry matter was 12.10 grams, or 2 grams per day. The separation of the fasting feces from those of the previous food was made by using currants. At the end of the fasting period a charcoal emulsion was used. An attempt to separate fasting feces was made on the subject J. A. (9). The fasting experiment was preceded by a 2-day experiment with food. Before the first meal of the food experiment the subject had fasted 15 hours. With the first meal 30 grams of dried blueberries were eaten. The presence of the residues of the berries in feces indicated that they belonged to the food of the food day.31 In order to separate the fasting feces from the second food period, the subject received in the first meal after the fast 0.5 gram of charcoal, which colored the corresponding feces gray. The total weight of air-dry82 fasting feces as separated in this manner was 12 grams, or about 2.4 grams per day. 81 From the statements of the authors it was assumed that the berry residues would become intimately incorporated with the feces for both days. This assump- tion seems hardly tenable. 32 9.08 per cent of water. 342 Influence of Inanition on Metabolism. Mueller,83 in an experiment with a patient with oesophagus stenosis, states that after four days complete fast there were 17.1 grams of dried feces, or 4.35 grams per day. He also cites two instances of the fasting insane in which the quantities per day were 5.9 and 4.8 grams, respectively, of dried feces. Since, however, in at least two of these cases, the fasting period extended beyond the actual period of the observations, these amounts would be somewhat diminished. For example, in the first instance, the four days of fast during which the patient was under observation were actually the 5th to 8th days of complete inanition. Mueller calls especial attention to the fact that it was very difficult to separate the feces in this period. Furthermore, in the second case of the fasting insane, the six days were from the 4th to the 9th fasting days. No evidence is given as to whether feces were passed on or before the beginning of this period, or whether the subject simply came to the attention of the observer at this time. In reporting the results of Luciani on Succi, Mueller has interpreted the weight of feces reported by Luciani, i. e., 150 grams, as the weight in the dried condition, while Luciani specifically states (4) that they were in all instances weighed in a pasty condition. On this basis the amount of dried material of feces excreted by Succi is reduced from 5 grams per day as computed by Mueller to about 1.5 grams. In the Naples (6) fast, Succi passed feces amounting to 72 grams (23 grams dry matter) on the second day of the fast. By means of purgatives 317 grams of feces (80 grams dry matter) were obtained on the 11th day. Thus there were 103 grams of dry feces collected during the first 11 days. No more feces were passed during the rest of the 21-day fast. The feces passed by Succi in the Vienna fast (10) weighed in the dry form 53 grams. They were collected twice during the 21-day fast. Unfortu- nately practically all the data regarding the feces were lost. The only observation on the defecation of Succi during the Hamburg fast is that of Brugsch (12), who states that during the last 14 days of the fast there was no defecation. It has frequently been considered, especially in the earlier experiments, that all feces passed after the beginning of a fast were fasting feces. In considering specifically the problem of the formation of feces in fasting men, it is import- ant to bear in mind that at the beginning of the fast the alimentary tract is more or less filled with material varying in composition from the partly digested food of the last meal remaining in the stomach or the upper part of the small intestine to the feces in the colon. 88 In criticism of Mueller's observations on fasting feces, it should be said that the quantity per day observed in Cetti's experiment is recorded differently in two places. On p. 17 (Untersuch. an zwei hungernden Mensch. Reference (7)) the amount is given as 3.4147 grams per day, while on p. 106 of the same article the amount is given as 3.818 grams per day. Feces. 343 While unfortunately in most experiments no data are given regarding the character and amounts of the diet and the time and nature of the last defe- cation before the fast began, it must be recognized that a part at least of the feces collected in this manner during fasting must have resulted from the ingestion of food prior to the fast. In the fasting experiments here reported, the data regarding the feces are extremely limited. The natural expulsion of feces is retarded by inanition and hence the defecation is not at all regular. It is of interest, therefore, to consider first the effect of inanition upon the regularity of defecation. Depending upon the amount of food consumed on the day previous, the defecation of the first day of fasting may be quite as regular as on the ordinary food days. If, however, as is the case in many of the experiments here reported, the last meal on the day before the fast was small, there may be entire absence of defecation on the first day. An examination of the record of body move- ments in the different experiments shows that in experiments Nos. 59, 68, and 69 there were no feces passed. In experiment No. 70, 119.2 grams of feces were passed at 7h 40ra p. m., December 22, 1904, i. e., 2| days after the first meal following the fast. This unusual delay in defecation is explained in part at least by the fact that this subject even under ordinary conditions defecates very irregularly. In experiment No. 71, with S. A. B., feces were passed on each of the first three days of fasting. The amounts were 241.5, 41.3, and 48.0 grams, respectively. In experiment No. 72 the feces were passed at 9h 45m a. m., January 11. In the 5 days of experiment No. 73 feces were passed with considerable regularity, i. e., on the first, second, and third days. The amounts passed were 71.6, 87.0, and 51.1 grams, respectively. No more were passed till the second day with food. During the 7-day fast (experiment No. 75) feces were passed only on the first day, i. e., 9h 10m a. m., March 4, 1905. The amount was 48.4 grams. Three days of food followed, but feces did not appear till the last food day, March 13, when 28.1 grams of feces were passed at 7h 40™ a. m. In experiment No. 77, 144.4 grams were passed on the first fasting day at 7h 36m a. m. During the evening of April 9, owing to the discomfort experi- enced by the subject, about 35 grams of fresh feces were removed by means of an enema. No more feces were passed until 24 hours after the first food was taken, i. e., after the subject had left the respiration chamber. In experiments in which the heat production is especially studied, it is desirable in so far as possible to maintain a constant muscular activity. It was found that frequently, especially during the earlier fasts, the subjects would, in many instances, on either the first or second day of the experiment, make ineffectual attempts to defecate. Aside from the disturbing element of the extraneous muscular exertion and exposure of the body, there was the dis- 344 Influence of Inanition on Metabolism. comfort to the subject which might produce abnormal results. Consequently, during the series of 2-day experiments Nos. 79 to 89, warm water enemata were given before the experiments, which resulted in completely removing the contents of the lower bowel so that in but one instance (experiment No. 82) were feces passed during the fasting period. The influence of the character of the diet taken by different subjects is probably considerable. For while the majority of the subjects of these experi- ments did not defecate after food was withheld, S. A. B., on the other hand, passed feces, in some fasting experiments at least, with considerable regularity. This subject commonly consumed rather large quantities of fresh and dried fruits, nuts, milk, eggs, and bread, as is seen by the menus given on pages 277 to 288. From the character of the feces obtained during the nitrogen metab- olism experiments, it is seen that ordinarily he passed relatively large quantities of feces. This is to be expected from the nature of the diet consumed. Hence, in the fasting experiments made with this subject, it is not at all surprising that in some instances at least we find defecation more or less regular on the first days of the fast. It is difficult to explain the fact that feces were passed only once during the 7-day fast on this assumption, unless it be that on the day preceding the fast the subject did not partake of his usual amount of food.34 Making due allowance, then, for the bulky nature of the diet, the influence of inanition in retarding the time of defecation as observed in these experiments is in accord with the observations made in connection with the experiments on Jacques (3) and Succi (4). In all of the experiments made in this laboratory the attempt has been made to recognize and separate any fecal matter that could properly be considered lasting feces. Having due regard for the possibility of the irritating action of solid material of any kind on the intestinal wall, lampblack has been used m all cases in attempting the separations. With short fasting experiments such as those previously made in this laboratory and reported elsewhere,85 the separation of feces was impossible. Indeed, even in experiments with food and with wide alterations in the nature of the diet between periods, it is a difficult matter to separate the feces from a 1 or 2-day experiment. "When it is considered that if we accept Mueller's view that the total dry matter of feces during fasting is 3 grams per day, corresponding to a total weight of fresh feces of 12 grams, it is seen that the separation of fasting feces for one day is practically impossible. With the beginning of the series of experiments in which the fasts were to last more than 2 days, it was hoped that some positive evidence regarding the fasting feces could be obtained. With the single exception of fasting experiment 4 The large nitrogen excretion on this day (p. 367) is opposed to this view. 35 U. S. Dept. Agr., Office of Expt. Sta. Bui. 136. Urine. 345 No. 73, during which 38.4 grams of fresh feces, weighing when air-dry 10.1 grams, were at first considered fasting feces, no definite indication of the formation of fasting feces could be seen. Indeed, during the longest fasting experiment, No. 75, which continued 7 days, it was impossible to distinguish with a reasonable degree of accuracy any fasting feces. While it is undeniably true that the intestinal canal throws off from its walls material as regularly as does the integument, it is highly improbable that any considerable portion of the material thus thrown off enters the large colon as the material thus deposited may be reabsorbed. Probably a certain amount of the epithelial debris from the walls of the large colon normally collects there, and may be considered as fasting feces. The amount thus formed is, however, in all probability very much less than that shown by any measurements thus far recorded. Chemical as well as microscopical examination of all feces passed during fasting experiments considerably longer than these are essential for a proper understanding of the nature of fasting feces. URINE. The chief end products of protein katabolism, at least those end products containing nitrogen, are eliminated in the urine, and while it is impossible to differentiate in the respiratory products between the carbon dioxide and water of protein katabolism, and that of fat and carbohydrate katabolism, a study of the compounds, especially the nitrogenous compounds in the urine, furnishes as accurate a measure of protein katabolism as is yet available. In the studies here reported, analyses of the urine were made with as great a degree of completeness as pressure of other work and the facilities in the laboratory would permit. Unfortunately the urines could not be analyzed with the completeness that characterizes the analytical scheme of Folin. Since the grosser study was more especially that of the gaseous exchange and heat transformations as affected by inanition, the determinations of potential energy in the urine were made. One difficulty which precluded the complete analysis of urine was the inability at times to secure sufficient material for samples. The determinations were invariably made in duplicate and in many instances in triplicate, hence large amounts of urine were necessary. In interpreting the results of the urine analyses, each component of the urine is given special consideration. VOLUME. Complete fasting during which no water is consumed results in lowering in a marked manner the total amounts of urine voided per day. That this is true is borne out by all the experimental data available, though unfortunately the number of instances of complete inanition in which accurate observations 346 Influence of Inanition on Metabolism. regarding the amounts of urine per day were made is relatively small, being confined to the few pathological cases in which the urine has been withdrawn each day by means of a catheter, and. some cases with the fasting insane. Schaefer 36 gives the amount of urine per day in 7 cases with fasting insane women and shows that in 4 cases of fasting without water the volume of urine varied from 550 to 162.5 cc. per day. In some of the cases in which water was taken the amounts of urine fell to 220 cc, while in no case did the quantity exceed 550 cc. per day. In a case of oesophagus stenosis in a 19-year-old girl reported by Mueller 37 the total quantity of urine, when the subject fasted without water, varied on the 4 days of the observation only between 130 cc. and 170 cc. The body weight was, however, unusually small (34.5 to 33.0 kgs.). The report of a case of partial fasting caused by carcinoma ventriculi, published by Seegen,38 shows that the patient took but 35 cc. of fresh cow's milk per day and during 12 days she passed but 2230 cc. of urine. The quantities ranged from 125 to 240 cc. per day. On the other hand, in an experiment made by Sadovyen on the subject " J," fasting without water, the quantity of urine voided was relatively large. On the first day it amounted to 898 cc. and on the second day to 913 cc. With the subject of Hoover and Sollman (8), the volume of urine varied considerably. On the day prior to the fast, the volume was 1350 cc, while on the first day of the fast which was essentially without water, the volume was 570 cc In general, when water is taken during the fast, the volume of urine approaches more nearly that voided by people under normal conditions. Indeed, when moderate amounts of water are consumed, the volume of urine presents as a rule no noticeable abnormalities. Cetti (7) voided in the 10 days of his fast 9433 cc. of urine. The quantities per day varied from 1310 cc to 620 cc. With Breithaupt (7) the volume of urine per day was much larger, varying from 1706 on the third day to 957 on the sixth and last day. Water was consumed on the 5 days of the fast with J. A. in the Stockholm laboratory (9) in the following quantities: 137, 560, 504, 965, and 650 cc The following quantities of urine were voided, 692, 537, 579, 650, and 579 cc. respectively. With Jacques (3) the collection of urine was more or less irregular as he did not always empty the bladder at a definite hour each day. The lowest amounts recorded for individual days were on the last 2 days of the fast, 88 Allgemeine Zeitschrift f. Psychiatrie (1897), 53, pp. 525-537. 37 Zeitschrift fur klinische Medicin (1889), 16, pp. 496-540. 38 Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Classe (1871) 63, Abth. II, p. 429. Volume of Urine. 347 in which the volumes of urine were 450 and 440 cc. respectively. At this period of the fast, the subject was taking about 900 cc. of liquid per day. It is moreover to be noted that since Jacques drank each day a part of his own urine, and as the amounts so drunk are not specified, no correction can be made to obtain the true intake of water. Among the observations on urine the records of the volumes of urine voided by Succi are of unusual interest. In the fast reported by Luciani (4) the volume of urine excreted by Succi remained quite constant. Only on the first day did the amount approximate normal. On the day before the experiment began the volume was nearly 1500 cc, while on the first day of the fast it had fallen to 900 cc, and on the second day to 500 cc. The lowest volume observed was 250 cc, on the 22d clay of the fast. On the first 2 days he took no water, hence the urine excretion is comparable on those days to experiments on fasting without water. During the remainder of the fast Succi partook of water as desired. Table 192. — Volume of urine eliminated daily by fasting subjects. Day of fast. Succi. At Flor- ence.1 At Na- ples. At Vien- na. J. A. Cetti. Breit- haupt. Lan- der- gren.5 J.3 Sohn. Sub- ject 1.5 Sub- ject II.5 Flora Tosca. Kel- lei.s Last food day. 1.. 2.. 3.. 4.. 5.. 6.. 7.. 8.. 9.. 10.. cc. cc cc. 1475 810 .... 900 960 1435 500 590 830 550 660 575 525 630 612 600 730 612 550 620 563 500 610 509 450 425 578 425 460 640 400 410 450 cc 1350 692 537 579 650 579 cc. 1150 990 940 1080 1310 980 945 995 790 775 620 cc 815 1403 1232 1706 1263 991 957 cc 1500 770 546 58S 583 cc. 2295 2490 1593 2613 cc 1350 570 470 530 4 1020 410 560 820 cc 1500 2780 2820 1540 cc 1460 970 1080 {::: cc 909 456 614 1115 853 616 594 509 775 712 619 cc 1020 460 1220 1540 1 Estimated from curve given by Luciani (4). 2Skan. Archiv. f. Physiol. (1903), 14, p. 112. 3 Reported by A. Sadovyen (2). 4 Amount for 2 days. 5 Reported by Scbreiber and Waldvogel. 6 Reported by A. Keller, Zeit. f. physiol. Chemie (1900), 29, p. 165. In the Naples fast (21 days) reported by Ajello and Solaro (6), the volume of urine voided by Succi varied from 960 cc. on the first day to 410 cc on the tenth and fourteenth days. The drinking-water varied in amount from 2170 cc. on the thirteenth day to 620 cc on the twentieth day. After the first 9 days of the fast the subject took regularly about 30 grams of citrate of magnesium. On 4 days from 200 to 300 cc of a saline water (Vichy eleventh day, Janos 12, 20, 21) were taken. 348 Influence of Inanition on Metabolism. In the Vienna fast (21 clays) reported by E. and 0. Freund (10) the volumes of urine varied from 1435 cc. on the first day to 235 cc. on the 21st day. The records of the amount of drinking-water are not given. The volumes of urine from the 23d to the 30th day of Succi's fast at Hamburg, reported by Brugsch (12), are twice as large as those of the last 10 days of the Vienna fast. In the Hamburg fast, Succi voided 1030 cc. of urine on the 30th day, and the smallest volume recorded was 520 cc. on the 29th day. The subject drank about 750 cc. of water each day of the fast. This amount of liquid was increased on the 30th day by 300 cc. of a sweetened lemonade. A tabular statement showing the daily volumes of urine voided by the subjects of the fasts discussed above, as well as those of a professional fasting woman (11), is given herewith in table 192. Vozarik39 in studying the influence of the character of the diet on the quantity of urine excreted, observed volumes excreted on single fasting days amounting to 905, 1405, 2240 cc, respectively. Since, as the author clearly points out, the character of the diet before the fasting period influenced in large measure the volume of urine excreted, and since unfortunately the nitrogen excretion was not determined, the results are of but little value in this discussion. Not only have the exact data of the amounts of urine, and the water consumed, been recorded in the Middletown experiments, but the amounts of water in urine and the daily ratios of the water of urine to ingested water have also been obtained. The results are presented in table 193, page 355. For convenience of discussion, and since the data above show that the volume is proportional to the drinking-water to a certain extent, the con- sumption of drinking-water is considered before proceeding to the particular point under discussion, i. e., the volume of urine. RELATION OF DRINKING-WATER TO VOLUME. The amount of drinking-water consumed by the subjects in different experi- ments varies greatly. In the experiments reported herewith the maximum consumption is 2763.2 grams on the third day of experiment No. 73, and the minimum 115.10 grams on the first day of experiment No. 89. Even in experiments with the same subject the variations are considerable. For example, in experiment No. 73 with S. A. B., the average amount of drinking water per day was 2124.74 grams, while in experiment No. 77, with the same subject, the average was 1405.45 grams. The maximum amount was consumed by this subject on the third day of experiment No. 73, 2763.20 grams, and the smallest amount on the last day of experiment No. 77, 962.00 grams. Experiments Nos. 68 and 69, both of which were made with another subject (A. L. L.), also show marked differences in the amount of water consumed. "Archiv f. die ges. Physiologie (1906), 111, p. 526. Volume of Urine. 349 With regard to experiment No. 68, however, it is to be noted that it was preceded by a metabolism experiment in the respiration chamber and the records show that the quantity of water consumed during the preceding days was probably much less than that consumed on the days preceding experiment No. 69. On the average much smaller amounts of drinking-water were taken by the subjects in the series of 2-day fasts, Nos. 79 to 89. With regard to the apparently excessive amounts of drinking-water consumed by S. A. B., it is to be noted that he was especially cautioned to consume considerable amounts of water. In some of the earlier experiments, especially those with A. L. L., an unusually high specific gravity was observed in the urine, and moreover the volume was insufficient to enable all the desired analyses to be made. It was considered also that the fast might be endured with less liability of causing discomfort if the volume of urine was kept relatively large and the specific gravity relatively low. Furthermore, it had been the custom of S. A. B. during his fasts in private to consume rather large amounts of water. Having seen from the above statement that the amounts of drinking-water vary considerably with different subjects, it is of interest to note the amounts cf urine passed and especially the water of urine. An inspection of the data on this point shows very wide fluctuations from experiment to experiment and even between the different experiments with the same individual. The lowest average weight per day of water in urine was in experiment No. 89 where there were but 545.89 grams. The largest daily elimination occurred on the second day of experiment No. 73, and amounted to 2928.23 grams. In experiment No. 69, there was an average of nearly 300 grams less water in the urine per day than in experiment No 68 with the same subject. The average amounts of water eliminated per day in the urine of the subject S. A. B. was more nearly constant, i. e., 1767.01 grams for experiment No. 71; 2156.51 grams for experiment No. 73; 1709.08 grams for experiment No. 75; and 1818.35 grams for experiment No. 77. The relationship between the amount of water in the urine and the amount of water ingested is of special significance. In column e of table 193 the ratios of the weight of the water in the urine to the weight of the ingested water are recorded. The wide variations in the amounts of water consumed and the water of urine noted in all the experiments are clearly indicated by these ratios. The lowest recorded for any given day is 0.628 on the first day of experiment No. 82. The highest ratio observed was that on the first day of experiment No. 80, when the water in the urine was over 8 times that ingested, i. e., 8.326. It is noticeable that both extremes in these ratios are found in the data for the shorter fasting experiments. When the differences in body condition previous to a fast are taken into consideration, it is not at all surprising that there are 350 Influence of Inanition on Metabolism. wide fluctuations in the amounts of urine voided, and therefore while the data for the 2-day experiments are of distinct value it is upon the results for the longer experiments that we must especially rely in discussing these ratios. In all the experiments continued 3 days and over, the ratios are much more nearly constant. Thus, the lowest average ratio is 0.763 for the 4-day experi- ment No. 69, while the highest is 1.294 in the 4-day experiment No. 77. The individual days in these longer experiments exhibit differences ranging from 0.643 on the first day of experiment No. 69 to 1.492 on the third day of experiment No. 77. Considering the long experiments with the same subject, namely, Nos. 71, 73, 75, and 77, the ratios range for the averages of the experiments from 0.941 to 1.294. A close examination of the data shows that in general on those individual days in which the amount of drinking-water is over 1000 grams, the ratios vary from 0.659 to 1.592, and for those experiments in which the average amount of water consumed per day was ever 1000 grams, the ratio varies from 0.846 to 1.294. Furthermore, when the amount of drinking-water on one day is materially larger than that on the following day, the ratio on the second day tends to increase. This is especially noticeable in the 2-day experiments. The results of these comparisons show, therefore, that the amount of water in urine and consequently the volume of urine is largely dependent upon the amount of water consumed. The generality of this rule is, however, strikingly interfered with in certain instances, notably those in which the ratios are the highest. Thus, on the first day of experiment No. 80, while there were but 132.80 grams of water consumed, the urine contained 1105.75 grains, and on the first day of experiment No. 89, there were but 115.10 grams of water consumed and the urine contained 599.55 grams of water. Even in those experiments in which the lowest ratios obtained, the quantities of urine voided are not unusually small for fasting experiments. Thus on the first day of experiment No. 82, when the ratio was but 0.628 there were 538.57 grams of water in the urine, although the subject had actually consumed 857.80 grams of water during the day. The anomalies noted in tins series of experiments almost invariably occur on the first day, a striking contrast to the records of Luciani. According to Luciani's table 1700 cc. of water were consumed on the 29th day of Succi's fast in Florence, while the volume of urine was but 350 cc. The retention of large amounts of water at the end of a fast has been frequently observed, and on the 10th and 13th days of Succi's fast at Naples the amounts of water consumed were 2025 and 2170 cc, accompanied by a urine excretion of but 410 and 480 cc, respectively. In general, then, during the earlier stages of a fast, with the exception of the first day, the volume of urine is in large measure determined by the quantity Specific Gravity of Urine. 351 of drinking-water consumed. If the volume of ingested water is small the volume of urine may exceed it several times. When the volume of drinking- water is over 1000 cc. the volume of urine is usually not far from that of the water consumed. REACTION. In all of the samples of urine, whether tested by periods or for the whole day, the reaction was acid. The pressure of other work prevented an accurate determination of the degree of acidity. According to Brugsch (12), however, the acidity, at least in the later stages of a prolonged fast, remains nearly constant from day to day. SPECIFIC GRAVITY. Normal urines vary widely in their specific gravity, large volumes of urine being generally accompanied by a low specific gravity and small volumes by a high specific gravity. In the first three experiments here reported the specific gravity was taken with a carefully calibrated urinometer with which, however, it was practically impossible to make accurate record to the fifth significant figure. In all subsequent experiments a Westphal balance was used. For purposes of comparison it is important that the Lemperature of the urine be constant when the specific gravity is taken, and hence all determinations were made at 20° C. If the urine was colder than 20° it was warmed by an electric heater. It is probably true, since the specific gravities in the experiments of earlier observers are as a rule recorded only to the third decimal, that a simple hydrom- eter was used. For comparing the different days of the same experiment, this method is fairly satisfactory, but it obviously renders difficult any satisfactory comparison between experiments not made in the same laboratory. In the case of Breithaupt, the specific gravity remained very constant from day to day, the lowest being 1.0110 on the second and third days, and the highest, 1.0135 on the fifth day. Unfortunately the records of specific gravity given for Breithaupt's urine show the specific gravity not of the urine as voided but of the volume of urine to which the original 24-hour amount of urine was diluted before the specific gravity was taken. Hence the results are for purposes of comparison of little value. The specific gravity of Succi's urine, in the fast at Naples, varied from 1.033 to 1.026, remaining for the most part very constant at about 1.028. In Succi's Vienna fast the specific gravity, which was 1.023 and 1.020 on the first and second days, respectively, remained almost constant at 1.030 during the remainder of the fast. Brugsch (12), in studying the urine from Succi during the fast at Hamburg, records the specific gravity. During the last days, 23d to 30th, the specific gravity varied only from 1.023 to 1.026, remaining for 5 successive days at 352 Influence of Inanition on Metabolism. 1.025. The volume of urine during this period varied from 530 to 600 cc. On the last day, however, the volume rose to 1030 cc., with a specific gravity of 1.026. The urine of the fasting woman Flora Tosca (11), varied in specific gravity from 1.0315 on the second fasting day to 1.023 on the last (15th) fasting day. The decrease was gradual as the fast progressed. The specific gravities of the urine for the different days of the experiments here reported are given in column / of table 193. The highest observed on any day was that on the second day of experiment No. 89, i. e., 1.0338. The lowest for any day was on the second day of experiment No. 73, 1.0032. In the first case, the volume of urine was 522 cc, while in the second it was 2958 cc. On certain of the periods even lower specific gravities were observed than that in experiment No. 73. Thus, on referring to table 94, it will be found that the specific gravity of the urine for the period from 1 p. m. to 7 p. m., January 30, was 1.0024. The volume of urine during this period was 888 cc. The highest specific gravity, 1.0360, observed in any period was on the first day of experiment No. 69. The amount of urine for this period was 63.1 grams. The urines of the subject S. A, B. are characterized by an exceedingly low specific gravity occasioned by the large volume of urine incident to the con- sumption of so large an amount of drinking-water. Aside from these exception- ally low records all the specific gravities observed come well within what would be termed normal limits. There is nothing like the constancy exhibited in the fasts of Succi in any of the experiments save Nos. 71 and 75. total solids. The volume and specific gravity are of interest only because together they afford a means of measuring the relative amounts of total solids in the urine, which latter have not often been determined directly by investigators. Since it seemed desirable to know more of the actual amount of these solids, especially during fasting, direct determinations of them were made in the experiments here reported. The determination of the total solids or water-free material was made as a rule by drying the urine in a vacuum to constant weight. Although there are unquestionably errors in this method, for purposes of relative comparison between different days of a given experiment or between different experiments, the data are of value. The largest amount of total solids recorded in any experiment is that on the second day of experiment No. 82, on which there were 54.93 grams 40 eliminated. The smallest amount recorded 40 While the data given in column g of table 193 show that in experiment No. 59 there was an elimination on the third day of 57.70 grams, this is hardly to be considered as an accurate statement of the actual weight of solids for this day, since it was determined indirectly by calculation as described on p. 28. The same is true of the results on the 2 days of experiment No. 68. However, the total solids for the average both for experiment No. 59 and No. 68 were determined directly on a total composite sample of urine for each experiment. Solid Matter in Urine. 353 is on the first day of experiment No. 71 with S. A. B., when less than half this amount, i. e., 25.51 grams, was eliminated. Considering the averages of the experiments, the lowest amount recorded is in experiment No. 71, 34.99 grams, and the highest amount is in experiment No. 59, 52.93. In general the average amount of total solids during the different experiments is not far from 40 grams per day. On the individual days of the same experiment, there is usually- an increase on the second day after which the amount remains fairly constant. An inspection of the data of the different experiments with the same subject, S. A. B., shows an average daily elimination of total solids as follows : Experi- ment No. 71, 34.99 grams; experiment No. 73, 37.13 grams; experiment No. 75, 44.22 grams; and experiment No. 77, 51.35 grams. Leaving out the first day of each of these experiments, since as a rule it is lower than the other days, the averages become 38.15, 36.79, 45.15, and 54.42 grams respectively. Conse- quently, it appears that in experiments Nos. 71 and 73, the subject S. A. B. eliminated practically the same amount of total solids each day. In experiment No. 75, there was an average elimination of nearly 10 grams more, while in experiment No. 77 the average amount per day was about 10 grams greater than that during experiment No. 75. While the average amount per day of total solids in the series of 2-day experiments is not far from 40 to 45 grams, yet individual fluctuations may be very considerable as, for example, in the 2-day experiment No. 82, in which on the first day there were 37.13 grams, and on the second, 54.93 grams. Ratio of the Total Solids to the Specific Gravity. Knowing the volume of urine excreted per day and having an accurate record of the specific gravity, it is possible by factors to compute in many instances the amount of total solids. This method of computation has frequently been used for the approximations of the clinician. The various organic and inorganic substances dissolved in normal urine have widely varying physical properties and hence solutions of varying density. A solution of 100 grams of sodium chloride in a liter of water has a density at 15° of 1.073, while a solution of urea 1 to 10 has a density of but 1.028. With ordinary conditions of diet, however, it has been found that the total solids in 1 liter of urine may be reasonably approximated by multiplying the last two figures of the specific gravity (as ordinarily expressed in three decimal places) by 2.33. Thus 650 cc. of urine of a specific gravity of 1.027 would contain total solids as follows : 27 X 2.33 X -650 = 40.89. It is stated by some writers, that if the specific gravity is less than 1.018, more accurate results can be obtained by using the factor 2.0. In normal urines, therefore, this factor is by no means constant. Since determinations were made of volume, specific gravity and total solids in the 23 354 Influence of Inanition on Metabolism. urines from the experiments here reported the factor for urines from fasting men may be found from these data. In any attempt to secure an accurate ratio between the amount of total solids and the specific gravity, it is important that all specific gravities should be taken at the same temperature. Each specific gravity was taken at 20° in the Middletown experiments. The volume of urine was not measured but was computed from the weight and the specific gravity. Hence, any errors that involve the determination of specific gravity would cause an error in the volume. Usually, however, it is difficult to see how any appreciable error on the volume could be made. The ratio (r) existing between the total solids and the specific gravity may be expressed by the following formula: r- I b X (tp. gr. — 1) in which (g) is the total weight of solids in volume (&) of urine. The ratios thus computed for the different days and experiments are recorded in table 193. The lowest average ratio is that of experiment No. 68, 2.1; the highest is that of experiment No. 73, 3.4. So far as the individual days are concerned, the lowest ratio is that of the first day of experiment No. 80, 2.2, and the highest, the second day of experiment No. 73, 4.1. In considering the values for the ratios, it must be borne in mind that in experiments Nos. 59 and 68, the total solids for individual days were apportioned by calculation. The average ratio for all of the experiments is somewhat higher than that commonly assumed for normal urines. In experiments with the same individual, there are also marked differences. The ratio for experiment No. 69 is higher than that for experiment No. 68, which was made with the same individual some 8 months earlier. In the series of experiments with S. A. B., the average ratio for the first three is fairly constant, 3.0, 3.4, and 3.3. In the last experiment (No. 77), the average falls to 2.5. In the longer fasts (experiments Nos. 69, 71, 73, 75, and 77) the average ratio is 2.9. Obviously during a fast, the relative amounts of organic and inorganic matter in the urine may vary considerably from those obtained when food is eaten. The amount of sodium chloride taken with the food, for example, may affect considerably the specific gravity. As has been pointed out, the specific gravity increases much more in proportion with an increase of sodium chloride than with urea. The larger the proportion of organic matter which consists in large part of urea, the lower the specific gravity with refer- ence to a given weight of total solids and hence the higher the ratio. Thus a knowledge of the relative amounts of organic and inorganic matter in the total solids is essential to an understanding of the variations in the relationship between specific gravity and total solids during fasting. Solid Matter in Urine. 355 Table 193. — Relations between amounts of drinking-water, icater in urine, specific gravity, and total solid matter in metabolism experiments ivithout food. Ex- peri- ment num- ber. Subject and date. (a) Am'nt of water con- sumed. (6) Vol- ume of urine. (c) UD Am'nt Water of in urine, urine. (e) Ratio of water in urine to water con- sumed (c-^a). if) Spe- cific grav- ity. (0) Total solids (h) Ratio of total solids to spe- cific grav- ity. Proportion of total solids. (i) Ash. U) Or- ganic mat- ter. 59. 68. 69. 71. 73. B.F.D.,Dec.l8, 1903.. Dec. 19, 1903... Dec.20,1903... Total 3 days. Av. per day . A.L.L.,Apr. 27-28,1904 Apr. 28-29,1904 Total, 2 days .... Av. per day A.L.L.,Dec.l6-17,1904 Dec. 17-18, 1904 Dec. 18-19, 1904 Dec. 19-20, 1904 Total, 4 days — Av. per day 76 77. S.A.B.,Jan. 7-8, 1905 Jan. 8-9, 1905 Jan. 9-10,1905 Jan. 10-11, 1905 Total, 4 days.. Av. per day . . S.A.B.,Jan.28,1905.. Jan. 29,1905.. Jan. 30, 1905.. J an. 31, 1905.. Feb. 1,1905.. Total, 5 days.. Av. per day... Gm 1342.50 1360.20 1188.00 3890.70 1296.90 c.c. 1517 962 908 Gms. 1540.80 981.80 928.90 Gmx. U494.50 i 927.00 i 871.20 1.113 .682 .733 1.0160 1.0210 1.0230 Gms. U6.30 154.80 157.70 1.9 2.7 2.8 P.Ct, 3387 1129 3451.50 1150.50 3292.70 1097.57 .'846 .... 158.80 52.93 k'.h .... 268.40 616.20 884.60 442.30 979 822 1801 901 1001.70 843.80 1845.50 922.75 i 956.40 i 795.70 1752.10 876.05 3.563 1.291 1.981 1.0230 1.0265 145.30 148.10 93.40 46.70 2.0 2.2 2.1 647.03 553.60 1085.30 828.50 437 572 743 725 3114.43 2477 778.61 619 S.A.B.,Mar. Mar. Mar. Mar. Mar. Mar. 4, 1905. 5, 1905. 6, 1905. 7, 1905. 8, 1905. 9, 1905. Mar. 10, 1905. . Total, 7 days... Av. per day S.A.B., Apr. 8,1905. Apr. 9,1905. Apr. 10. 1905. Apr. 11, 1905 . Total, 4 days... Av. per day — H.E.S., Oct. 13, 1905.. Oct. 14, 1905.. Total, 2 days... Av. per day — 1204.40 1973.20 2312.90 1485.30 6975.80 1743.95 2,082.30 2,746.90 2,763.20 1,955.40 1,075.90 2,252 2,958 2,725 1,953 1,026 2,264.30 2.225.81 2,966.80 2,928.23 2.735.20, 2,698.27 1,964.10 1.928.35 1,037.80 1,001.89 10,623.70 10,914 10 968.2010,782.55 2,124.74 2,183 I 2.193.64 2,156.51 2048.20 1592.90 1018.70 962.00 5621.80 1405.45 2552 2156 1554 1136 7398 1850 2570.30 2528.15 1.234 2177.601 2122.94' 1.333 1574.30 1520.30 1.492 1156.60 1102.01 1.146 7478.80 7273.40' .... 1869.70 1818.35 1.294 1.0072 1.0101 1.0132 1.0178 42.15 54.66 54.00 54.59 205.40 51.35 2.3 2.5 2.6 2.7 2.5 782.50 399.90 1182.40 591.20 1015 840 1855 928 1027.20 859.10 996.08 809.96 lssii.:ii"i 943.15 1806.04 903.02 1.273 11.0118 2.025 ! 1.0223 1.527 31.12 49.14 2.6 2.6 80.26- 40.13 2.6 33 22 20 17 23 28 23 25 1 Not determined for individual days, calculated as shown on p. 28. P.ct. 451.80 415.75 590.10 545.08 762.30' 714.81 742.10; 700.32 .643 .985 .659 .845 1.0330 1.0310 1.0250 1.0230 36.05 45.02 47.49 41.78 2.5 2.5 2.6 2.5 18 14 14 13 2546.301 2375.96 636.58; 593.99 .'763 .... 170.34 42.59 'k'.'t "is 1149 2009 2529 1473 1159.50 1133.99 2021.10 1984.11 2541.60 2500.68 1485.80 1449.25 7208. 001 7068.03 1802.00 1767.01 .942 1.006 1.081 .976 1.0089 1.0058 1.0048 1.0087 25.51 36.99 40.92 36.55 2.5 3.2 3.4 2.9 30 16 14 16 7160 1790 i.oih .... 139.97 34.99 '3.0 "19 1.069 1.0057 38.49 3.0 26 1.066 1.0032 38.57 4.1 16 .976 1.0036 36.93 3.8 16 .986 1.0056 35.75 3.2 17 .931 1.0119 35.91 2.9 17 185.65 1.615 37.13 3.4 is 1,973.30 1,496 1,508.20, 1,469.59 .745 1.0079 38.61 3.3 16 1,728.90 1,871 2,117.60 2,208 1,885.70 1 839.88 1.064 1.0077 45.82 3.2 16 2,221.00' 2,175.69 1.027 1.0057 45.31 3.6 14 1,911.30 1,986 1,999.80 1,953.00 1.022 1.0070 46.80 3.4 17 1,581.00 1,423 1,437.50, 1,391.64 .880 1.0100 45.86 3.2 16 1,702.00 1.611 1,624.40, 1,580.70 .929 1.0081 43.70 3.3 13 1,693.10 1,584 1,596.50 1,553.08 12,273.1011,963.58 .917 1.0080 43.42 3.4 12 12,707.80 12,179 309.52 1,815.40: 1.740 1,753.30! 1,709.08 .941 44.22 3.3 is 82 86 86 87 85 70 84 86 84 81 74 84 84 83 83 81 84 84 86 83 84 87 85 67 78 80 83 77 72 77 11 356 Influence of Inanition on Metabolism. Table 193. — Relations beticeen amounts of drinking-water, water in urine, specific gravity, and total solid matter in metabolism experiments without food — Continued. Ex- peri- ment num- ber. Subject and date. (a) Am'nt of water con- sumed. (b) Vol- ume of urine. (C) Am'nt of urine. (d) Water in urine. • • • 92.06 46.03 'z.'h "28 "72 1467.10 884.20 2351.30 1175.65 1187 1046 1204.50 1065.00 1159.21 1012.39 .790 1.145 1.0146 1.0186 45.29| 2.6 52.6l| 2.7 18 17 82 83 2233 1117 2269.50 1134.75 2171.60 1085.80 .'924 .... 97.90 .... 48.951 2.7 "ii "81 704.50 707.70 1412.20 706.10 mm 666 2 34.62 1188.00 679.40 2 33.37 1145.11 642.44 1820.92 910.46 1.673 .908 1.0154 1.0203 21.25 42.89 36.96 2.4 2.7 29 15 71 85 1836 918 1902.02 951.01 L289 .... 81.10 40.55 'z.k "22 "78 115.10 357.20 472.30 236.15 626 522 644.40 639.60 599.65 492.22 5.209 1.378 1.0296 1.0338 44.85 47.38 2.4 2.7 30 16 70 84 1148 574 1184.00 692.00 1091.77 545.89 2^312 — 92.23 46.12 "z'.'s "*23 "77 1209.58 1350 1367.44' 1323.77 1.095 .... 43.67 2.8 3 21 3 79 1 Includes volume of urine calculated as lost. 2 Calculated. See p. 243. 3 For 38 days. Constituents of Total Solids. To interpret intelligently the elimination of total solids, it is necessary to make a more careful study of the constituents of the solid matter in urine. The elimination of the total quantity of solids in the urine during inanition is affected in no slight measure by the amounts and character of the food for the day or two preceding the fast. This is especially true on the first days of fasting. If considerable amounts of sodium chloride are taken with the food, this will be excreted more or less rapidly on the first days of the fast. It is important, therefore, to determine if possible the nature of the total solids Ash in Urine. 357 eliminated. It is very much to be regretted that complete proximate analyses of urine could not have been made, as such data would undoubtedly make much clearer many troublesome points which arise in interpreting the results of the partial urinary analyses that were possible. Ash. — In addition to the total solids, the ash was determined by incinerating the dried material at as low a temperature as possible, extracting the charred mass with water, and igniting the carbonaceous material. The aqueous extract was evaporated to small bulk, added to the ignited residue, and after com- pleting the evaporation, the mass was dried for a few moments at a low red. heat. Admittedly the errors in this determination are such as to make it only approximate, and the results are valuable only for the purposes of comparison, but since the same method was followed with all experiments, comparisons probably have more value than would be the case if different, though even more accurate methods, had been employed to determine the total solids and ash in the different experiments. The quantities of total solids are recorded in column g of table 193. The total ash as determined is recorded in table 194. Table 194. — Ash excreted in urine in metabolis m, experiments without ; food. Exper- iment Subject and duration of First Second Thirc 1 Fourth Fifth Sixth Seventh num- ber. experiment. day. day. day. day. day. day. day. Grams. Grams. Grams . Grams. Grams. Grams. Grams. 59 B.F.D., Dec. 18-20, 1903.. 1 6.94 1 8.20 18. 6r r .... .... 68.... A.L.L., Apr. 27-28, 1904.. U0. 91 U1.60 . . . .... .... . . .... 69 A.L.L., Dec. 16-19, 1904.. 6.64 6.43 6.5( 5 5.57 .... . . .... 71 S.A.B., Jan. 7-10, 1905.. 7.65 6.06 5.8! ) 5 . 94 .... , . .... 73 ... . S.A.B., Jan. 28-Feb. 1, 1905 9.96 6.23 5.7' I 6.09 6.02 . . .... 75 S.A.B., Mar. 4-10, 1905.. 6.03 7.54 6.4< { 7.80 7.48 5 85 5.27 77 S.A.B., Apr. 8-11,1905.. 13.88 11.98 10.5! ) 9.14 .... 79 H.E.S., Oct. 13-14, 1905.. 8.73 11.17 • • a .... . . .... 80 C.R.Y.,Oct. 27-28, 1905. . 18.93 12.52 . . ... .... 81 A.H.M.,Nov.21-22, 1905.. 12.61 11.22 • . . . . .... 82 H.C.K., Nov. 24-25, 1905.. 9.67 16.33 . . . . . .... 83 H.R.D., Dec. 5- 6, 1905.. 8.19 8.73 . . .... 85 N.M.P.,Dec. 9-10, 1905.. 212.83 5.57 . . . . . 89 D.W., Jan. 10-11, 1906. . 13.34 7.45 .... 10.45 9.36 7.3< ) 6.91 6.75 5.85 5.27 1 Not determined for individual days, calculated as shown on p. 28. 2 Includes amount calculated in urine spilled. See p. 243. As has been pointed out before, the ash in experiments Nos. 59 and 68, was apportioned over the different days, and hence the figures do not represent actual determinations. In all subsequent experiments, the ash and total solids for each day were determined. Delaying for the present the discussion of the constituents of ash, the total ash determinations are of value in showing what proportion of the total 358 Influence of Inanition on Metabolism. solids were of inorganic nature. The lowest amount of total ash recorded is on the last day of experiment No. 75, 5.27 grams; the highest amount is on the first day of experiment No. 80, 18.93 grams. While the amount of ash per day remains fairly constant in the longer fasting experiments, and except in experiments No. 68 and No. 77, never exceeds 10 grams per day, in the series of 2-day fasts, beginning with No. 79, much larger amounts of ash are eliminated, the lowest average being 8.46 grams, in experiment No. 83 and the highest, 15.73 grams, in experiment No. 80. In the case of all the long fasting experiments except Nos. 68 and 77, the ash elimination is not far from 6 to 7 grams per day. On the first day of experiment No. 73, there is an unusually large amount of ash, 9.96 grams, while on the second day, it immediately falls to 6.23 grams. In experiment No. 68, the ash, although possibly erroneously distributed over the two days, is nevertheless for the average of the two days relatively large, and in experi- ment No. 77, the ash elimination varies from 13.88 grams on the first day to 9.14 grams on the last day. Comparing experiments with the same subject, it may be seen that the ash in experiment No. 69 is but a little more than half as large as in the earlier experiment, No. 68, while with the series of experiments with S. A. B., the ash is practically constant at not far from 6 grams for all save the last experiment, No. 77, in which the average elimination is over 11 grams. In the series of 2-day experiments, the ash is usually smaller on the second day than on the first, although marked exceptions to this rule are seen in experiment No. 79, and especially No. 82. It is, however, necessary to consider the data regarding the elimination of sulphur, phosphorus and chlorine for an intelligent interpretation of the elimination of ash. The only data regarding the ash elimination during fasting with which we are familiar are the quantities in the urine of J. A. (9). On the last day with food the total ash of urine amounted to 23.0 grams; on the 5 fasting days the total ash eliminated was 14.7, 6.7, 5.7, 5.0, and 4.5 grams, respectively. Although complicated by the fact that considerable amounts of sodium chloride were taken with some meat extract on each day of the experiments reported by Pettenkofer and Voit,41 the ash determinations are here given. For the three one-day fasting experiments the amounts were 19.7, 18.89, and 14.40 grams, respectively. Deducting the salts in the meat extract and the weight of sodium chloride used, the subject lost 2.1, 2.9, and 1.0 grams of ash per clay in the three experiments. Organic matter. — While as has been shown in the discussion regarding the total ash, the mineral constituents of fasting urines may vary considerably in "Zeit. f. Biologie (1866), 2, p. 479. Organic Matter in Urine. 359 amount, especially on the first days of a fast, it is important to see what variations exist in the actual amounts of organic matter excreted per day. Since the organic matter is made up in large part of nitrogen containing material, it may be taken as a rough measure of the amount of protein metab- olism, and therefore measurements of this total amount are of especial interest. The total organic matter of the urine is estimated by deducting the weight of ash from the weight of total solids, and hence the values thus obtained are affected by the errors incidental to the determinations of both ash and total solids. In determining total solids, it is probably true that the analyses give results which are somewhat too low, since even in drying in a vacuum, there is unquestionably loss of material. It is likewise true that in the determination of crude ash there must be more or less volatilization of inorganic material which tends to make the results for crude ash too low. Hence it is seen that the two most striking errors in the determinations of these two factors may be said to more or less compensate when the values thus obtained are used for the indirect determination of the total organic matter. It is reasonable to suppose then, that the determinations of organic matter are approximately correct. Furthermore, as has been pointed out previously, since the same method was employed for all experiments the results are strictly comparable. Table 195. — Total organic matter excreted in urine in metabolism experiments without food. Exper- iment num- ber. Subject and duration of experiment. First day. Second day. Third day. Fourth day. Fifth day. Sixth day. Seventh day. 59 B.F.D., Dec. 18-20, 1903.. Grams. •39.36 Grams. 1 46.60 Grams. 1 49.03 Grams. Grams. Grams Grams. 68 A.L.L., Apr. 27-28, 1904. . » 34.39 i 36 . 50 ... - . . . . . • • • .... . , 60 A.L.L., Dec. 16-19, 1904.. 29.41 38 . 59 40.93 36. 21 . . • . .... , . 71 S.A.B., Jan. 7-10, 1905.. 17.86 30.93 35.07 30. 31 > . • .... , , 73 S.A.B., Jan.28-Feb. 1, 1905 28 . 53 32.34 31.19 29. 56 29 .89 .... . , 75 S.A.B., Mar. 4-10, 1905. . 32.58 38.28 38.87 39. 30 38 .38 37.85 38 15 77 S.A.B., Apr. 8-11, 1905.. 28.27 42.68 43.45 45. 15 , . 79.... H.E.S., Oct. 13-14, 1905.. 22.39 37.97 . . • ■ • . . • . . 80 C.R.Y.,Oct. 27-28,1905.. 22.72 30.08 • • • • .... . . 81 A.H.M.,Nov. 21-22, 1905.. 26.13 36.82 . . ■ • . • 82.... H.C.K.,Nov. 24-25, 1905.. 27.46 38.60 .... .... . , 83 H.R.D.,Dec. 5- 6, 1905.. 37.10 43.88 • • • • . . . • . . 85 N.M.P.,Dec. 9-10, 1905. . 231.31 31.39 ■ ■ ■ ■ .... . . 89 D.W., Jan. 10-11, 1906.. Average 31.51 39.93 • 29.22 37.47 39.76 38.19 34-14 37.85 38 , 1 S 1 Not determined for individual days, calculated as shown on p. 29. 2 Includes amount calculated in urine spilled. See p. 243. The amounts of organic matter found by deducting the total ash from the total solids are recorded for each day of each experiment (table 195). It is here to be noted that the apportionments of the total organic matter on 3G0 Influence of Inanition on Metabolism. the different days of experiments Nos. 59 and 68, are open to the same objection as has been pointed out before, namely, that they are mathematical apportionments and do not represent actual determinations. The irregularity in the amounts of ash elimated on the different days of the different fasts, is likewise noted in the amounts of total organic matter. The lowest amount of organic matter, 17.86 grams, is on the first day of experi- ment No. 71, while the highest amount, 49.03 grams, is that mathematically apportioned to the third day of experiment No. 59. The largest amount actually determined, 45.45 grams, is on the last day of experiment No. 77. These limits, however, do not give a correct idea as to the relative constancy which seems to obtain in almost all the experiments with regard to the total amount of organic matter eliminated. In general the amount ranges some- where between 30 and 40 grams. The greatest discrepancy generally appears on the first day of the longer fasts, after which the amount remains fairly constant for the different days of the fast. In all the experiments, the amount is larger on the second day than on the first. In the fasts with S. A. B., the amounts of organic matter per day, especially after the first day are worthy of note. Excepting the first day, in experiments Nos. 71 and 73 the average amount is not far from 30 grams. It is about 38 grams in experiment No. 75, while in experiment No. 77, the total amount of organic matter per day for the last three days rises to nearly 44 grams, thus indicating that although on the second and succeeding days of fasting the amount is relatively constant, there may be wide variations during different fasts with the same subject, variations fully as wide as those obtained in the 2-day fasts with different subjects. It is noticeable that while the variations from day to day in the same fast after the first day are insignificant, such gross variations between different experiments with the same subject obtain. The determinations of total solid matter and ash in the urine of three one- day fasting experiments reported by Pettenkofer and Voit42 permit a compu- tation of the amounts of organic matter. On the three days there were excreted 29.82, 25.26, and 31.89 grams of organic matter, respectively. Further discussion of the organic matter of urine will be found with the data for the amounts of nitrogen, carbon, hydrogen of organic matter, and the proximate constituents of the urine. Proportions of Ash and Organic Matter in Total Solids. The greater amount of mineral matter excreted on the first days of the fast produces wide variations in the proportions of organic and inorganic consti- tuents of the total ash. In columns i and ; of table 193, the proportions of ash and organic matter in the total solids are given. With the aid of these proportions the wide variations in the ratio between the total solids and the "Loc. cit Nitrogen in Urine. 361 specific gravity discussed on page 354 can best be interpreted. The proportion of ash. in total solids is as a rule, greatest on the first day and markedly less on the second day. In the longer fasts the proportion is relatively constant after the second day, although in experiment No. 77, an unusually high percentage of mineral matter was excreted on all four days. A ratio apparently exists between the proportion of mineral matter shown in column i and the relation between specific gravity and total solids as given in column h of the table. The higher the percentage of ash in the total solids, the lower the ratios shown in column h. This is especially noticeable in considering different days of the same experiment, but does not obtain for different experiments even with the same subject. In the shorter fasts the ratio between columns i and h is more nearly uniform. Indeed, even from experiment to experiment and with different subjects, the general rule may be noted that the higher the data in column i, the lower the corresponding data of column h. Thus the highest percentage of ash is 45 on the first day of experiment No. 80, and the lowest relation observed in the shorter experiments was 2.2 on the same day. Similarly, while the lowest percentages of ash occurred on the second days of experiments Nos. 83, 85, and 89, the highest ratios of total solids to specific gravity were observed on these days. These observations are fully in accord with the well known fact that the density of solutions of sodium chloride (a typical urine salt) is greater than that of solutions containing equal weights of urea (a typical organic urinary constituent). NITROGEN. The elimination in the urine of the partially oxidized protein in the form of a number of nitrogenous products has given to the determination of nitrogen an especial significance. In all the earlier fasting experiments, the study of the nitrogenous ingre- dients of the urine has received by far the greatest attention of any individual factor. The collection and analysis of urine were comparatively simple matters, while the analyses of the respiratory products were in general precluded. Consequently we find in the literature of the subject a large number of deter- minations of nitrogen in fasting urines which are of especial interest in discussing the nitrogen elimination in the experiments here reported. Eecent investigations, notably those of Folin, have shown the importance of determining not only the total nitrogen but also the different nitrogenous ingredients, and of apportioning the nitrogen among these ingredients. In earlier experiments, however, the partition of the total nitrogen was not attempted and in the Middletown experiments here reported, it was impossible to determine the nitrogenous compounds directly, except in the instances where the determinations of creatine and creatinine, and a very few of uric acid, were made. 362 Influence of Inanition on Metabolism. Total Nitrogen. In many of the earlier experiments on fasting made before the development of the Kjeldahl method, urea, the chief nitrogenous product of the urine, was determined either by the Liebig method, or by the action of sodium hypobromite, and the results thus obtained were considered as the measure of the total nitrogen elimination, although it is now recognized that other nitro- genous ingredients which are not precipitated by mercury salts or decomposed by sodium hypobromite are present in all urines. Indeed even in the long experiment with Succi at Florence, the method for determining nitrogen was distinctly open to objection, as has been pointed out by Munk (7). Table 196. — Amounts of nitrogen eliminated in urine daily by fasting subjects. Day of fast. Succi. At Flor- ence. 63.3K. At Na- ples. 63.6K At Vien- na. J. A. 67. 8K. Cetti. 17. OK Breit- haupt. 60.1K. Lan- der- gren. 78.6K J.1 Sohn. 80. 2K. 58. 6K. Sub- ject Sub- ject II.2 Flora Tos-i ca. Kel- ler.3 74. 7K. Last food day. 1.. 2.. d . . 4. . 5.. 6.. 7.. 8.. 9.. 10.. Gms. *17.85 15.19 12.13 15.25 14.08 14.12 11.18 10.31 9.37 8.56 7.43 Gms. ^8.99 8.72 8.45 9.05 8.51 9.87 62 62 s-t 6.90 5.37 Gms. Gms 22 .41 17.00 12.04 11.2 j 12.72 10.55 13.48 10.8 13.56 ll.19lll.34 11.01 8.79 9.74 10.05 7.12 Gms. 13.49 13.55 12.59 13.12 12.39 10.70 10.10 10.89 8.90 10.83 9.47 Gms. 13.02 10.01 9.92 13.29:15.13 12.78:13.87 10.95J 9.88 Gms. \ Gms. 19.71 13.60ll2.44 13.43) 5.67 9.71 9.47 Gms. 20. 9S 12.37 12.37 14.01 .6 27.99 10.79; 14.50J 21.58 Gms. 7.15 8 10 11 Gms. 9.48 11.36 14.33 Gms. 13.99 8.761 8.38 10.73 9.40 7.87 7.73 6.11 7.70j 7.35: 6.80 Gms. 8.21 6.75 7.91 11.48 1 Reported by A. Sadovyen (2). 2 Reported by Schreiber & Waldvogel. 3 Reported by A. Keller, Zeit. f . physiol. Chemie (1900), 29, p. 165. 4 The figures in this column are given as corrected by Munk (7), p. 118. 5 Given by Ajello and Solaro (6) as urea and here converted to nitrogen for purposes of comparison. Since the authors do not give the method employed, no attempt is here made to correct the figures. 0 Amount for 2 days. The urea determinations in these earlier experiments are therefore of little value for comparison with results obtained by the Kjeldahl method, but they do serve to show the approximate relative amount of urea on the different days of the same experiment. Munk (7) has collected the references to the older literature of the excretion of nitrogen (usually expressed as urea) in the fasting insane, cases of oesophagus stricture, etc. The subjects were in nearly all cases of small body-weight and the nitrogen output was frequently as low as four grams. These figures are of value in indicating the possibilities of a minimum proteid katabolism, but the results of the determination of the nitrogen output of Nitrogen in Urine. 363 persons in normal health made with modern methods of analysis are of much greater interest in connection with a discussion of the results obtained in the Middletown experiments. The results of the more recent fasting experiments are given in table 196. Although the nitrogen excretion per kilo of body-weight is not given in the table, the data are readily obtained since the body-weight at the beginning of the fast *3 is given in the column headings. Eanke 44 made three experiments on himself while fasting. Each experiment lasted one day. The amounts of nitrogen 45 eliminated were 10.4, 8.0, and 8.6 grams, respectively. In the last experiment no water was taken. Pettenkof er and Voit 4B reported three experiments, each continuing one day in which the subject took no food other than a small amount of meat extract. The extract contained 1.2, 1.3, and 1.7 grams of nitrogen on the three days. The excretion of nitrogen in the urine was 12.5 (7.42 day, 5.09 night), 12.3 (6.72 day, 5.55 night) and 12.3 (5.55 day, 6.21 night) grams, respectively. From 10 to 15 grams of common salt were taken in connection with the meat extract. In the first two experiments the subject was resting, in the last he performed considerable muscular work. In both Eanke's and Pettenkofer and Yoit's experiments, the determination of nitrogen was made by the Liebig method and hence did not represent the total urinary nitrogen. It is a matter of historic interest that the first experiment made with Pettenkofer's respiration apparatus was that of Eanke studying the metabolism during inanition. Likhachev " in a one-day fasting experiment found the total nitrogen output to be 8. 5 2 grams. In a 2-day fast Eitter48 found that the subject excreted 11.68 and 12.96 grams of nitrogen, respectively. In fasting experiments made in this laboratory and reported elsewhere,49 the nitrogen excretion on the first day of fast in four experiments was 11.5, 16.0, 14.1, and 11.7 grams, respectively. The experiments were all made with the same subject. The first was after a carbohydrate diet during 4 days of rest; the second after a 4-day work experiment, in which the diet consisted largely of fats; the third likewise followed a 4-day work experiment with a fat diet; and the fourth was the first day of a 2-day fast experiment following a day of rest. On the second fasting day 12.2 grams of nitrogen were excreted. 43 In computing the nitrogen excreted per kilo of body-weight the average body- weight for each day of the fast should be used to secure the greatest accuracy. These weights may be obtained by means of the daily losses shown in tables 184 and 185. 44Archiv Anat. u. Physiol. (1862), p. 340. 45 The total nitrogen was computed from the weights of urea and uric acid found. 49Zeit. f. Biol. (1866), 2, p. 478. 47 Dissertation St. Petersburg (1893). ^Ritter, Miinchener medic. Wochenschr., 1893, Nos. 31 and 32. 49 U. S. Dept. of Agr., Office of Expt. Sta. Bui. 136, p. 120 (1903). 364 Influence of Inanition on Metabolism. Woods & Merrill s° report the average daily nitrogen excretion in a two- day fast as 11.96 grams. Eosemann " found 9.89 grams of nitrogen in the urine on the first day of a fast. In cooperation with Dr. A. E. Diefendorf the writer has presented * the results of a 6-day fast with an insane woman weighing 50 kilos. The daily nitrogen output was 4.19, 6.05, 6.38, 6.93, 6.16, and 4.41 grams. Brugsch and Hirsch f have reported the total nitrogen output of a profes- sional fasting woman weighing 56.3 kilos from the second to the sixteenth day of fasting. The output for the successive days of fasting beginning with the second was 8.41, 6.59, 7.78, 7.86, 7.82, 7.13, 6.20, 5.40, 4.38, 5.17, 5.38, 8.11, 5.96, 5.10, and 4.07 grams respectively. Dr. Otto Folin $ reports the total nitrogen output on six days with a fasting man weighing 50 kilos as 4.4, 6.5, 7.7, 10.8, 11.0, and 12.2 grams respectively. The total nitrogen excretion for each 24 hours of the fasting experiments here reported is given in table 197. For the details of the amounts of nitrogen excreted during the different periods of the day, reference is made to the tables in the statistical data for the experiments and to table 198. The results of the nitrogen determinations in the urine of the subjects in the experiments made in this laboratory here reported show that the nitrogen elim- inated on the first day of fasting ranges from 5.84 grams in experiment No. 71 to 13.25 grams, in experiment No. 83. The average for the first day for all the experiments is 10.03 grams. The unusually small amount of nitrogen eliminated in experiment No. 71 has been the subject of much investigation. An examination of the statistical tables shows that on that particular day, the nitrogen determinations by periods and in the composite sample agree as well as could possibly be expected. The weight of total solids observed on this day is lower than on any other day of fasting (see column g of table 193), and the weight of organic matter (see table 195) for this day is likewise much lower than on any other day. Hence there seems to be no doubt that there was an unusually low excretion of nitrogenous material in the urine on this day. It is also of interest to note in this connection that the largest amount of organic matter on the first day of the fasting experiments, aside from the imperfectly apportioned amount in experiment No. 59, is found on the first day of experiment No. 83, i. e., it corresponds with the largest excretion of nitrogen. Considering experiments with the same individual, S. A. B., in experiment No. 71, excreted the lowest amount (5.84 grams), of any of the subjects on the first day, and yet, on the first day of experiment No. 75, he excreted 12.24 grams, over twice as much as on the first day of experiment No. 71. 00 U. S. Dept. of Agr., Office of Expt. Sta. Bui. 85, p. 41 (1900). "Archiv f. die ges. Physiol. (1897), 65, p. 360. * Amer. Jour. Physiol. (1907), 18, p. 362. |Zeit. f. experimented Pathologie u. Therapie (1906), 3, p. 640. % Private communication. Nitrogen in Urine. 365 The smallest amount of nitrogen excreted on any day was 5.84 grams on the first day of experiment jSTo. 71. Table 197.— Nitrogen excreted in urine in metabolism experiments without food. Ex- peri- ment num- ber. Subject and duration of experiment. 59.. 68.. 69.. 71.. 73.. 75.. 77.. 79.. 80.. 81.. 82.. 83.. 85.. 89.. B. F. D., Dec. 18-20, 1903. . . . A. L.L., Apr. 27-28, 1904. ... A. L. L., Dec. 16-19, 1904. . . . S. A. B., Jan. 7-10, 1905 S. A. B., Jan. 28-Feb. 1, 1905 S. A. B., Mar. 4-10, 1905 S. A. B., Apr. 8-11, 1905 H.E.S.,Oct. 13-14, 1905... C. R.Y., Oct. 27-28, 1905. .. A. H. M., Nov. 21-22, 1905. . H. C. K., Nov. 24-25, 1905... H. R. D., Dec. 5-6, 1905 N.M. P., Dec. 9-10, 1905.... D.W.,Jan. 10-11, 1906 First day. Total. Per kilo- gram of body weight . Second day. Total. Grms. 11.84 12.26 10.09 5.84 10.29 12.24 8.81 8.11 7.78 9.11 9.38 13.25 KS2 11.05 9.99 Average . 10.03 Grms . 0.177 .170 .137 .101 .176 .206 .144 .143 .114 .148 .132 .239 .170 .128 Per kilo- gram of body wt. Third day. Total Per kilo- gram of body wt. Fourth day. Total. Grms. Grms. 14.110.213 13.03 14.26 11.04 11.97 12.45 10.78 14.35 9.95, 13.05 14.36 13.53 11.35 14.46 0.156 .184 .196 .194 .208 .211 .179 .259 .149 .216 .205 .248 .173 .188 Grms. 14.82 15.04 13.10 11.54 13.02 10.98 12.760.202 13.08 Grms. 0.226 .209 .232 .203 .223 .186 Grms. Per kilo- gram of body wt. Grms. 12.970.182 10.74 .193 10.39 .185 11.63 .202 11.45 .198 0.213 11.440.192 Experi- ment number. Subject and duration of experiment. Fifth day. Total. Per kilo- gram of body weight. Sixth day. Total. Per kilo- gram of body weight. Seventh day. Total. Per kilo- gram of body weight. 73.. 75.. S. A. B., Jan. 28-Feb. 1, 1905. S. A. B., Mar. 4-10, 1905 Grams. 9.98 10.87 Average 10 . 43 Grams. 0.180 .191 Grams. lb. n Grams. o!i9o 0.186 10.74 0.190 Grams. lb'.is 10.13 Grams. 0.'i8l 0.181 1 Determined for urine spilled. See p. 243. In the records of the nitrogen output during fast given in table 196, aside from the uncorrected low figures of the Naples fast of Succi, the lowest nitrogen excretion is 5.67 grams on the second day with " J." The observa- tions during the earlier days of fasting of so low an amount of nitrogen in the urine of the subjects " J " and " S. A. B." are strikingly exceptional. Even with the fasting girl, Flora Tosca, presumably of smaller body-weight, the nitrogen excretion is larger than in the two instances cited above. The largest amount of nitrogen excreted on any fasting day in the Middle- town experiments is 15.04 grams on the third day of experiment No. 69. 366 Influence of Inanition on Metabolism. While the data in table 196 show that in general the excretion of nitrogen is less as the fast progresses, the determination of 21.58 grams of nitrogen in the urine of Sohn, on the eighth day of fast, is remarkable as being the largest excretion of nitrogen thus far observed with any fasting man. It should further be noted, that inasmuch as Sohn was well nourished at the beginning of the fast and the experiment continued but eight days, we have not here to do with the premortal rise in nitrogen excretion occasionally observed in animals after a prolonged fast. Excretion of nitrogen on days preceding fast. — In discussing the nitrogen excretion during fasting, the excretion on the day preceding the fast is of value in showing the rate of elimination when the ingestion of food ceased, since a large nitrogen excretion on the day preceding the fast would indicate a large nitrogen consumption. In certain cases the subjects of fasting experiments have made a special effort to eat an unusual amount on the day before the fast as a preliminary to the fast. Thus, the hypnotic subject of Hoover & Sollman (8) excreted 20.98 grams of nitrogen on the day before the fast began, while on the first day without food the nitrogen excretion fell to 12.37 grams. The nitrogen excreted by Landergren was 19.71 grams on the day before the fast and 13.60 grams on the first day of the fast. Perhaps the most marked difference in the excretion of the nitrogen on the day before the fast and the first fasting day thus far recorded is that of J. A., in the Stockholm laboratory (9), the nitrogen on the first day of the fast, 12.04 grams, being but a little more than one-half that excreted on the day before the fast, namely, 22.41 grams. In nearly all of the Middletown experiments, the subjects were cautioned not to eat heartily during the day preliminary to a fast, and to eat no meat. It was possible in many cases to measure the nitrogen excretion on the day preceding the fasting period. For use in interpreting the results of the fasting days all the data available for the nitrogen determinations for the day before the fast and made in connection with these experiments are here given. In a number of instances, only the urine for the 12 hours before the fast began, was analyzed. In others, the analyses are made in the usual four periods for the day, the data being given in full below. An examination of the data obtained with the subject S. A. B. for the preliminary night or day, shows that on the day preceding the first fast day of experiment No. 71 the subject excreted 3.21 grams of nitrogen between 7 p. m. and 7 a. m., an unusually small excretion for this subject, as is seen by com- parison with the corresponding amounts of nitrogen eliminated during the night period on other preliminary days. The total nitrogen output for the day preceding experiment No. 73 was 12.50 grams, of which 7.153 grams was elimi- nated during the night from 7 p. m. to 7 a. m. On this preliminary night, Nitrogen in Urine. 167 therefore, the nitrogen excretion was over twice that of the night preceding experiment No. 71. The corresponding elimination on the first day of the fast is also twice as great as in experiment No. 71. On the day preceding experiment No. 75, S. A. B. excreted 19.495 grams of nitrogen, and the greatest elimination of nitrogen on the first day in his fasts occurred in this experiment. Preliminary No. 59. Dec. 17-18, 1903: Grams. 7 p. m. to 7 a. m 6.234 No. 68.1 Apr. 25-26, 1904: 7 a. m. to 1 p. m 4.332 1 p. m. 7 p. m 5.098 7 p. m. 11 p. m 3.302 11 p. m. 7 a. ni 5.720 Total for 24 hours. .18.452 Apr. 26-27, 1904 (2d day, No. 67) : 7 a. m. to 1 p. m 3.766 1 p. m. 7 p. m 4.714 7 p. m. 11 p. m 2.991 11 p. m. 7 a. m 4.949 Total for 24 hours. .16.420 No. 71. Jan. 6-7, 1905: 7 p. m. to 11 p. m 702 11 p. m. 7 a. m 2.508 No. 73. Jan. 27-28, 1905: 7 a. m. to 1 p. m 1.845 lp.m. 7 p. m 3.504 7 p. m. 11 p. m 2.949 11 p. m. 7 a. m 4.204 Total for 24 hours. .12.502 1 Experiment No. 68 followed immediately No. 6 Nitrogen. No. 75. Mar. 3-4, 1905: Grams. 7 a. m. to 7 p. m 11.239 7 p. m. 11 p. m 2.893 11 p. m. 7 a. m 5.363 Total for 24 hours. .19.495 No. 77. Apr. 7-8, 1905: 7 a. m. to 1 p. m. . . . 3.104 1 p. m. 7 a. m 8.259 Total for 24 hours. .11.363 No. 79. Oct. 12-13, 1905: 7 p. m. to 11 p. m 1.95 11 p. m. 8h33m a. m. 2.70 No. 80. Oct. 26-27, 1905: 7 a. m. to 7 a. m 7.78 No. 81. Nov. 20-21, 1905: 7 a. m. to 7 a. m 8.61 No. 82. Nov. 23-24, 1905: 7 a. m. to 7 a. m 11.20 No. 83. Dec. 4-5, 1905: 7 a. m. to 7 a. m 8.719 No. 85. Dec. 8-9, 1905: 7 a. m. to 7 a. m 16.597 No. 89. Jan. 9-10, 1906: 7 a. m. to 7 a. m 13.684 7, a rest experiment with food. Furthermore, on the day preceding experiment No. 77 the nitrogen elimi- nation was 11.363 grams, while the nitrogen excretion on the first day fell to 8.81 grams. So far, therefore, as the different experiments with the same subject are concerned there would appear to be a relationship existing between the nitrogen elimination of the day preliminary to the fast and the nitrogen secretion of the first day. When the series of 2-day fasting experiments for different individuals is considered the relationship is by no means as„ clear. The largest elimination on the day prior to the fast was that in experiment No. 85, 16.597 grams, and yet the nitrogen elimination on the first day of the fast was nearly 2 grams less than the maximum "first day" excretion which occurred in experiment No. 83. On the other hand, one of the lowest nitrogen excretions during the preliminary days was that on the day preceding the latter experiment. 368 Influence of Inanition on Metabolism. Unfortunately in the series of 2-day fasts, duplicate experiments were not made with the same subject, and hence it is not possible to substantiate for the same individual the wide variation between the amounts of nitrogen excreted on the preliminary day and the first day of the fast. It is furthermore much to be regretted that the data are also lacking for this comparison in the several fasts made by Succi. Prausnitz (5), in a critical discussion of the subject, reports two experiments made with the same subjects in which the food on the day before the fast contained a much larger amount of protein in one case than in the other. The effect of the ingestion of increased amounts of protein on the nitrogen excretion of the first fasting day is very evident. In fifteen 2-day fasting experiments on 12 different subjects, Prausnitz observed a nitrogen excretion on the first day of the fast varying from 4.6 " to 17.3 grams. Sadovyen (2) reports also a second experiment with "J" in which the subject fasted 2 days without drinking water. The nitrogen excretion in the urine was for the day before the fast, 20.842 grams; for the first day of fast, 12.6 grams; and for the second fasting day 11.55 grams. It is undoubtedly true that the nitrogen of protein ingested is not completely eliminated for some 36 hours, yet it is nevertheless a fact that a large excretion of nitrogen on the day preceding a fast may not necessarily be accompanied by a similar excretion of nitrogen on the first fasting day. Although the nitrogenous excretion in the urine of the first fasting day is influenced to a marked degree by the protein ingestion of the day prior to the fast, the amounts of nitrogen eliminated on the second day of the experi- ments (table 197), are much more nearly constant ranging from 9.95 grams in experiment No. 80 to 14.46 grams in experiment No. 89. The average for all the experiments is 12.76 grams. Comparison of experiments with the same subject (S. A. B.) shows that the nitrogen excreted in experiment No. 77 on the second day was 10.78 grams and in experiment No. 75, 12.45 grams. On the second day of the other two experiments (Nos. 71 and 73) with S. A. B., it was 11.04 and 11.97 grams, respectively. Thus, while marked variations in the total quantity of nitrogen eliminated on the first day of the fast are observed in experiments with different individuals and in experiments with the same individual, on the second day of the fast the maximum difference in the amount of nitrogen excreted by the same individual in four fasts was but 1.7 grams. Prausnitz (5) on the second day of fasting found in fifteen experiments with twelve subjects a nitrogen excretion of 13.0, 4.4, 10.6, 13.0, 11.0, 10.3, 12.5, 14.9, 13.8, 14.5, 12.6, 16.0, 14.9, 13.0, and 19.3 grams, an average of 12.9 "This unusually low nitrogen output, followed by a still lower amount (4.4 grams) on the second fasting day, has been the subject of special comment by Prausnitz. Nitrogen in Urine. 369 grams. In comparing Prausnitz's results with those obtained in this laboratory, it is important to note that while wide differences in size, weight, and dietetic habits existed in Prausnitz's subjects those used in the Middletown experiments were as a rule college students eating more nearly the same fare. The approaching constancy in the nitrogen elimination found on the second day of the fast with different individuals would incline one to the belief that on the third day the constancy would be even greater, yet an inspection of the results in table 197 shows that the variations ranged from 10.98 in experiment No. 77 to 15.04 in experiment No. 69. In the four experiments with S. A. B., the variations were from 10.98 to 13.10 grams, or a little over 2 grams. The average elimination for the six experiments was 13.08 grams. Five experiments were made in which the fast was continued for 4 or more days. In these the nitrogen excretion varied from 10.39 to 12.97 grams, the average elimination for the five experiments being 11.44 grams. In the four experiments with S. A. B., the excretion varied from 10.39 to 11.63 grams, a difference of 1.24 grams. Not only is the variation for all the experiments much less, therefore, than on the previous days, but in different experiments with the same subject, there is a tendency for the nitrogen excretion to approach constancy on the fourth clay. Two experiments, both with S. A. B., continued for 5 or more days. The nitrogen in these experiments amounted on the fifth day to 9.98 and 10.87 grams, averaging 10.43 grams. One experiment continued during the sixth and seventh days and on these days there were excreted 10.74 and 10.13 grams of nitrogen, respectively. Variations in nitrogen elimination as the fast progresses. — While as has been seen from the previous discussion, the gross fluctuations in the nitrogen excre- tion for the first day of different fasts render any interpretation of these results for short fasting experiments difficult, in the longer fasts the disturbing features of the first day are eliminated and the data are such as to permit of definite conclusions. In the majority of the experiments, there is an excretion of at least 2 grams more on the second day than on the first. In experiment No. 85, there was a greater excretion on the first day by 0.02 gram which, however, may well be within the limit of experimental error. On the contrary, Prausnitz (5) found in three out of fifteen cases a dimin- ished excretion on the second day. Of the thirteen fasting experiments cited in table 196 there was an increased nitrogen elimination on the second day of fasting in only three instances. The lack of harmony between these results and those obtained in the Middletown experiments can not readily be explained. Physical condition, previous dietetic habits, muscular exercise and other factors, yet too little known, may all have contributed to this result. The great uniformity in the results of the Middletown experiments would tend to uphold the view, that with like bodily activity and environment, the results should all be strictly comparable. 24 370 Influence of Inanition on Metabolism. For the third day of each fast (table 197) the data show that in all cases (except experiment No. 73, where there was a diminished excretion of 0.43 gram), the nitrogen is greater on the third day than on the second, thus indicating an average progressive increase in the nitrogen excretion. For the average of all the experiments, the daily excretion is 10.03, 12.76, and 13.08 grams on the first, second, and third days, respectively. The evidence is sufficient to show a positive increase in nitrogen on the third day over the second amounting to 0.32 gram. This is also observed on the third day of the earlier fasts (table 196) with the single exception of the Vienna fast by Succi. Of the five experiments which continued for 4 days or more, 4 were made with one subject, and with the exception of experiment No. 77, there was a decrease in the total nitrogen excretion on the fourth day as compared with the third. The average excretion for the five experiments was 11.44 grams, or 1.64 grams less than on the third day. There was a further decrease of 1 gram in the nitrogen excreted on the fifth day while on the sixth and seventh days, it remained about the same. In general, then, according to our experiments, the average nitrogen excretion on the second and third days is larger than on the first day, the maximum being reached on the third day. On the fourth and fifth days there is a steady diminution, and so far as the few observations recorded show, the excretion for the fifth, sixth, and seventh days is practically constant. Relation of body-weight to nitrogen excretion. — Since proteid katabolism is proportional in general to the size of the body and in all probability to the active mass of protoplasmic tissue, it is important to note the nitrogen excretion per kilo of body-weight in the different fasting experiments. These have been also recorded in table 197. On the first day of the fast, the variation in the nitrogen excreted per kilo of body-weight is from 0.101 gram in experiment No. 71 to 0.239 gram in experiment No. 83. The low excretion per kilo of body-weight in experiment No. 71 is to be expected from the unusually small amount of nitrogen excreted on this day. It is important to note that in experiment No. 75, made with the same subject, the amount of nitrogen excreted per kilo of body-weight is 0.206 gram. The average excretion for the first day of the fast is 0.156 gram. On the second day the excretion per kilo varies from 0.149 gram in experi- ment No. 80 to 0.259 gram in experiment No. 79, averaging 0.202 gram. With the same individual, S. A. B., the variations ranged from 0.179 gram in experi- ment No. 77 to 0.211 gram in experiment No. 75. The excretion, then, per kilo of body-weight on the second day of the experiments is much more nearly constant than on the first day. On the third day, the per kilo excretion varies from 0.186 gram in experiment No. 77 to 0.232 gram in experiment No. 71, the average for the six experiments being 0.213 gram. The variations on the fourth day of the fast are even smaller than on the third, ranging from 0.182 gram in experiment No. 69 to 0.202 gram in Nitrogen in Urine. 371 experiment No. 75, the average for the 5 experiments being 0.192 gram. On the fifth, sixth, and seventh clays, the average excretion per kilo of body- weight is 0.186, 0.190, and 0.181 gram, respectively. The same differences that exist in the total excretion of nitrogen in the different experiments on the first day of the fast likewise appear in the excretion per kilo of body-weight. On the second and succeeding days, the excretion per kilo of body-weight is on an average much more nearly constant, and the maximum is reached on the third day. In discussing the possible causes of variations in the elimination of nitrogen with different subjects, and indeed, in different experiments with the same subject, certain questions other than that as to the nitrogenous excretion on the day before the fast may be raised. For example, does the relative amount of protein in the body influence the rapidity with which it is disintegrated during the fast? In discussing the experiments on Cetti and Breithaupt, the Berlin investigators were of the opinion that the large increase in the elimination of nitrogen by these two subjects was due to the fact that they were not very fat and hence had a larger proportion of protein in the body. With the subject S. A. B., considerable light may be thrown upon the storage of protein inasmuch as between experiments Nos. 76 and 77 and also for a period of two weeks after experiment No. 77, a careful record was kept of all food ingested and the nitrogen was determined for each day's food, urine, and feces thus furnishing a complete balance of income and outgo of nitrogen. The details of these balances are reported beyond (see page 534), but it may be stated here that the results show that during the period between experiments Nos. 76 and 77, the subject actually gained 45.0 grams of nitrogen more than were lost during fasting experiment No. 75 and food experiment No. 76, thus indicating a marked storage of nitrogen in the body. In the period following experiment No. 77 the data show that the body gained in addition to the nitrogen lost during the fast 11.1 grams of nitrogen. Under these conditions, the increased amount of nitrogen is very noticeable and hence we should expect to find, that if the actual amount of nitrogenous material present in the body, be it either " organized " or " circulating," at the beginning of the experiment influences the subsequent nitrogen elimination during fasting, there would be an increase in the nitrogen elimination in the last experiment with this subject (experiment No. 77) over that excreted in the earlier ones. As a matter of fact, aside from the unusually low excretion noted on the first day of experiment No. 71, the absolute excretion, as well as the excretion per kilo of body-weight, was lowest rather than highest in experiment No. 77. Unquestionably the gain in body-weight after fasting was due in large measure to the replacement of the water lost, but there must have been a not inconsiderable gain of fat, since the subject weighed considerably more at the end of the series of experiments than at the beginning. It may then be true 372 Influence of Inanition on Metabolism. that the proportion of protein, when compared to that of fat, may have been no greater in the later than in the earlier experiments. The increase in water content of the body, however, may well lower the proportion of total protein in the body even with the material gain observed in the long nitrogen metabolism experiments. Although the evidence is insufficient for drawing any definite conclusions, the results obtained imply that after the first fasting day the supply of reserve protein in the body has little, if any, effect on the subsequent excretion of nitrogen. Further experiments in which the partition of the nitrogen is more carefully studied are needed.1 Excretion of nitrogen in different periods of the day. — The samples of urine for the four periods of each day were separately analyzed during a portion of the series of experiments in the effort to obtain information concerning the distribution of nitrogen excretion. These data are presented in table 198, in which the excretion for the two 6-hour day periods and the 12-hour night period are given. In this table the results for the periods from 7 p. m. to 11 p. m., and 11 p. m. to 7 a. m. are combined. The proportions of the total nitrogen excreted during the day period and the night period have likewise been computed and the percentages expressed in the table. The data in the table, which includes all the fasting experiments in which determinations by periods were made, show that during the period from 7 a. m. to 7 p. m. the per cent of total nitrogen may vary from 47.5 per cent, as on the second day of experiment No. 89, to 60.5 per cent, as on the first day of experiment No. 75. On the average 51.9 per cent of the total nitrogen is excreted during the day period. There is a general tendency for the per cent of nitrogen excreted during the day period to increase perceptibly as the fast progresses in experiments Nos. 69 and 73. On the contrary, in experiment No. 75, the highest percentage is on the first, the lowest on the third day. The proportions of nitrogen eliminated during the periods from 7 a. m. to 1 p. m. and 1 p. m. to 7 p. m. have not been shown separately in the table, but in general the larger excretion takes place during the period from 7 a. m. to 1 p. m., although the average for all the experiments is only about 2 per cent larger. Thus 51 per cent of the total nitrogen excreted during the first 12 hours of the day appears in the urine of the six hours from 7 a. m. to 1 p. m. A few noticeable exceptions are observed especially in the second day of experi- ment No. 59, both days of experiment No. 68 and in the second day of experiment No. 89. 1 Folin's result on a fasting (see p. 364) man are of interest in that the subject had subsisted on a low nitrogen diet for some days before the fast began. Nitrogen in Urine. 373 Table 198. — Periodic distribution of nitrogen in urine in metabolism experiments without food. Ex- peri- ment num- ber. Subject and date. 7 a. m. to 1 p. m. 1 p. m. to 7 p.m. Total for first 12 hours. Total for last 12 hours. Amt. Pro- portion of total for 24 hours. Amt. Pro- portion of total for 24 hours. 59 68 69 71 73 75 B.F.D. Dec. 18-19, 1903 Grams. 3.59 3.37 4.37 Grams. 3.22 4.19 4.17 Grams. 6.81 7.56 8.54 Per ct. 57 5 53.6 57.6 Grams. 5.03 6.55 6.28 Per ct. 42.5 46.4 42.3 Dec. 19-20, 1903 Dec. 20-21, 1903 Total, 3 days 11.33 3.78 11.58 3.86 22.91 7.64 56.2 17.86 5.95 43! 8 A. L. L. Apr. 27-28, 1904 2.89 3.08 3.31 3.47 6.20 6.55 50.6 50.3 6.06 6.48 49.4 49.7 Apr. 28-29, 1904 Total, 2 days 5.97 2.99 6.78 3.39 12.75 6.38 .... 12.54 50.4 i 6.27 4^96 Average per day A. L. L. Dec. 16-17, 1904 2.45 3.94 4.32 4.12 2.51 3.80 4.03 3.47 4.96 7.74 8.35 7.59 49.2 54.3 55.5 58.5 5.13 6.52 6.69 5.38 50.8 45.7 44.5 41.5 Dec. 17-18, 1904 Dec. 18-19, 1904 Dec. 19-20, 1904 Total, 4 days 14.83 3.71 13.81 3.45 28.64 7.16 54.7 23.72 5.93 45.3 S. A. B. Jan. 7- 8, 1905 2.74 3.47 2.85 3.29 5.78 6.93 6.21 56.3 '< 2.55 52.4 5.26 52.9 6.17 57.8 i 4.53 43.7 47.6 47.1 42.2 Jan. 8- 9, 1905 3.04 3.46 3.36 Jan. 9-10, 1905 Jan. 10-11, 1905 Total, 4 days 9.86 J3.29 9.06 x3.02 22.21 5.55 .... 18.51 54.5 4.63 45!5 Average per day S.A.B. Jan. 28-29, 1905 2.58 3.13 3.18 3.10 3.14 2.56 3.23 2.72 2.92 2.61 5.14 6.36 5.90 6.02 5.75 50.0 53.1 51.1 57.9 57.6 5.15 5.61 5.64 4.37 4.23 50.0 46.9 48.9 42.1 42.4 Jan. 29-30, 1905 Jan. 30-31, 1905 Jan. 31-Feb. 1, 1905 Feb. 1-2, 1905 Total, 5 days 15 13 3.03 14.04 2.81 29.17 5.83 53^9 25.00 5.00 46 .1 Average per day S.A.B. Mar. 4- 5, 1905 4.10 3.82 3.55 3.36 3.19 3.26 2.98 3.30 3.30 3.73 3.39 3.13 2.82 2.83 7.40 60.5 7.12 57.2 4.84 5.33 5.74 4.88 4.55 4.66 4.32 39.5 42.8 44.1 42.0 41.9 43.4 42.6 Mar. 5- 6,1905 Mar. 6- 7, 1905 7.28 6.75 6.32 6.08 5.81 55.9 58.0 58.1 56.6 57.4 Mar. 7- 8, 1905 Mar. 8- 9, 1905 Mar. 9-10, 1905 Mar. 10-11, 1905 Total, 7 days 24.26 3.47 22.50 3.21 46.76 6.68 57.7 34.32 4.90 42.3 Average per day 1 Average for 3 days. 374 Influence of Inanition on Metabolism. Table 198. — Periodic distribution of nitrogen in urine in metabolism experiments icithout food — Continued. Ex- peri- ment num- ber. Subject and date. 7 a. m. to 1 p. m. 1 p.m. to 7 p. m. Total for first Total for last 12 hours. 12 hours. Am't. Pro- portion of total Am't. for 24 hours. Pro- portion of total for 24 hours. 89 D.W. Jan. 10-11, 1906 Grams. 2.82 3.18 Grams. 2.73 3.69 Grams. 5.55 6.87 Per cfc. 55.6 47.5 Grams. 4.44 7.59 Per ct. 44.4 52.5 Jan. 11-12, 1906 Total, 2 days 6.00 3.00 6.42 3.21 12.42 6.21 50^8 12.03 6.02 49^2 Average of above experiments l3.36 J3.24 1 fi.48 54.9 5.33 45.1 1 Average for 26 days. Data regarding the periodic excretion of nitrogen during fasting are also furnished in the experiments made in this laboratory and reported elsewhere." In the 5 clays during which the subject fasted, three 1-day experiments and one 2-day experiment, the proportion of total nitrogen excreted between 7 a. m. and 7 p. m. was 58.6, 52.7, 56.6 per cent for the three 1-day experiments, and 53.8 and 45.7 per cent for the 2 consecutive days of fasting. The Swedish investigators in reporting the results for J. A. (9) give data for computing the proportion of total nitrogen excreted between 10 a. m. and 10 p. m. for the 5 days of the fast, The percentages are 59.1, 51.0, 50.7, 58.3, and 56.1, respectively. In a one-day fast Eosemann54 found 59.8 per cent of the total nitrogen was excreted between 7 a. m. and 7 p. m. Influence of drinking-water on nitrogen excretion. — It has long been observed that the ingestion of large amounts of drinking-water results frequently in an increased nitrogen excretion. The significance of the relation between the volume of urine and the nitrogen excreted during fasting was pointed out by the Berlin scientists (7) in their discussion of the experiments on Cetti and Breithaupt. The influence of excessive amounts of drinking-water on fasting animals has also repeatedly been observed, and hence it is of importance to consider the volume of urine in the fasting experiments here reported, since the variations in volume may have an influence upon the excretion of nitrogen, and thus possibly account for the fluctuations in nitrogen excretion observed from day to day. The daily nitrogen excretion in grams and also the volume of urine are recorded in table 200, beyond. SSU. S. Dept. Agr., Office of Expt. Sta. Bui. 136 (1903). "Archiv f. die ges. Physiol. (1897), 65, p. 360. Nitrogen in Urine. 375 A comparison of the data for the total 24 hours of a given experiment shows that in general large volumes of urine are accompanied by large amounts of nitrogen. For the consecutive days of each experiment this is generally true but no comparison is permitted for different experiments. This general- ization is true for the large majority of the fasts. Especially is it true in the first 3 days of experiment No. 69, and the whole of experiments Nos. 71, 73, 75, 81, and 82. On the other hand, there are striking contradictions noticed more especially in experiment No. 77. In this experiment the largest volume of urine appears on the first day and is coincident with the smallest excretion of nitrogen, while on the last day the smallest volume of urine is accompanied by the largest excretion of nitrogen. A similar statement holds true for experiment No. 59. (See table 3.) In considering this question, it is doubtful whether the 2-day experiments, influenced as they undoubtedly are by the food of the day before, can be considered as of much value. It thus appears that, while in the majority of the experiments it may be reasonably inferred that there was an increased elimi- nation of nitrogen coincident with large volumes of urine, on the other hand, the data of experiments Nos. 59 and 77 render the conclusion doubtful. The influence of drinking large quantities of water on the nitrogen excretion has been studied very carefully, the majority of the experiments, however, having been made with animals. All of the observations have demonstrated that with increased volume of urine there is an increased nitrogen excretion. The results have, however, permitted of two interpretations of the cause of the increase, and scientists have differed in their belief. According to one view the increase is due simply to the flushing out of the urea and other products preformed in the tissues, while the other maintains that the drinking of large amounts of water results in a stimulated protein katabolism. The more recent investigations of Hawk," and Heilner,56 have considered this problem in a more elaborate manner, stating not only the excretion of nitrogen but also of phosphorus, sulphur and chlorides. An examination of the more recent experimental evidence leads to the conviction that experiments on fasting men especially with complete urine analyses would aid materially in settling this much disputed point. In spite of the fact that in several of the experiments here reported unusually large amounts of drinking-water were consumed, the conditions under which these experiments were made are such as in our judgment to preclude any direct conclusions regarding the influence of increased consumption of water on nitrogen excretion. It is important, however, in considering the causes for the variations in the nitrogen excretion on the different days, not to overlook the fact of the possible influence of the varying amounts of water consumed. 58 Univ. of Pennsylvania Medical Bulletin (1904). 56Zeit. f. Biol. (1906), 47, p. 538. 376 Influence of Inanition on Metabolism. In all the experiments with S. A. B., the volume of urine was large, in many cases very large. It should be noted, however, that the highest nitrogen excretion by this subject on any day was 13.10 grams, while in experiment No. 69 with A. L. L., an excretion of 15.04 grams of nitrogen appeared on the third day with but 743 cc. of urine. It would appear, then, that no very marked washing out of the urea or, indeed, increased protein katabolism followed the ingestion of the excessive quantities of water consumed by S. A. B. In the experiments with this latter subject the volume of urine was less than 1200 cc. on only three days. On the first day of experiment No. 71, the volume was 1149 cc. with a nitrogen excretion of 5.84 grams; on the last day of experiment No. 73, the volume was 1026 cc. with a nitrogen excretion of 9.98 grams; and on the last day of experiment No. 77, the volume was 1136 cc, and the nitrogen excretion was 11.45 grams. The large amounts of drinking-water consumed by the subject S. A. B. were as a matter of fact not much, if any, greater than his customary consumption. While the data for the amount of drinking-water on the preliminary days is missing, the volumes of urine are recorded in the following table. For com- parison the volumes on the first days of the experiments are included, as well as the nitrogen excretion for the corresponding days. Experiment number. Preliminary. First day. Volume. Nitrogen. Volume. Nitrogen. 71 73 75 77.. 1482 2265 1135 1111 13.21 12.50 19.49 11.36 1149 2252 1496 2552 5.84 10.29 12.24 8.81 1 For 7 p. m. to 7 a. m. only. The corresponding quantities for the same period on the first day of experiment 71 are : Volume = 170 cc ; nitrogen = 2.55 grams. The only noticeable increase in the volume of urine on the first day of fast is observed in experiment No. 77, in which nearly two and one-half times the volume of urine of the preliminary day was excreted. In spite of the influence of the large amount of water on this day the nitrogen excretion can in no wise be construed as indicating a flushing out of the nitrogenous wastes in the tissues. A more detailed statement of the elimination of nitrogen by different periods in which the volume of urine as well as the total weight of nitrogen are taken into consideration is given in table 199. While from the total volume of urine in 24 hours, it would appear that the nitrogen excretion followed roughly the volume of urine, when the volumes and excretion by periods are considered, the relationship is by no means as clear. Thus in experiment No. 73, first 6-hour period of the first day, there were passed 871 cc. of urine with but 2.58 grams of nitrogen. During the same period on the last day of the same experiment there was a little over half this Nitrogen in Urine. 377 Table 199. — Volume of urine excreted and nitrogen elimination by periods in metabolism experiments without food. Ex- peri- ment num- ber. Subject and date. 59 68 69 71 73 75 B.F.D., Dec. 18-19, 1903 Dec. 19-20, 1903 Dec. 20-21, 1903 Total, 3 days Average per day A.L.L., Apr. 27-28, 1904 Apr. 28-29, 1904 Total, 2 days Average per day A.L.L., Dec. 16-17, 1904 Dec. 17-18, 1904 Dec. 18-19, 1904 Dec. 19-20, 1904 Total, 4 days Average per day S.A.B., Jan. 7- 8, 1905, Jan. 8- 9, 1905 Jan. 9-10,1905, Jan. 10-11, 1905, Total, 4 days Average per day .... S.A.B., Jan. 28-29, 1905. Jan. 29-30, 1905 Jan. 30-31, 1905. Jan. 30-Feb. 1,1905 Feb. 1-2, 1905 Total, 5 days. . . . Average per day . 89 S.A.B., Mar. Mar. Mar. Mar. Mar. Mar. 4- 5, 1905 5- 6, 1905 6- 7, 1905 7- 8, 1905 8- 9, 1905 9-10, 1905 Mar. 10-11, 1905 Total, 7 days Average per day. . . D.W., Jan. 10-11,1906. Jan. 11-12, 1906. Total, 2 days Average per day. . . . Average of all above ex- periments 7 a.m. to 1 p.m. 1 to 7 p.m. Vol- ume of urine. Nitro- gen. c.c. 434 188 252 874 291 Volume NHro. urine. Grms. c.c. 3.59 652 3.37 4.37 11.33 3.78 284 298 1234 411 gen. Grms 3.22 4.19 4.17 11.58 3.86 First 12 hrs. Vol- ume of urine. c.c. 1086 472 550 2108 703 Nitro- gen. Last 12 hrs. Volume of urine. Nitro- gen. Grms. 6.81 7.56 8.54 22.91 7.64 c.c. 431 490 358 Grms. 5.03 6.55 6.28 1279 426 17.86 5.95 262 217 479 24C 2.89 3.08 259 217 5.97 2.99 476 238 3.31 3.47 6.78 3.39 521 434 6.20 6.55 955 12 . 75 478 6.38 458 388 846 423 6.06 6.48 12.54 6.27 112 2.45 119 2.51 231 4.96 206 169 3.94 163 3.80 332 7.74 240 250 4.32 203 4.03 453 8.35 290 245 4.12 223 3.47 468 7.59 257 776 14.83 708 13.81 1484 28.64 993 194 3.71 177 3.45 371 7.16 248 5.13 6.52 6.69 5.38 23.72 5.93 ■ • * . 979 3.29 170 827 3.04 547 2.74 1374 5.78 635 750 3.46 605 3.47 1355 6.93 1174 640 3.36 519 2.85 1159 6.21 314 J2217 '9.86 U671 l9.06 4867 22.21 2293 1 739 *3.29 1 557 '3.02 1217 5.55 573 2.55 5.26 6.17 4.53 18.51 4.63 871 2.58 549 2.56 1420 5.14 832 815 3.13 937 3.23 1752 6.36 1206 633 3.18 888 2.72 1521 5.90 1204 605 3.10 765 2.92 1370 6.02 583 451 3.14 241 2.61 692 5.75 334 3375 15.13 3380 14.04 6755 29.17 4159 675 3.03 676 2.81 1351 5.83 832 5.15 5.61 5.64 4.37 4.23 25.00 5.00 407 774 500 605 310 448 389 3433 490 4.10 3.82 3.55 3.36 3.19 3.26 2.98 24.26 3.47 464 451 807 818 568 452 550 4110 587 3.30 3.30 3.73 3.39 3.13 2.82 2.83 22.50 3.21 871 1225 1307 1423 878 900 939 7.40 7.12 7.28 6.75 6.32 6.08 5.81 7543 46 . 76 1078 6.68 625 646 901 563 545 711 645 4636 662 4.84 5.33 5.74 4.88 4.55 4.66 4.32 34.32 4.90 219 123 342 171 2.82 3.18 6.00 3.00 195 138 333 167 2.73 3.69 6.42 3.21 414 261 675 338 5.55 6.87 12.42 6.21 212 261 473 237 4.44 7.59 12.03 6.02 442 23.362 458 2 3. 24 903 6.48 544 5.33 For 3 days. 2 For 26 days. 378 Influence of Inanition ox Metabolism. volume of urine eliminated, but with a very considerable increase in the nitrogen. When the total quantity of urine for the first 12 hours of the first and last days of this experiment are taken into consideration a similar peculiarity may be observed. On the first and fourth days of experiment No. 75, in the first 12 hours there was a marked increase in the volume of urine accompanied by an actual decrease in the elimination of nitrogen. Similar anomalies are to be found in all of the experiments. It is clear, therefore, that the deduction that large volumes of urine induced by copious water drinking result in an invariable increase in the quantities of nitrogen excreted can not be made from the results of the Middletown experiments. It may be said, moreover, that the other conditions surrounding the experiment, namely, state of nutrition, body-weight, etc., vary too much in the different experiments to permit the deduction that copious water drinking does not affect the excretion of nitrogen. Comparing the excretion of the day and the night periods in all the experi- ments given in table 199, there was on the average a greater excretion of nitrogen during the day than during the night. Considering the individual days, there were only two exceptions to this, namely, on the first day of experi- ment No. 69 and on the last day of experiment No. 89, on both of which days the excretion was somewhat greater during the night period. In one case the excretions for the day and for the night were identical. It is furthermore true that the volume of urine was, in general, greater during the day than during the night, there being but one exception to this, namely, on the second day of experiment No. 59. In general, the larger amount of nitrogen excreted during the day period from 7 a. m. to 7 p. m. appears in the urine from 7 a. m. to 1 p. m., and an inspection of the volumes of urine shows that in experiments Nos. 73 and 75, the larger volume of urine is collected as a rule in the second period (1 p. m. to 7 p. m.). This is likewise true for experiment No. 59. In each of the experiments Nos. 68, 69, and 89, the volumes average about the same for both 6-hour periods of the day. Only in experiment No. 71 do we find a noticeably larger volume of urine during the first 6 hours. It is interesting to note that this difference in the volume of urine is accompanied by a larger increase in nitrogen during the first 6-hour period. Ratio of Total Solids to Nitrogex. The nitrogenous ingredients of urine are of organic origin and hence the larger the proportion of ash in the urine, the wider the variations in the ratio of total solids to nitrogen. The ratios, however, are of particular interest as use was made of certain of them in experiments Nos. 59 and 68 in apportioning the amounts of total ash and total solids over the several days of the experiments as explained above. The ratio of total solids to nitrogen or the number of grams of total solids present in the urine for each gram of nitrogen is shown in table 200. Nitrogen ix Urine. 379 Table 200. — Ratio of total solids and organic matter to nitrogen in urine in metabolism experiments with and without food. Ex- peri- ment num- ber. Subject and date. (a) Vol- ume. (6) Nitro- gen. (c) Total solids. (d) Ratio of total solids (e) ch- Ratio of organic matter 'tonitro-^^ to nitro- gen (c-f-ft). gen (e-*-6). 59 68 69 71 73 75 Experiments without food. B. F. D., Dec. 18-21, 1903, aver- age per day l A. L. L., Apr. 27-29, 1904, average per day1 A. L. L., Dec. 16-17, 1904 Dec. 17-18,1904 Dec. 18-19, 1904 Dec. 19-20, 1904 c.c. \Grams. 1129 13.59 Grams. 52.93 Total, 4 days. . . . Average per day . S.A. B., Jan. 7- 8,1905. Jan. 8- 9,1905. Jan. 9-10,1905. Jan. 10-11, 1905. Total, 4 days. . . . Average per day . S.A.B. Jan. 28-29, 1905 Jan. 29-30, 1905 Jan. 30-31, 1905 Jan. 31-Feb. 1,1905. Feb. 1-2, 1905 Total, 5 days. . . . Average per day . 77 79 S. A. B., Mar. Mar. Mar. Mar. Mar. Mar. 4- 5, 1905, 5- 6, 1905. 6- 7, 1905. 7- 8, 1905. 8- 9, 1905. 9-10, 1905. Mar. 10-11, 1905. Total, 7 days Average per day . S.A. B., Apr. 8- 9,1905. Apr. 9-10, 1905. Apr. 10-11, 1905. Apr. 11-12, 1905. Total, 4 days Average per day . . . H. E. S., Oct. 13-14, 1905. Oct. 14-15, 1905. Total, 2 days Average per day . 3.895 901 12.65 46.70 3.692 35.45 2.802 Grams. 45.00 3.311 437 572 743 725 2477 619 10.09 14.26 15.04 12.97 52.36 13.09 36.05 45.02 47.49 41.78 170.34 42.59 3.573 3.157 3.158 3.221 3.254 29.41 38.59 40.93 36.21 145.14 36.29 2.915 2.706 2.721 2.792 2.772 1149| 5.84 2009 11.04 2529 13.10 1473 10.74 7160 1790 40.72 10.18 25.51 36.99 40.92 36.55 139.97 34.99 4.368 3.351 3.124 3.403 3.437 17.86 30.93 35.07 30.61 114.47 28 62 3.058 2.802 2.677 2.850 2.811 2252 2958 2725 1953 1026| 10.29 11.97 11.54 10.39 9.98 10914 2183 54.17 10.83 38.49 38.57 36.93 35.75 35.91 185.65 37.13 3.741 3.222 3.200 3.441 3.598 3.428 28.53 32.34 31.19 29.66 29.89 151.61 30.32 2.773 2.702 2.703 2.855 2.995 2.800 1496 12 1871 12 2208, 13' 1986 11 1423 1611 1584 12179 1740 10 10 10 .24 .45 .02 .63 .87 .74 .13 38.61 45.82 45.31 46.80 45.86 43.70 43.42 81.08 309.52 11.581 44.22 3.154 680 480 024 219 069 4.286 3.819 32.58 38.28 38.87 39.00 38.38 37.85 38.15 263.11 37.59 2.662 3.075 2.985 3.353 3.531 3.524 3.766 3.246 2552 8.81 2156 10.78 1554 10.98 1136 11.45 7398 1850 42.02 10.51 42.15 54.66 54.00 54.59 205.40 51.35 4.784 5.071 4.918 4.768 4.886 28.27 3.209 42.68 3.959 43.45 3.957 45.45 3.969 159.85: 39.96 3.802 1015 8.11 31.12 840 14.35 49.14 3.837 3.424 1855 22.46 80.26 928 11.23 40.13 3.573 1 Not determined for individual days. 3S0 Influence of Inanition on Metabolism. Table 200. — Ratio of total solids and organic matter to nitrogen in urine in metabolism experiments with and without food — Continued. Ex- peri- ment num- ber. 80 81 82 83 85 89 70 72 74 Subject and date. Experiments without food (cont'd). C. R. Y., Oct. 27-28, 1905 Oct. 28-29, 1905 Total, 2 days. . . . Average per day. A. H. M., Nov. 21-22, 1905. Nov. 22-23, 1905. Total, 2 days Average per day .... H. C. K., Nov. 24-25, 1905. Nov. 25-26, 1905. Total, 2 days Average per day . . H.R.D., Dec. 5-6,1905 Dec. 6-7, 1905. Total, 2 days Average per day . N.M.P., Dec. 9-10,1905 Dec. 10-11,1905. Total, 2 days Average per day .... D.W., Jan. 10-11,1906. .. Jan. 11-12, 1906. .. Total, 2 days Average per day Average of experiments without food Experiments with food. A. L. L., Dec. 20-2 1,1904.... Dec. 21-22, 1904.... Dec. 22-23, 1904.... Total, 3 days Average per day . S.A. B.; Jan. 11-12, 1905. S. A. B., Feb. 2-3, 1905. . . Feb. 3-4, 1905... Feb. 4-5, 1905. . . Total, 3 days. . . . Average per day . (a) Vol- ume. c.c. 1128 759 1887 944 (&) Nitro- gen. Grams. 7.78 9.95 17.73 8.87 (c) Total solids. Grams. 41.65 42.60 84.25 42.12 (d) Katioof total solids to nitro- gen (c-s-fe). (e) Or- ganic matter. If) Ratio of organic matter to nitro- gen (6-S-W. 5.353 4.281 I Grams. 22.72 30.08 4.749 52.80 26.40 2.920 3.023 2.976 643 812 9.11 13.05 1455' 22.16 728 11.08 38.74 48.04 S6.78 43.39 4.252 3.681 3.916 26.13 36.82 62.95 31.48 2.868 2.821 2.841 561 17721 9.38 14.36 37.13 54.93 3.958 3.825 27.46 38.60 2.928 2.688 23331 23.74 1167 11.87 92.06 46.03 3.878 66.06 33.03 2.783 1187 1046 2233 1117 13.25 13.53 26.78 13.39 45.29 52.61 97.90 48.95 3.418 3.888 3.656 37.10 43.88 80.98 40.49 2.800 3.243 3.024 1170 666 1836 918 1.32 .11.05 11.35 22.72 11.36 21.25\| 42.89J | 36.96 3.882 3.256 r 3.89\ 130.42/ 31.39 81.10 40.55 3.570 62.70 31.35 2.754 2.766 2 '760 626 522 1148 574 9.99 44.85 14.46 47.38 24.45 12.23 92.23 46.12 4.489 3.277 3.771 31.51 39.93 3.154 2.761 71.44 35.72 2.921 1350 11.54 43.67 3.784 34.82 3.017 1014 1026 608 2648 883 13.04 9.84 10.15 33.03 11.01 40.13 35.66 38.54 114.33 38.11 3.077 3.624 3.797 3.461 35.59 29.30 29.61 94.50 31.50 2.729 2.978 2.917 2.861 14S7 10.66 33.07 3.102 29.48 2.765 1509 10.74 1S86 8.25 1597 6.78 32.96 28.23 25.52 3.069 3.422 3.764 29.47 24.06 20.70 4992 25.77 1664 8.59 86.71 28.90 3.364 74.23 24.74 2.744 2.916 3.053 2.SS0 1 Determined for urine lost. See p. 243 -Calculated. See p. 243. Nitrogen in Urine. 381 Table 200.— Ratio of total solids and organic matter to nitrogen in urine in metabolism experiments icith and icithout food — Continued. Ex- peri- ment num- ber. Subject and date. (a) Vol- ume. (b) Nitro- gen. (O Total soiids. (d) Batio of total solids to nitro- gen (e+6). (e) Or- ganic matter. Ratio of organic matter to nitro- gen (e + b). 76 Experiments with food {cont'd). S. A. B., Mar. 11-12, 1905 c. c. 1725 Grams. 10.17 7.15 7.82 Grams. 40.62 23.25 27.09 3.994 3.252 3.464 Grams. 38.88 21.41 23.40 3.823 2.994 2.992 Mar. 12-13, 1905 1308 Mar. 13-14, 1905 1596 Total, 3 days 4629 1543 25.14 8.38 90.96 30.32 3.618 83.69 27.90 3.329 Average of experiments with food 1376 9.46 32.51 3.437 28.19 2.980 In experiments Nos. 59 and 68, only the average for the experiment is given as the determinations were made on the composite urine. In the other experiments, the determinations were made daily. The ratios range from 3.124 on the third day of experiment No. 71 to 5.353 on the first day of experiment No. 80. The average of all the experiments without food is 3.784. Comparing the individual days with this average ratio, the variations are in general very slight. The comparison shows even a greater uniformity in the ratios for the averages of the different experiments. The lowest average ratio is 3.254 in experiment No. 69 and the highest 4.886 in experiment No. 77. The experiments on S. A. B. show that save in the case of experiment No. 77, the ratio is fairly constant. The two experiments with A. L. L. show similar uniformity. There is an unusually high ratio, 4.886, in experiment No. 77, which marks it as distinct from all the others. Ratio of Organic Matter to Nitrogen. The presence of varying amounts of ash and especially sodium chloride in the total solids of the urine would cause marked variations in the ratios of total solid matter to nitrogen. The nitrogen of urine is present in the organic matter and not in the ash of the urine, and hence if the ash be deducted from the total solids, the ratio of organic matter to nitrogen would be expected to be more nearly constant. Here again the ratio for experiments Nos. 59 and 68 are given only for the average per day for the whole experiment. The lowest ratio of organic matter to nitrogen is 2.646 on the second day of experiment No. 79. The highest ratio, 3.969, is found on the last day of experiment No. 77. For each gram of nitrogen^excreted, there is on the average 3.017 grams of organic matter. The ratio of organic matter to nitrogen is, therefore, much more nearly constant than that of total solid matter to nitrogen. 383 Influence of Inanition on Metabolism. The averages for the different experiments range from 2.687 to 3.802 grams. In the experiments with S. A. B., Nos. 71 and 73 show ratios that are not far from the average for all the experiments. In experiment No. 75 the ratio after the first day increases as the fast progresses and in experiment No. 77 there is an unusually high ratio on all days of the experiment. The excretion of non-nitrogenous organic matter is therefore indicated. While no tests were made for pathological constituents other than sugar or albumen (see p. 397) the possible formation of /? oxybutyric acid and similar compounds must not be lost sight of in this connection. (See p. 395.) excretion of carbon in the urine. There is a considerable amount of carbon excreted in the urine, the total quantity of which, although relatively small when compared with the total excretion of carbon as carbon dioxide from the lungs, is nevertheless sufficiently large to be taken into consideration in obtaining the carbon balance. The determination of nitrogen in urine is readily and rapidly made by the Kjeldahl method, but the determination of carbon presents a number of difficulties which render the absolute amount determined much less accurate than the corresponding determinations of nitrogen. Since, however, the amount of carbon excreted in the urine is only a small part of the total carbon elimi- nation of the body, a relatively large error in the determination of carbon in the urine may exist without noticeable effect on the carbon balance. Pettenkofer and Voit °° determined the carbon elimination through the urine for the first fasting day, compared it with the total nitrogen and obtained as a result of three different experiments a ratio of carbon to nitrogen of 0.7. In comparing the ratio of carbon to nitrogen obtained by other investigators, the differences in the methods emplo}red may explain whatever differences in ratios appear. Munk (7) in determining the carbon in the urine dried 5 or 10 cc. of the acidulated urine with powdered copper oxide on the water bath and after mixing the material with coarsely powdered copper oxide, burned it in the combustion tube as usual. Determinations of the urine obtained in the experiment with Cetti were possible on only two days, namely, on the first and fifth fasting days. On the first day, during which 13.55 grams of nitrogen were eliminated, there were 12.63 grams of carbon giving the ratio C : N = 0.9. On the fifth fasting day, there were 14.85 grams of carbon and 10.69 grams of nitrogen, the ratio being 1.39. Munk states that in his opinion, the determinations made on the fifth day are too large. The determinations of Breithaupt's urine were made very carefully and the ratios obtained are given in table 201, which also includes the results of the more recent experiments on J. A. (9) in which the carbon was determined by the moist combustion process of Kjeldahl. 80Zeit. f. Biol. (1866), 2, p. 479. The amount of carbon in the urine for each day was 8.25, 8.05, and 9.30 grams, respectively. Carbon in Urine. 383 In only one of the several fasting experiments previously made in tins laboratory n was the carbon determined in the urine. In experiment No. 36, which lasted one day there were 8.54 grams of carbon excreted and 11.34 grams of nitrogen. The ratio was C : N = 0.753. The method of determining the carbon in urine at present in use in this laboratory is as follows : Five or 10 cc. of urine (the amount depending upon the specific gravity) is drawn into an aluminum dish and evaporated nearly to dryness in a vacuum desiccator. The partially dried mass is transferred from the aluminum dish to a copper combustion boat and dried again, prior to combustion in the Liebig combustion tube. In this process there is unques- tionably a loss of nitrogenous and carbonaceous material, which it has as yet Table 201. — Ratio of carbon to nitrogen of urine Cetti, and J. A. in experiments with Breithaupt, Subject. Day of fast. First. Second. Third. Fourth. Fifth. Sixth. Breithaupt 0.88 .90 .667 0.95 .654 0.83 .734 0.68 .762 0.79 1.39 .822 0.89 Cetti J. A Note. — It is important to note that in all three experiments, the ratio is above unity only on the fifth day of Cetti's fast, and the author distinctly questions the accuracy of the determination for that day. been impossible to prevent. The method does, however, give as high results for carbon as any method with which we are familiar and consequently it has been used in all the experiments here given. The total quantity of carbon and the total nitrogen excreted daily in the urine together with the ratio of carbon to nitrogen are given in table 202. It should be remembered that in experiments Nos. 59 and 68, the apportionment of the carbon over the different days was made according to the nitrogen elimi- nation, and hence only the ratio for the total excretion for each experiment can be used. In all the other experiments, the carbon was determined for each day and hence the daily ratios may properly be used. The lowest carbon elimination in any experiment was on the first day of experiment No. 71, 5.22 grams, while the highest was on the fourth day of experiment No. 77, 14.57 grams. Both these experiments were with the same subject and thus they serve to show the wide variations that may occur in the carbon elimination in the urine of a fasting man. The most striking variation in any one experiment is seen in experiment No. 77, when on the first day the elimination of carbon was 7.97 and on the fourth day, 14.57 grams. In consid- ering the long experiments, it is noteworthy that the carbon elimination is invariably lowest on the first day and on the remaining days is relatively 61 U. S. Dept. of Agr., Office of Expt. Sta. Bui. 136 (1903), 384 Influence of Inanition on Metabolism. constant. The two most marked instances occur in experiments Nos. 77 and 75. In the former the carbon rose from 7.97 grams on the first day to 13.94 grams on the second, and in the latter from 8.14 grams on the first day to 11.50 grams on the second day, remaining practically constant thereafter. Table 202. — Ratio of carbon to nitrogen of urine in metabolism experiments without food. Ex- peri- ment num- ber. Subject and date. (a) Ni- tro- gen. CM Car- bon. (c) » Ratio of carbon to nitro- gen (b + a). Ex- peri- ment num- ber. Subject and date. (a) Ni- tro- gen. (b) Car- bon. (c) 1 Ratio of carbon to nitro- gen (b+a). 59.. 68.. 69.. 71.. 73.. 75.. B.F.D., 1903: Dec. 18-21, av- A.L.L., 1904: Apr. 27-29, av- A.L.L., 1904: Dec. 16-17 Dec. 17-18 Dec. 18-19 Dec. 19-20 Average. . . . S.A.B., 1905: Jan. 7-8 Jan. 8- 9 Jan. 9-10 Jan. 10-11 Average. . . . S.A.B., 1905: Jan. 28-29 Jan. 29-30 Jan 30-31 Jan. 31-Feb. 1.. Feb. 1-2 Average. . . . S.A.B., 1905: Mar. 5- 6 Mar. 6- 7 Mar. 7-8 Mar. 8- 9 Mar. 9-10 Mar. 10-11 Average. . . . Gins. 13.59 O-ms. 10.01 0.737 77.. 79.. 80.. 81.. 82.. S3.. 85.. 89.. S.A.B., 1905: Apr. 8- 9 Apr. 9-10 . , . Apr. 10-11 Apr. 11-12 Average. . . . H.E.S., 1905: Oct. 13-14 , . . Oct. 14-15 Average. . . . C.R.Y., 1905 : Oct. 27-28 Oct. 28-29 Average. . . . A.H.M., 1905 : Nov. 22-23 . Average. . . . H.C.K., 1905: Nov. 25-26 Average. . . . H.R.D., 1905: Dec. 5-6 Dec. 6-7 Average. . . . N.M.P., 1905: Dec. 9-10 Dec. 10-11 Average. . . . D.W., 1906: Jan. 10-11 Average Average, 43 days. Gms. 8.81 10.78 10.98 11.45 Oms. 7.97 0.905 13.94 1.293 13.70 1.248 12.65 10.06 .795 14.57 1.273 10.5l|l2.54 1.193 10.09 14.26 15.04 12.97 8.31 9.97 10.37 9.42 .824 .699 .690 .726 8.11 14.35 11.23 6.27 10.22 .773 .712 8.24 .734 13.09 9.52 .727 7.78 9.95 7.11 9.02 .914 .907 5.84 11.04 13.10 10.74 5.22 8.29 8.64 7.73 .894 .751 .660 .720 8.87 8.06 .909 9.11 13.05 7.52 9.48 .826 .726 10.18 7.47 .734 11.08 8.50 .767 10.29 11.97 11.54 10.39 9.98 7.47 8.31 7.93 7.66 7.89 .726 .694 .687 .737 .791 9.38 14.36 7.71 10.77 .822 .750 11.87 9.24 .778 10.83 7.85 .725 13.25 13.53 9.76 12.78 .737 .945 12.24 12.45 13.02 11.63 10.87 10.74 10.13 8.14 11.50 11.11 12.20 12.65 12.02 11.34 .665 .924 .853 1.049 1.164 1.119 1.119 13.39 11.27 .842 » .32 11.05 11.35 11.36 3 .22 7.48 8.22 .677 .724 7.96 .701 11.58 11.28 .974 9.99 14.46 8.38 9.87 .839 .683 12.23 9.12 .74 6 11.54 9.57 .829 1 Averages in this column obtained as ft -^ a. 2 Determined for urine lost. See p. 243. 3 Calculated. See p. 243. Carbon in Urine. 385 Thus it appears that the influences which affect the nitrogen elimination on the first day also affect the carbon elimination, and, indeed, in an even more striking manner. The ratio of the carbon to the nitrogen presents wide variations, the minimum being observed on the third day of experiment No. 71, namely, 0.660, and the maximum on the second day of experiment No. 77, 1.293. While in experi- ments Nos. 73 and 75, there appears to be a tendency for the ratio to increase as the fast progresses, in experiment No. 77, the ratio remains almost constant after the first day. Practically the same can be stated regarding experiments Nos. 69 and 71. In the series of 2-day experiments, the ratios exhibit rather wide fluctuations on the different days of different experiments, but the averages for the two days of each experiment agree quite closely, the extremes being 0.909 and 0.701. There seems to be no general rule regarding the fluctu- ations seen in the ratios for the first and second days of the different experi- ments. Of special interest is the fact that with the same subject, S. A. B., the average ratio varied from 0.725 in experiment No. 73 to 1.193 in experiment No. 77. In fact, in the four experiments with 8. A. B., there is a constancy in the ratio in the first two experiments and a marked increase in experiment No. 75, with a still further increase in experiment No. 77. If the high ratios obtained in the Middletown experiments are the results of errors in determinations, they may be accounted for in only two ways — first, the determination of nitrogen may be low, and second, the determination of carbon may be high. Eegarding the former point, it is hardly to be considered that the results as determined by the Kjeldahl method are too low. In fact, check tests with organic materials of known nitrogen content, which are frequently made, show the method and apparatus to be unusually accurate. Furthermore, in all these cases, the nitrogen was determined in all of the four periods as well as in the composite and the agreement was invariably very satisfactory. On the other hand, a critical inspection of the method for determining carbon renders it almost impossible to believe that the carbon results are too high. While Munk (7), in discussing his determination of carbon in Cetti's urine, was of the opinion that the excessive amount of water in the dried urine might have escaped absorption in the first calcium chloride tube of the absorp- tion system and thus be subsequently held in the potash tube, here the urine was completely dried in a vacuum desiccator and sulphuric acid was used for the first tube to absorb the moisture resulting from the combustion. Further- more, it is the general experience in all laboratories that in organic elementary analysis, the determination of carbon is somewhat lower than the theoretical. Considering, then, the possibilities of loss of carbon dioxide by the decom- position of urea or ammonium carbonate during the process of drying, it would appear that the results for carbon, if incorrect, are somewhat too low, and 25 386 Influence of Inanition on Metabolism. hence the ratios from the experiments with S. A. B. too low rather than too high. The apparently abnormal ratios between carbon and nitrogen in these fasting experiments have confirmation so far as the determination of the amount of carbon is concerned in the ratio of the energy to the nitrogen. (See discussion under energy section, p. 490.) From an inspection of the data given there, it would appear that the ratio of carbon to nitrogen observed is wholly in accord with what would be expected from the ratios of energy to nitrogen and hence support the view that the high ratios here observed actually exist and are not due to errors in analysis or computation. The possibility of an excretion of non-nitrogenous or low-nitrogenous material in the urine, thereby increasing the ratio of carbon to nitrogen, will be considered in discussing the elimination of creatine and creatinine. creatinine and creatine. Although the pressure of other work in this laboratory precluded analyses of the urine to secure information as to the partition of the nitrogen, Prof. Lafayette B. Mendel, of Yale University, kindly offered to make determinations of creatinine in the urine of experiments ISTos. 73 and 74, suggesting that, in view of the recent appearance of Folin's theory of protein metabolism, deter- minations of the creatinine output of a fasting man would be of interest. A Dubosc colorimeter was obtained later and in all experiments subsequent to No. 74, the determinations were made in this laboratory. Observations regarding the excretion of creatinine by fasting men have hitherto been confined to those of Baldi M and E. and 0. Freund (10) on Succi, and by Yan Hoogenhuyze and Verploegh (11) on a fasting girl. Baldi determined the creatinine elimination of Succi during the Florence fast by using the ISTeubauer method. On the seventh fasting day he found by this method 0.8011 gram of creatinine; on the twelfth, 0.7159 gram, and on the seventeenth, 0.4029 gram. After the seventeenth day, although creatinine could be determined qualitatively, it was not present in weighable amounts. E. and O. Freund (10) likewise using the Neubauer method, determined the creatinine in the 21-day fast of Succi made in Vienna in 1896. According to their observations, the creatinine-nitrogen increased from 0.134 gram, on the first day to 0.578 gram, on the ninth day. The authors explain that the increase was probably caused by the exercise which Succi had with the sabre. On the tenth and eleventh days the creatinine-nitrogen fell rapidly. The twelfth day the output was but 0.08 gram. The decrease continued as the fast progressed, there being but 0.025 gram of creatinine-nitrogen excreted on the twenty-first day. K Centralblatt f. klin. Medic. (1889), 10, p. 651. Creatinine and Creatine. 387 Folin's es method for determining creatinine was used in the experiments with the fasting girl Flora Tosea. It is not stated definitely whether the urine was heated with hydrochloric acid before making the colorimetric observations. It is not, therefore, clear whether the amounts of creatinine include preformed creatine (as creatinine). Urine was collected in three periods of the day, from 10 a. m. to 4 p. m., 4 p. m. to 10 p. m., and 10 p. m. to 10 a. m. The last food was taken on the morning of June 10, 1905, and no more food was consumed until June 25. During the fasting period the subject remained at rest, save for 2 hours of gymnastic exercise on June 17. The quantity of creatinine in the urine on the day preceding the fast was 1.087 grams. The excretion de- creased regularly and rapidly to the eighth day. On the day when muscular exercise was taken, the creatinine amounted to only 0.469 gram, while on the following day it rose to 0.689 gram. During the three days before muscular work was done a total of 1.662 grams was excreted; the total output for the three following days was 2.006 grams. The excretion then decreased to about 0.5 gram daily, remaining fairly constant during the remainder of the fast. The new method of Folin for determining creatinine by means of the Jaffe reaction was used in many of the experiments here reported. This reaction is not given by creatine and hence it is necessary to heat the urine with hydro- chloric acid to convert the creatine to creatinine. A determination of the creatinine in the urine before and after heating with hydrochloric acid gives, therefore, a measure of the amount of creatine (in terms of creatinine) which is present in the urine. It has been found in a large number of experiments, that the amount of preformed creatine in the urine is normally very small. In table 203 the results of the creatine and creatinine determinations are given for the fasting experiments, for the two food experiments following fasting, namely Nos. 74 and 76, and for the two nitrogen metabolism experi- ments Nos. 1 and 2 made with S. A. B. In all experiments after No. 74, the determinations were generally made of both preformed and total creatinine and consequently in the last column of the table is recorded the weight of creatine (expressed as creatinine) excreted. In experiments Nos. 73 and 74 the quantities of urine that could be spared for samples to be sent to New Haven were small, and hence the results were not as satisfactory as those made in our laboratory where analyses could be repeated as desired, and where they could be made on the day immediately following the collection of the sample. It is, therefore, more especially with the experi- ments beginning with No. 75 that the results are of especial importance. The excretion of total creatinine, namely, preformed creatinine plus creat- inine formed by heating the creatine of the urine with acid, remains singularly constant on all days of the fast, even during the 7-day fast, experiment No. 75. 63 Zeitschrift f. physiol. Chemie (1904), 41, p. 223. 388 Influence of Inanition on Metabolism. The quantity of preformed creatinine on the contrary, is seen to diminish as the fast progresses. In general, the diminution is quite regular for the first 4 days of the experiment and indeed, when the results for experiments Nos. 75 and 77 are compared, the agreement between the two experiments is remark- able. On the fifth day of experiment No. 75, the lowest excretion of preformed creatinine is observed. After this day, the excretion increases slightly until the end of the experiment. Table 203. — Creatinine and creatine excreted in urine in metabolism experiments with and without food. Creatinine. Creatinine. Ex- peri- ment num- ber. Ex- peri- ment num- ber. Subject and date. Total, includ- ing crea- tine. Pre- form- ed. Crea- tine.1 Subject and date. Total, includ- ing crea- tine. Pre- form- ed. Crea- tine.1 73.. S.A.B. Jan. 1905: 28-29 Gms. 1.250 Qms. Gms. 77.. S.A.B., 1905: Apr. 8- 9. . . Gms. 1.342 Gms. 1.202 Gms. 0.140 Jan. 29-30 1.200 .... .... Apr. 9-10... 1.343 1.027 .316 Jan. 30-31 .871 • • ■ • .... Apr. 10-11... 1.383 .968 .415 Jan. 31_Feb. 1. .791 .... .... Apr. 11-12... 1.386 .848 .538 Feb. 1-2 1.110 • • ■ ■ .... N2.. S.A.B., 1905: 74.. 8.A.B., 1905: Apr. 12-13... 1.450 1.407 .043 Feb. 2-3 1.06 .... • ■ • • Apr. 13-14... 1.420 1.384 .036 Feb. 3-4 1.01 • • ■ ■ .... Apr. 18-19 . . . 1.334 1.300 .034 Feb. 4-5 .86 .... .... 79.. H.E.S., 1905: 75.. S.A.B. , 1905: Oct. 13-14... 1.225 1.212 .013 Mar. 4- 5.... 1.237 1.212 0.025 Oct. 14-15... 1.418 1.254 .164 Mar. 5- 6. . . . 1.294 1.061 .233 80.. C.R.T., 1905: Mar. 6- 7. . . . 1.407 .956 .551 Oct. 27-28... 1.505 1.505 > . . • Mar. 7- 8. . . . 1.325 .865 .460 Oct. 28-29... 1.538 1.398 .140 Mar. 8- 9. . . . 1.214 .712 .502 81.. A.H.M., 1905: Mar. 9-10 1.318 .733 .585 Nov. 21-22. . . 1.226 1.184 .042 Mar. 10-11.... 1.270 .782 .488 Nov. 22-23. . . 1.357 1.229 .128 76.. S.A.B., 1905: 82.. H.C.K., 1905: Mar. 11-12 1.411 1.137 .274 Nov. 24-25. . . 1.726 1.643 .083 Mar. 12-13. . . . .... 1.292 Nov. 25-26... 1.795 1.730 .065 Mar. 13-14 .... 1.346 .... 83.. H.R.D., 1905: Nl.. S.A.B. 1905: Dec. 5-6 1.317 1.217 .100 Mar. 14-15 1.566 1.493 .073 Dec. 6-7 1.363 1.129 .234 Mar. 15-16 .... 1.433 .... 85.. N.M.P., 1905: Mar. 16-17.... 1.251 .... Dec. 9-10... 2 1.543 2 1.525 2 .018 Mar. 21-22.. .. • • ■ ■ 1.487 .... Dec. 10-11... 1.703 1.541 .162 Mar. 22-23. . . . 1.523 53 89.. D.W., 1906: Jan. 10-11... Jan. 11-12. . . 2.091 1.966 2.028 1.922 .063 .044 1 In terms of creatinine. 3 Does not include possible amount in urine lost. While the quantity of preformed creatinine gradually diminishes as the fast progresses, the amount of creatine, which in normal urines is extremely small, gradually increases, and on the sixth day of the fast (No. 75), there is excreted 0.585 gram of creatine (expressed in terms of creatinine). Creatinine and Creatine. 389 The same may be said regarding the excretion of creatine in experiment No. 77. In the shorter experiments, the amount of total creatinine, in general, remains fairly constant. The results for creatine show a greater amount on the second day of the experiment than on the first day in five of the experiments, while in Nbs. 82 and 89, there is but a small quantity of creatine excreted on both days, the lesser amount being excreted on the second day. The interpretation of these results, according to Folin's theory of protein metabolism, would indicate that the amount of tissue protein disintegrated (endogenous protein katabolism), even during a prolonged fast, remains constant from day to day. While on the one hand it might reasonably be expected that the tissue protein would be more rapidly katabolized during fasting, when the body has to subsist on its own material, than during the ingestion of food, yet on the other hand it is not difficult to conceive that during fasting the body has the power to protect its organized or tissue protein more thoroughly than when well supplied with food. Hence the constancy in the total creatinine elimination, viewed in the light of the Folin theory, furnishes no evidence to show either an increased or diminished endogenous protein katabolism during inanition. While with a normally fed individual creatine is wholly converted by the body to creatinine before being excreted, apparently with the fasting individual the body gradually loses this power of converting creatine to creatinine as the fast progresses.64 The data thus far obtained are insufficient to show clearly the true significance of this abnormality in the power of the body to convert creatine. Further experiments are necessary to solve this specific problem. The fact, however, that during prolonged fasting the body loses to a marked degree the power of converting creatine to creatinine is suggestive in interpreting results of the katabolism of tissue protein and also of the synthetic power of the body in general. It is furthermore clear that while the majority of the short experi- ments indicate a loss of power of conversion, experiments should be continued for more than two days to throw definite light on this subject. In a recent series of experiments made by Folin,85 he has advanced the suggestion that creatine when ingested is a food serving a purpose different from that of the ordinary amino acids. These experiments imply that when the body is flooded with protein, as after a diet rich in nitrogenous material, the ingestion of creatine is followed by its immediate excretion, while creatine ingested after the supply of reserve protein has been depleted by a low protein diet is wholly retained in the body. Neither the creatine, creatinine, nor, indeed, the total nitrogen excretion are affected by the ingestion of creatine under these conditions. 84 For another possible explanation of the increase in creatine excretion see p. 458. 86 Festschrift fur Olof Hammersten, Upsala (1906). 390 Influence of Inanition on Metabolism. Somewhat similar results were obtained by Klercker.68 While in some ways it is difficult to harmonize the results of Folin and Klercker with those here reported, both sets of experiments emphasize the fact that it is difficult at times for the body to convert creatine to creatinine. It would appear, then, that if the body possesses a ferment which can dehydrate creatine to creatinine, the secretion or possibly the activity of this ferment is markedly decreased by fasting. In this connection, it is important to observe the rapidity with which the body recovers its power to convert creatine to creatinine after the ingestion of food. Thus, in experiment No. 76, the excretion of creatine which on the day previous (seventh fasting day) was 0.488 gram was reduced to 0.274 gram, although this day differs in no wise from the day preceding except in that the following amounts of food were eaten: 651 grams modified milk, 123 grams apple, 313 grams orange juice, 178 grams shredded wheat, and 10 grams gluten crackers. Unfortunately, the determinations of total creatinine on the two following days of experiment No. 76 could not be made, but from the determinations of creatinine, made before heating with acid, the deduction may be made that the normal elimination of creatinine had been reached on the second day of the experiment, and that in all probability the amount of creatine excreted on this day was very small. In fact, 3 days later, namely, March 14-15, the amount of creatine was again determined and found to be but 0.073 gram. The power to convert creatine was regained in an even more striking manner in nitrogen metabolism experiment No. 2. The creatine excretion (as creatinine) of April 11-12 was 0.538 gram. On the next day with food (April 12-13) it was only 0.043 gram. This practical disappearance of creatine from the urine was incidental to the ingestion of food on the first day that the subject was out of the respiration chamber and subsisting on a creatine-free diet of his own selec- tion. The kinds and amounts of food eaten this day are given in table 176. While in all cases, the creatine practically disappeared upon the ingestion of food, there was no marked increase in the total creatinine from day to day, which indicates that the creatine had simply been converted to creatinine and excreted as such. In all of the food experiments following fast, care was taken to avoid foods containing creatine and creatinine, though throughout the nitro- gen metabolism experiments the diet was practically creatine-free, as the sub- ject ate meat on only one or two occasions. On all days when samples were taken for creatinine determinations no meat had been taken for 24 hours. Excretion of creatinine by periods. — In experiments Nos. 75 and 76 the determinations of preformed creatinine, i. e., creatinine determined before heating with acid, were made on the samples collected during each period. ^Zeit. f. gesammte Biochemie (1906), 8, p. 59. Creatinine and Creatine. 391 The data, therefore, furnish evidence as to the excretion of creatinine during the different periods of the day. Such determinations as are available have been placed in table 204. A comparison of the total amount in 24 hours as recorded in this table and the corresponding amount as recorded in table 203 shows slight discrepancies on the different days. The record in table 203 is that determined on the daily composite sample, while that in table 204 is the sum of the amounts of preformed creatinine for the four periods of each day. When the large volumes of urine are taken into consideration the agreement is as satisfactory as could well be expected. The table shows the actual amount of preformed creatinine excreted in the periods from 7 a. m. to 1 p. m., from 1 p. m. to 7 p. m., and the total for the first 12 hours of the day, the amount for the night being given in one column as was the case in table 198, showing the distribution of nitrogen by periods. The unequal division of the night into a 4-hour and an 8-hour period makes this presentation of the results more satisfactory. Table 204. — Preformed creatinine by periods eliminated in urine in metabolism experiments Nos. 75 and 16. Exper- iment num- ber. Subject and date. 7 a.m. to 1 p.m. 1 p.m. to 7 p.m. Total for first 12 hours. Am'nt. Propor- tion of total for 24 hours. Total for last 12 hours. Am'nt. Propor- tion of total for 24 hours. Total amount in 24 hours. 75.. 76. 1905. S.A.B.: Mar. 4- 5 Mar. 5- 6 Mar. 6- 7 Mar. 7- 8, Mar. 8- 9 Mar. 9-10 Mar. 10-11 S.A.B.: Mar. 11-12, Mar. 12-13. Mar. 13-14, Orams. 0.340 .313 .236 .243 .169 .157 .148 .263 .365 .348 Orams. 0.330 .256 .222 .212 .148 .158 .175 .302 .304 .350 Orams. Per ct. Orams. 0.670 54.6 0.558 .569 54.4 .476 .458 52.5 .415 .455 58.3 .325 .317 48.8 .332 .315 48.5 .335 .323 44.4 .405 .565 50.7 .549 .669 54.3 .564 .698 51.9 .648 Per ct. 45.4 45.6 47.5 41.7 51.2 51.5 55.6 49.3 45.7 48.1 Grams. 1.228 1.045 .873 .780 .649 .650 .728 1.114 1.233 1.346 Considering first the apportionment of the total preformed creatinine excretion between the day and the night periods, the figures show that during the day the per cent may be as low as 44.4 of the total and as high as 58.3. The percentage excreted during the day has a distinct tendency to become lower as the fast progresses. Aside from the 58.3 per cent on the fourth day of the fast, the decrease is very constant. On the ingestion of food, the excretion of performed creatinine during the first 12 hours of the day averages about 52 per cent. 392 Influence of Inanition on Metabolism. Table 205. — Proportions of nitrogen of total creatinine and of creatine in total nitrogen excreted in urine in metabolism experiments with and without food. Experiment number. Subject and date. (a) Nitrogen in total creatin- ine. (6) Nitrogen in creatine. (c) Total nitrogen in urine. Proportion of total nitrogen. (d) In total creatin- ine (o c ). (e) In creatine {h+c). 73. 74. 75. 76. 1 Nitrogen metabolism. 77. 2 Nitrogen metabolism. 79 80 81 82 83 85 89 S.A.B. S.A.B. S.A.B., S.A.B., S.A.B. , S.A.B., S.A.B., H.E.S., C.R.T., A.H.M., H.C.K., H.R.D., N.M.P., D.W., 1905. Jan. 28-29 Jan. 29-30 Jan. 30-31 Jan. 31-Feb. 1 Feb. 1- 2 Feb. 2- 3 Feb. 3- 4 Feb. 4- 5 Mar. 4- 5. . . . Mar. 5- 6 Mar. 6- 7 Mar. 7- 8 Mar. 8- 9 Mar. 9-10 Mar. 10-11 Mar. 11-12 Mar. 12-13 Mar. 13-14 Mar. 14-15 Apr. 8- 9 Apr. 9-10 Apr. 10-11 Apr. 11-12 Apr. 12-13 Apr. 13-14 Apr. 18-19 Oct. 13-14 Oct. 14-15 Oct. 27-28 Oct. 28-29. .. Nov. 21-22 Nov. 22-23 Nov. 24-25 Nov. 25-26 Dec. 5- 6 Dec. 6- 7 Dec. 9-10 Dec. 10-11 1906. Jan. 10-11. ... Jan. 11-12 Gram. 0.465 .447 .324 .294 .413 .395 .376 .320 .460 .482 .524 .493 .452 .491 .473 .525 .583 .499 .500 .515 .516 .540 .529 .497 .456 .528 .560 .572 .456 .505 .642 .668 .490 .507 .574 .634 •778 .732 Gram 0.008 .075 .177 .148 .161 .188 .157 .088 .023 .045 .101 .133 .173 .014 .012 .011 .004 .053 .000 .045 .013 .041 .027 .021 .032 .075 1.006 .052 .020 .014 Grams. 10.29 11.97 11.54 10.39 9.98 10.74 8.25 6.78 12.24 12.45 13.02 11.63 10.87 10.74 10.13 10.17 7.15 7.82 12.61 8.81 10.78 10.98 11.45 11.14 11.65 12.72 8.11 14.35 7.78 9.95 9.11 13.05 9.38 14.36 13.25 13.53 2 11.05 11.35 9.99 14.46 zr ct. Per ct. 4.52 .... 3.73 | 2.81 .... 2.83 .... 4.14 . . . . 3.68 4.56 • . • ■ 4.72 3.76 3.87 4.03 4.24 4.16 4.57 4.67 5.16 4.62 5.67 4.64 4.69 4.51 4.85 4.54 3.91 5.62 3.68 7.20 5.75 5.01 3.87 6.84 4.65 3.70 3.75 5.20 5.59 7.79 5.06 0.07 .60 1.36 27 .48 .75 .55 .87 .18 .51 .94 1.21 1.51 .13 .10 .09 .05 .37 .00 .45 .14 .31 .29 .15 .24 .55 •05 .46 .20 .10 1 Does not include possible amount In urine lost. 2 Does not include nitrogen determined in urine lost. Creatinine and Creatine. 393 On comparing the excretion of creatinine during the first 12 hours of the day with the total nitrogen (see table 198) it is seen that while in experiment No. 75 on the average 57.7 per cent of the total nitrogen for 24 hours is excreted between 7 a. m. and 7 p. m., but 51.6 per cent of the preformed creatinine is excreted during the same period. Furthermore, while the proportion of nitrogen excreted during the first 12 hours of the day remains quite constant on all days of the fast, the proportion of creatinine has a distinct tendency to diminish as the fast progresses. Proportions of creatine nitrogen and total creatinine nitrogen excreted. — The proportions of the total urinary nitrogen derived from creatinine nitrogen and creatine nitrogen are of interest. The amounts of total nitrogen, total creatine- nitrogen and of total creatinine-nitrogen, as well as the proportions, are given in table 205. The absolute amounts of total creatinine-nitrogen vary from 0.294 gram to 0.778 gram. While these wide variations are noted with different individuals, with the same individuals the absolute amount is relatively constant. The nitrogen of creatine is inappreciable save on the later days of the longer fasts, the highest amount observed being on the sixth day of experiment No. 75, 0.188 gram. The relative constancy in the creatinine-nitrogen from day to day is not accompanied by a similar constancy in the total nitrogen excretion, hence the proportion of creatinine-nitrogen in total nitrogen varies somewhat. In experiment No. 75, there is a distinct tendency for the proportion of creatinine-nitrogen to increase as the fast progresses, while in experiment No. 77 the reverse is true. An inspection of the figures shows that in almost every instance the variation is due to the fluctuations in the total nitrogen of the urine rather than to the nitrogen of creatinine. The proportion of creatine nitrogen in the total nitrogen is exceedingly small on all save the later days of fast. The highest per cent observed was 1.75 on the sixth day of experiment No. 75. It is highly probable that errors in determination may account for the minute quantities of creatine found on the first day of fasting. Relation of total creatinine excreted to body-weiglit. — As Folin has pointed out, the total creatinine excretion is relatively unaffected by the total nitrogen excretion when creatinine-free diets are consumed. The total creatinine elimi- nation does vary, however, according to the individual, and in general, with body-weight. With persons of the same body-weight, nevertheless, marked differences in the proportion of body fat, and height, may result in marked variations in the amount of creatinine excreted per kilo of body-weight. For purposes of comparison, the data regarding the body-weight and the total creatinine, as well as the total creatinine per kilo of body-weight, are given in table 206. 394: Influence of Inanition on Metabolism. The average total creatinine excreted per kilo of body-weight is 23.2 mg. The lowest observed amount was 17.6 mg. in experiment No. 74 and the highest 27.8 mg., during a 1-day observation in the first nitrogen metabolism experiment. Table 206. — Total creatinine excreted per day in urine per kilogram of body-weight in metabolism experiments with and without food. Experiment number. Subject and duration of experiment. (o) Body- weight.1 (b) Total creatinine (including creatine). (c) Total creatinine per kilogram of body- weight (6-s-a). 73 S.A.B., Jan. 28 to Feb. 2, 1905 S.A.B., Feb. 2 to 5,1905 Kilos. 57.1 55.4 57.7 255.7 4 56.3 59.5 6 57. 7 860.1 56.0 67.5 60.9 70.4 54.9 66.5 77.8 Grams. 1.044 .977 1.295 31.411 51.566 1.364 U.435 9 1.334 1.322 1.522 1.292 1.761 1.340 1.623 2.029 Mgms. 18.3 17.6 22.5 3 25.3 527.8 22.9 '24.9 922.2 23.6 22.5 21.2 25.0 24.4 24.4 26.1 74 75 . 76 S.A.B., Mar. 11 to 14, 1905 S.A.B., Mar. 14 to 23, 1905 S.A.B., Apr. 8 to 12, 1905 S.A.B., Apr. 12 to 19, 1905 C.R.Y., Oct. 27 to 29, 1905 H.R.D., Dec. 5 to 7,1905 N.M.P., Dec. 9 to 11, 1905 D.W., Jan. 10 to 12, 1906 1 Nitrogen metabolism. . 77 2 Nitrogen metabolism. . . 79 80 . 81 82 83 89 1 Average of weight at beginning and than underclothes, and deducted from end of experiment. weighing of April 14. a Average of weighings of first day. 7 Amount for first two days. 8 Amount for first day. 8 Average weight of April 18-19 less l 4 Weight at 7 a. m., "March 14. kg. assumed for clothing other than under- 8 Amount for first day. clothes. a Average weight for first two days. One • Amount for April 18-19. kg. assumed as weight of clothing other In comparing different experiments on the same subject, as, for example, experiments Nos. 73-77 and the two nitrogen metabolism experiments, it is to be noted that in some of them the observations were confined to one day. Aside from the results obtained in experiments Nos. 73 and 74, the uniformity, not only with the same individual in the different experiments, but likewise with different individuals, is very striking. The results here obtained are wholly in accord with those observed by Folin,67 Closson,*8 Klercker," and Van Hoogen- huyze & Verploegh (11) in normal individuals. In discussing the results of the determinations of creatine and creatinine in the urine of fasting men by the Folin method, the possibility of the presence of other bodies in the urine giving color reactions with sodium picrate must be "Amer. Journ. Physiol. (1905), 13, p. 85. "Amer. Journ. Physiol. (1906), 16, p. 252. 6»Beitrage zur chem. Physiol, u. Path. (1906), 8, p. 59. Uric Acid. 395 considered. As was pointed out by Jaffe/0 acetone gives the same color with sodium picrate that creatinine produces. Folin71 found that acet-acetic acid, acet-acetic ether, and hydrogen sulphide also give the reaction, but, as he points out, these constituents are all pathological and can easily be removed from urine. With the urine from fasting men, the researches of Brugsch (12), especially, have shown the presence of (3 oxybutyric acid as a constant constit- uent as well as amino acids and acetone. Kuelz " showed that even after 3 or 4 days of fasting in the case of insane patients, (3 oxybutyric acid was present. The influence of these acids on the Jaffe reaction has not to our knowledge been studied. Since unquestionably acetone 73 is formed in the urine of fasting man, it is at least possible that constituents other than creatinine may produce the color with sodium picrate. No precautions were employed to remove these abnormal constituents of the urine, but during the heating with hydrochloric acid the acetone would in large part be expelled. It is possible to conceive of an increased production of compounds giving a color with sodium picrate as the fast progresses, and thus, in part at least, account for the apparently constant elimination of total creatinine. On the other hand, it is hardly probable that the production of other color-producing materials should proceed at such a rate as to exactly compensate for any possible falling off in the production of creatinine, and hence while the results must be accepted with some reserve, it would appear that during fasting, the output of total creatinine remains constant, while the relative proportion between preformed creatinine and creatine indicates a constant change, the proportion of creatine increasing as the fast progresses. Since this report was written evidence regarding the presence of creatine in pathological urine has been collected which would tend to sustain the alternate explanation of the presence of creatine given in a subsequent section of this report. (See p. 458.) URIC ACID. Unfortunately, the excretion of uric acid in fasting man has been only imperfectly studied, and in the experiments here reported but few observations were made. In some of the samples of the urine sent to New Haven for determination of creatinine, Professor Mendel kindly made determinations of the uric acid. It was found impracticable to carry out these determinations in the subsequent experiments with the limited amount of urine at our disposal. T0Zeit. f. physiol. Chemie (1886), 10, p. 399. nZeit. f. physiol. Chemie (1904), 41, p. 224. 73 Zeitschrift f. Biologie (1887), 23, p. 338. "E. & 0. Freund also report acetone and acet-acetic acid in Succi's urine during the Vienna fast. 396 Influence of Inanition on Metabolism. The earliest records of the determination of uric acid in the urine of fasting man with which we are familiar are those of Banke." In three 24-hour fasts he found 0.236, 0.033, and 0.24 gram of uric acid, respectively. The author makes no comment on the unusually small amount obtained in the second experiment. Pettenkofer and Voit75 determined the uric acid in a 24-hour fast. The authors report that the subject excreted 0.5584 gram of uric acid. Unques- tionably this excretion was somewhat augmented by the considerable quantities of beef extract that were consumed. Uric acid was not determined by Luciani (4) in the experiments on Succi. Monaco 76 determined the uric acid in a 20-day fast made by Succi. Using the Salkowski method he obtained on the last day with food 0.8228 gram of uric acid, and on the eighteenth and twentieth days of the fast 0.2565 and 0.244 gram, respectively. On the second day after the fast the uric acid amounted to 0.5539 gram. According to the author the relation between the total nitrogen and uric acid did not appear constant. In the two fasting experiments reported by Sadovyen (2) uric acid was determined. In the first experiment, which lasted 2 days, and during which no water was consumed, the subject excreted 0.301 gram of uric acid in 24 hours. On the second day 0.291 gram was excreted. In a subsequent 4-day fasting experiment, the uric acid excretion was 0.412, 0.201, 0.301, and 0.357 gram, respectively. The most elaborate series of observations recording the excretion of uric acid in fasting man is that of E. & O. Freund (10). The uric acid-nitrogen was determined on nearly every day of the 21-day fast. The largest amount recorded was on the first day, 0.29 gram (0.87 gram uric acid), and the smallest amount on the twenty-first day, 0.046 gram (0.138 gram uric acid). The quantity of uric acid-nitrogen gradually diminished as the fast progressed, although but little change occurred after the fifth day. Brugsch (12) reports only the total purin-nitrogen, no attempt being made to isolate the nitrogen of uric acid. For the last 7 days of the 30-day fast, the total purin-nitrogen varied from 0.104 gram to 0.146 gram. Schreiber & Waldvogel" observed the uric acid output of two individuals each of whom fasted 3 days. The uric acid excretion for the 3 successive days was 0.290, 0.233, and 0.197 gram with one subject, and 0.718, 0.405, and 0.205 gram with the second subject. E. Orgler 78 reports that during a 1-day fast he excreted 0.480 gram of uric acid. "Arch. Anat. u. Physiol. (1862), p. 340. "Zeitschr. f. Biol. (1866), 2, p. 479. "Bull, della Soc. Lancis degli ospedali d. Roma (1894), xrv, 2, p. 102. Ab- stracted in Schmidt's Jahrbiicher, 252, p. 109. "Archiv f. experimentelle Path. u. Pharm. (1899), 42, p. 69. "Allg. med. Ctrl.-Zeitung, 1896, No. 66. Cited by Magnus-Levy, Physiologie des Stoffwechsels (1905), p. 134. Pathological Constituents. 397 The quantities of uric acid observed in the fasting experiments here reported are unusually small. For the five days of fasting experiment No. 73, the quantities of uric acid per day were 0.172, 0.122, 0.082, 0.059, and 0.054 gram, respectively. It is to be noted, however, that on the last 3 days of the fast, the quantities of sample available for the uric acid determinations were so small as to render the results reasonable estimates rather than accurate determina- tions. The fact remains, however, that remarkably small amounts of uric acid were excreted by this subject on the last 3 days of the fast. With the1 resumption of the ingestion of food, which was of a purin-free nature, i. e., milk, the uric acid excretion increased. On the 3 days of experiment No. 74, in which the subject remained inside the chamber, the uric acid excretion was 0.329, 0.537, and 0.407 gram, respectively. On the first day after the subject left the respiration chamber, when he partook of an elaborate diet of varied nature, the uric acid excretion increased to 0.744 gram. In the absence of more complete data, little can be said regarding the uric acid excretion in the fasting experiments here reported, save that the evidence seems to indicate that remarkably small amounts of uric acid are excreted by fasting man after the first one or two days of the fast. This observation agrees in general with that of E. & O. Freund (10) save that the estimated amounts excreted by S. A. B. are considerably less than the quantities for the corresponding fasting days observed on Succi in the Vienna fast. The unusually low uric acid excretion observed in one day fasts by Ranke 79 likewise indicates that during even a short period of inanition the uric acid output may be greatly reduced. pathological constituents of the urine. Of the ordinary pathological constituents in urine (albumen and sugar) albumen has not been found in any of the long fasts made by Succi, and in but one, E. & O. Freund (10), was sugar present. The observations of Brugsch (12) on acidosis showed the presence of /? oxybutyric acid. Careful tests of all the samples of urine of the experiments here reported showed in no instance the presence of albumen or sugar. The large increase of carbon in the urine in experiment No. 77, may, as has been before stated, have been caused by the formation of /3 oxybutyric acid, although no direct evidence of its presence was obtained. sulphur. Sulphur in the urine is derived almost wholly from the oxidation of the sulphur in the protein molecule, for a very small amount of preformed sulphuric acid is taken with the food or drink. Consequently, as an index of protein katabolism, the total elimination of sulphur in the urine may be of value. 79Loc. cit. 398 Influence of Inanition on Metabolism. Sulphur occurs in the urine in at least three different forms: First, preformed sulphuric acid; second, oxidized sulphur which is combined with some organic radical to form the so-called ethereal sulphates; and third, the unoxidized or " neutral " sulphur. Not only did some of the earlier investi- gators fail to recognize clearly the different forms of sulphur or their signifi- cance, but in the records of the determinations which have been made there has been more or less lack of uniformity in the method of expressing the results. Some authors have expressed their data as S, others as H2S04 and still others as S03. Since but little is known of the nature of the neutral sulphur, it seems most logical to record the total sulphur elimination as S, rather than H2S04 or S03, and in subsequent reports it will be so expressed. In consideration of the fact that much of the earlier work is here presented, and that physiological chemists have not been accustomed to interpret the results of sulphur determinations in terms of sulphur, in all the experiments here reported the old usage is retained. To pave the way for the subsequent expression of the results of analyses in terms of sulphur, the averages for the different days of the experiment are given in heavy faced type in tables 208, 210, and 211 as sulphur (S) along with the results expressed as sulphur trioxide (S03). From a careful computation of the results as presented, the data for earlier fasting experiments (all calculated on the same basis, namely, S03, for purposes of comparison) are given in table 207 herewith. Unfortunately the determinations on Succi in Florence and Naples did not include the total sulphur, hence there are no data for these fasts regarding the neutral sulphur. E. & 0. Freund (10) reported the total, the inorganic, the ethereal, and the neutral sulphur. The results reported by Sadovyen (2) did not include the neutral sulphur and hence are comparable to the results in other experiments in which the inorganic and ethereal alone were determined. It has recently been pointed out by Folin 80 that the determination of sulphuric acid as barium sulphate involves technicalities that have in many instances been overlooked. Hence the comparison of the results obtained by different investigators by the determination of sulphuric acid in the urine is somewhat questionable. In fact, it is tolerably clear that much of the older work involves errors which render even the determinations of total sulphur unreliable. Moreover, any errors in the determinations of either total sulphur or ethereal sulphates affect the measure of the neutral sulphur, since the latter is determined indirectly as the difference between the total sulphur and the ethereal (and inorganic) sulphates. So far as the inorganic and ethereal sulphur is determined, it is probable that with certain limitations the results may, in general, be compared. 80 Journal of Biological Chemistry, 1906, 1, p. 131. Sulphur in Urine. 399 Total sulphur. — The excretion of total sulphur expressed as S03 in the fasting experiments here reported is given in table 208. The averages, ex- pressed as S and S03, are both included in the table. At the time these determinations were made Folin's critical discussion of the method of determining sulphur had not appeared and hence these deter- minations are probably subject to all the errors there pointed out. Nevertheless, they represent a determination as accurate as most of the earlier work, and also, in all probability, give results that are comparable so far as the different days of the same experiment are concerned. Table 207. — Amounts of sulphur (as SOJ eliminated in urine daily by fasting subjects. Day of fast. Succi. Cetti. Breithaupt. J.1 (a) 6 (6) At Vienna. (f) (.7) a 05 (h) («) 0") a (*) (1) (c) (d) OS (e) a .... 80 C.R.Y., Oct. 27-28, 1905.. .969 1.710 • • ■ • • • • . ■ • 81 A.H.M., Nov. 21-22, 1905.. 1.427 2.036 ... ... .... .... 82.... H.C.K., Nov. 24-25, 1905.. 1.423 1.915 • . • . . > .... .... 83 H.R.D., Dec. 5- 6, 1905.. 2.173 1.755 > • • . . . .... .... 85 N.M.P., Dec. 9-10, 1905.. 1 1.507 1.506 • ■ • ... . • ■ • .... 89 D.W., Jan. 10-11, 1906.. 1.730 2.148 1.468 .58S 1.806 .733 1.867 .748 1.765 .707 1.717 .688 1.648 .660 1.553 .623 1 Does not include possible amount in urine lost. There were 5 experiments in which the fast continued for 4 days or more. The output ranges from 1.651 grams in experiment No. 77 to 1.971 grams in experiment No. 69. In every instance there was a decrease from the amount excreted on the third day. Observations regarding the fifth day of the fast are found in but 2 experi- ments, Nos. 73 and 75. In No. 73, the excretion was 1.765 grams, a slight increase over the fourth day, while in experiment No. 75, the excretion was somewhat less than on the fourth day, i. e., 1.668 grams. The sulphur trioxide excretion for the sixth and seventh days of experiment No. 75 showed a slight but persistent decrease. It is interesting to note that the excretion on the seventh day was almost identical with the excretion of the first day of experiment No. 75. Considering, then, the results as obtained from these experiments, it is seen that the excretion of sulphur increases on the second day. There is an increase on the third day and a steady diminution on the succeeding days of the fast. Ratio of nitrogen to total sulphur. — Since practically all the sulphur in the urine results from the katabolism of protein, a parallelism in the excretion of Sulphur in Urine. 401 nitrogen and sulphur is to be expected. Indeed, it has long been urged that the sulphur excretion is as true a measure of protein katabolism as is the nitrogen excretion. In the fasting experiments with Cetti and Breithaupt, Munk (7) computed the ratios between the nitrogen and sulphur N/S. These ratios are given herewith. Day. 12345 6789 10 Cetti 13.7 .. 14.2 .. 15 ... 14 16 .. 15.3 Breithaupt 13.5 13 13.5 17 17 16.6 The average ratio for the 6 days on which it was determined for Cetti was 14.7 and the average of 6 fasting days for Breithaupt was 15.1. In general, the quotients remained fairly constant, although there is a minimum of 13 and a maximum of 17 observed in the case of Breithaupt. Commenting on the ratios obtained by Pellizzari & Luciani (4, p. 145), Munk shows that the Italian authors determined only the inorganic and ethereal sulphur and hence their values for total sulphur are somewhat too low. It should also be said, however, that the values for nitrogen are likewise too low,81 and hence the discrepancies tend to compensate. The ratios found in the Florence fast of Succi range during the first 10 days of fasting from 19.5 on the first to 16.2 on the tenth day. Table 209. — Ratio of nitrogen to total sulphur (8) in metabolism experiments without food. Exper- iment num- ber. Subject and date. First day. Second day. Third day. Fourth day. Fifth day. Sixth day. Seventh day. 69.... A.L.L., Dec. 16-19,1904. . 18.33 19.31 17.97 16.43 71 S.A.B., Jan. 7-10, 1905. . 11.25 15.53 17.27 16.04 ... 73 S.A.B., Jan.28-Feb. 1,1905 15.97 17.10 16.20 14.99 14.15 5 75 S.A.B., Mar. 4-10,1905.. 19.59 18.62 17.38 16.11 16. 2( 5 16.27 16.28 77 S.A.B., Apr. 8-11,1905.. 18.49 16.53 16.03 17.30 79 H.E.S., Oct. 13-14,1905.. 14.94 18.47 • • • • .... 80 C.R.Y.,Oct. 27-28,1905.. 20.03 14.52 • • • • ■ • . ■ 81 A.H.M., Nov. 21-22, 1905. . 15.94 15.96 • • • * • • • • 82 H.C.K.,Nov. 24-25,1905.. 16.45 18.71 .... • • ■ • 83 H.R.D.,Dec. 5- 6,1905.. 15.22 19.24 • . • . . • • • 85 N.M.P.,Dec. 9-10,1905.. 18.84 18.81 • • • • 89 D.W., Jan. 10-11,1906.. 14.41 16.80 16.62 17.47 16.97 16.17 15.lt ) 16.27 16.28 Luciani discusses at considerable length the fluctuations in this ratio, but in the light of our present knowledge of sulphur determinations, his discussion is at best inconclusive. The ratios of nitrogen to total sulphur for the Middletown experiments have been computed and are given in table 209. 81 See discussion by Munk (7), p. 118. 26 402 Influence of Inanition on Metabolism. The ratio is highest for the first day of experiment No. 80, 20.03, and lowest for the first day of experiment No. 71, 11.25. The average ratios for all of the experiments remain relatively constant throughout the fasts. It is evident, however, that marked variation may occur in the experiments, even in those with the same subject. For example, in experiment No. 71 the ratios ranged from 11.25 on the first day to 17.27 on the third. It will be remembered that on the first day of this experiment an unusually small amount of nitrogen was excreted and hence the numerator of the fraction is small. Evidently, then, the low excretion of nitrogen was not accompanied by a correspondingly low excretion of sulphur. The average ratio for all of the experiments is 16.78. Further discussion of the significance of this ratio is deferred pending the consideration of the quantities of protein, fats, and carbohydrate katabolized during fasting. Ethereal sulphates. — The work of Baumann,82 on ethereal sulphates in the urine has led to the almost general acceptance of his views on the use of the amount of ethereal sulphates as an index of bacterial decomposition in the intestine. This view has been recently opposed by Folin,83 who implies that the ethereal sulphates have a much more complicated origin than bacterial action. The ethereal sulphur was not determined in the fasting experiments here reported, but in some of the earlier fasts, especially those with Cetti and Breithaupt, the amounts of ethereal sulphates were found. An inspection of the data presented by Munk & Mueller (7) shows that the amount of ethereal sulphates is considerable on all the days of the fast. In one instance, namely, on the ninth day of Cetti's fast, the amount of the ethereal sulphate was nearly one-third of the total sulphuric acid. These writers point out that the amounts they found with Cetti and Breithaupt are ten times greater than those found by Luciani (4) in Succi's Florence fast. The results of Munk & Mueller (7), viewed from the standpoint of Baumann, indicate a bacterial decomposition continuing throughout the fast and, indeed, in very considerable measure. On the other hand, in the observations of E. & 0. Freund (10) on Succi during the Vienna fast, the amounts of ethereal sulphates (expressed as S03), range from 0.20 gram on the first fasting day to 0.058 gram on the twentieth fasting day, there being a gradual decrease as the fast progressed. These quantities are similar to those found by Luciani in the Florence fast, and, viewed from the standpoint of Baumann, indicate a very much lower degree of bacterial fermentation than do the results of Munk & Mueller on the Berlin fasters. Inorganic and ethereal sulphates. — Although the data for computing the ethereal sulphates are lacking in the experiments here reported, the sum of the inorganic and ethereal sulphates has been recorded for all experiments in 82Zeit. f. physiol. Chemie (1879), 3, p. 156. 83Amer. Journ. Physiol. (1905), 13, p. 97. Sulphur in Urine. 403 table 210. These data may be compared to the inorganic and ethereal sulphates of the earlier writers (see table 207). Except in the Vienna fast of Succi reported by E. & 0. Freund (10) and Sadovyen's experiment (2) the amount of inorganic and ethereal sulphates (S03) is not far from 1.5 grams per day in all fasting experiments. Table 210. — Proportions of total sulphur (SOs) excreted as inorganic and ethereal and as neutral sulphur in metabolism experiments without food. Exper- iment number. Subject and duration of experiment. First day. Total. Inor- ganic and ethereal. Neutral. In per cent of total. Inor- ganic and ethereal. Neutral. 75 77 79 80 81 82 83 85 8.A.B., Mar. 4 to 10, 1905 8.A.B., Apr. 8 to 11, 1905 . H.E.S., Oct. 13 to 14, 1905 C.R.Y., Oct. 27 to 28, 1905 A.H.M., Nov. 21 to 22, 1905 H.C.K., Nov. 24 to 25, 1905 H.R.D., Dec. 5 to 6, 1905 N.M.P., Dec. 9 to 10, 1905 D.W., Jan. 10 to 11, 1906 . . . Grams. 1 . 559 1.190 1.355 .969 1.427 1.423 2.173 1 1.507 1.730 Grams. 1.354 .934 1.162 .827 1.295 1.230 1.961 U.219 1.472 Grams. 0.205 .256 .193 .142 .132 .193 .212 i .288 .258 Per cent. 86.8 78.5 85.7 85.3 90.8 86.4 90.2 80.9 85.1 Per cent. 13.2 21.5 14.3 14.7 9.2 13.6 9.8 19.1 14.9 1.482 .593 1.273 .510 0.209 .083 I 85.9 14.1 Exper- iment number. Subject and duration of experiment. Second day. Total. Inor- ganic and ethereal. Neutral. In per cent of total. Inor- ganic and ethereal. Neutral. 75 77 79 80 .... 81 82 83 89 S.A.B., Mar. 4 to 10, 1905 S.A.B., Apr. 8 to 11, 1905 H.E.S., Oct. 13 to 14, 1905 C.R.Y., Oct. 27 to 28, 1905 A.H.M., Nov. 21 to 22, 1905 H.C.K., Nov. 24 to 25, 1905 H.R.D., Dec. 5 to 6, 1905 N.M.P., Dec. 9 to 10, 1905 D.W., Jan. 10 to 11, 1906 A (SO, Grams. 1.669 1.628 1.940 1.710 2.036 1.915 1.755 1.506 2.148 Grams. 1.480 1.322 1.701 1.483 1.891 1.746 1.580 1.331 1.831 Grams. 0.189 .306 .239 .227 .145 .169 .175 .175 .317 Per cent. 88.7 81.2 87.7 86.7 92.9 91.2 90.0 88.4 85.2 Per cent. 11.3 18.8 12.3 13.3 7.1 8.8 10.0 11.6 14.8 1.812 .726 1.596 .639 0.216 .087 I 88.1 11.9 1 Does not include possible amount in urine lost. Neutral sulphur. — It is greatly to be regretted that the nature of neutral sulphur in fasting urines has not been more definitely studied. The differential method of obtaining neutral sulphur presumes so great an accuracy that it is 404 Influence of Inanition on Metabolism. difficult to interpret the results obtained on Succi, Cetti, and Breithaupt with any degree of satisfaction. Indeed in the experiments on Breithaupt, in at least three instances, the amount of inorganic and ethereal sulphur was greater than the total sulphur and it is fair to assume that the determinations on the Berlin fasters were made as accurately as any up to that date. The more recent determinations of E. & 0. Freund (10) on Succi are probably less open to objection, although the methods employed are not given and hence the accuracy of the results is uncertain. Furthermore, in the fasts of Succi at least, various amounts of alkaline waters were consumed. Presumably these contained but small amounts of preformed sulphates, although at times aperient water ( Janos) was taken. The data obtained in the earlier experiments regard- ing the excretion of neutral sulphur are, therefore, extremely unsatisfactory. In connection with some of the experiments here reported, the amounts of neutral sulphur were determined. The data are recorded in tables 210 and 211. For convenience in computing, the " total " and " inorganic and ether- eal" sulphur are also included in the first table, as are also the proportions of the total sulphur represented by the " inorganic and ethereal " and the neutral sulphur. Table 211. — Proportions of total sulphur (SOJ excreted as inorganic and ethereal and as neutral sulphur in metabolism experiments without food. Day of experiment. Experiment number. Inorganic and ethereal. Neutral. Amount. Proportion of total. Third day. Seventh day. . . . 77 75 77 Average . . 75 75 75 Orams. (1.484 ( .594 1.618 1.425 j 1.522 I .610 J 1.520 \ .609 ( 1.477 \ .592 j 1.414 \ .566 Grams. 0.225 ) .090 \ .184 .226 .205 ) .082 f .148 ) .059 f .171 ) .068 \ .139 ) .056 [ Per cent. 13.2 10.2 13.7 11.9 8.9 10.4 8.9 The neutral sulphur (S03) elimination on the first day of the fast varied from 0.132 to 0.288 gram, and the average for the 9 experiments was 0.209 gram. The proportion of total sulphur represented by the weight of neutral sulphur varied from 9.2 to 21.5, averaging for all experiments 14.1 per cent. On the second day of the fast, the weight of neutral sulphur in the urine ranged from 0.145 to 0.317, averaging 0.216 gram, while the proportion of total sulphur represented by the neutral sulphur varied from 7.1 to 18.8 per cent. The Sulphur in Urine. 405 average is 11.9 per cent. Unfortunately, the data for the third day were secured from only one experiment in which the neutral sulphur was 0.225 gram, or 13.2 per cent of the total sulphur eliminated. On the fourth day, the determinations made in two experiments gave 0.184 and 0.226 gram respect- ively. Expressed in per cents of total sulphur, these weights corresponded to 10.2 and 13.7 per cent. On the fifth, sixth, and seventh days, the neutral sul- phur was determined in experiment No. 75 only. The amounts excreted were less than on previous days, being 0.148, 0.171, and 0.139 gram, respectively, these weights corresponding to 8.9, 10.4, and 8.9 per cents of the total sulphur. There is, then, a tendency for the actual weight of neutral sulphur to decrease as the experiment continues, and the per cent of total sulphur represented by the neutral sulphur likewise diminishes. The effect of the bile flow on the neutral sulphur has generally been main- tained, since it was believed that the taurin contributed in large measure to the amount of neutral sulphur found in the urine. Luciani (4) maintained that the secretion of bile continued throughout the 30 days of the Florence fast, since from time to time Succi vomited material stained with bile pigments. In common with all other secretions, however, the bile flow must be distinctly diminished and the results obtained in the Middletown experiments might be taken as indicating that there is a relationship between the bile flow and the amount of neutral sulphur, for, as has been pointed out above, not only the total but the relative amount of neutral sulphur persistently diminishes as the fast progresses. This conclusion is, however, distinctly at variance with that drawn from the recent experiments of Shaffer84 on a woman with a biliary fistula. Judged from the standpoint of Folin's theory of protein metabolism, the variations in the amounts of sulphur excreted indicate that there is scarcely any greater disintegration of tissue protein during fasting, than under normal conditions with food. That this view is in marked contrast to the many physical observations made on the size of the liver and other organs, as well as of the muscles, during fasts no longer than some of these recorded here, would lead to the belief that in these organs the actual structure is not necessarily materi- ally drawn upon during fasting, but that the whole organism becomes deprived of its fluid to a considerable extent and hence diminishes in volume. While in the paper presenting his views of protein metabolism Folin 85 maintained that there was a distinct relation between the neutral sulphur and the endogenous protein katabolism, it is of interest to note that in a subsequent statement8" he says that his more recent researches would indicate some relationship between the food ingested and the neutral sulphur. MAmer. Journ. Physiol. (1906), 17, p. 340. S5Amer. Journ. Physiol. (1905), 13, p. 117. 88 Private communication reported by Shaffer, loc. cit., p. 375. 406 Influence of Inanition on Metabolism. While in Folin's metabolism experiments there was a marked parallelism between the excretion of creatinine and neutral sulphur, in these experiments the total creatinine remains constant during the fast, and the neutral sulphur persistently decreases. The preformed creatinine on the other hand more nearly follows the neutral sulphur elimination. The results here obtained during inanition serve to complicate the matter still further, and there is obviously much research to be done upon the problem of neutral sulphur before definite conclusions can be drawn. PHOSPHORUS. Under normal conditions, with food, material amounts of phosphorus are excreted in the feces as well as in the urine. In fasting experiments, however, and specifically in the experiments under discussion in this report, only the phosphorus excretion of the urine has been adequately studied. The difficulties experienced in the proper separation of fasting feces precludes any discussion of the amount of phosphorus excreted through the feces during fasting. The phosphorus of the urine, under normal conditions when food is given, may originate from several sources : First, preformed phosphates of the food, such as the phosphates of milk; second, phosphorus of the nucleo-proteids of food ; and third, phosphorus already stored in the body, including of course the calcium phosphate of the bones, which might possibly become disintegrated to a certain extent. In fasting urine, especially, this last factor should not be overlooked. Determinations of phosphorus in urine were common in the earlier fasting experiments, and since the method, i. e., volumetric titration with uranium salts, has not been materially modified, the results are much more nearly comparable than is the case with almost any other element involved in katabolism. While there is not complete uniformity on the part of writers in expressing the results of phosphorus determinations, it has been common to express them in the form of phosphorus pentoxide (P205). For the reason set forth in the discussion of sulphur in the preceding section, it has been thought desirable to report the elimination of phosphorus expressed as the element in subsequent experiments. The daily amounts of phosphorus (as P205) elimi- nated in the urine of fasting subjects are presented in table 212. For subse- quent discussion the ratios of nitrogen to phosphorus pentoxide (N/P20B) are given in heavy-faced type in the same table. The phosphorus pentoxide excretion is by no means uniform in the three experiments with Succi. For example, there were but 1.46 grams excreted on the fourth day of fasting in the Naples experiment, while there were 2.54 grams on the corresponding day of the fast in Vienna. On the whole, there was a much larger amount of phosphorus excreted per day at Vienna than during either of the two earlier fasts. In Cetti's experiment, the phosphorus excretion Phosphorus in Urine. 407 was much larger than in any of the experiments with Succi, while Breithaupt, on the other hand, excreted phosphorus in about the same amounts as did Succi. The lowest phosphorus output is found in the case of the subject J. Sohn eliminated extremely large amounts and, singularly enough, the excre- tion increased throughout the whole experiment. Table 212. — Ratio of nitrogen to phosphorus (P2OJ and amount of phosphorus (P2OJ eliminated daily in urine by fasting subjects. Day of fast. Succi. Cetti. Breit- haupt. At Flor- ence. At Na- ples. At Vi- enna. J.l Suhn. Sub- ject I.* Flora Tosca. Keller.5 Last food day . . . 1 2 3 4 5 6 8. 9. 10. Av.ratio^ (for fasting- days only).. Grams. 7.87 1.930 91 051 30 090 6.64 2.120 5.90 2.394 5.18 2.150 53 865 85 601 6.29 1.360 5.86 1.246 6.24 Grams. 4.73 1.90 4.90 1.78 4.64 1.82 4.64 1.95 5.83 1.46 3.74 2.64 3.49 2.47 3.28 2.32 4.18 1.48 4.63 1.49 4.37 1.23 4.37 Grams 5.70 2.98 4.07 2.75 4.19 2.52 4.25 2.54 4.46 2.51 4.85 2.27 4.13 2.13 4.22 2.31 4.19 2.40 4.24 1.68 4.43 Grams. 4.89 2.76 5.22 2.597 4.30 2 . 925 3.99 3.289 4.17 2.974 3.73 2.871 3.79 2 . 667 4.09 2.663 5.17 1.722 5.24 2.065 10.00 .948 4.97 Grams. 4.84 2.69 6.42 1.56 5.25 1.89 5.25 2.53 5.42 2.360 00 19 32 29 5.28 Grams. 40 682 20 091 51 491 03 571 Grams. 6.21 3 . 381 5.37 2.303 5.45 2.268 6.17 2.270 5.54 2 5.052 3 3 4.43 2.434 60 150 4.S6 4.442 6.29 5.20 Grams. 4.53 1.58 3.97 2.22 3.81 2.65 4.41 2.65 4.06 Grams. 5.24 2.670 5.65 1.550 4.58 1.830 4.04 2.654 20 934 50 749 7.23 1.069 8.57 0.713 4.64 1.658 4.32 1.702 4.65 1.461 5.14 Grams. 41 86 55 90 24 44 54 53 3.94 1 Reported by A. Sadovyen (2). 2 Amount for 2 days. "See Note (2). 4 Reported by Schreiber and Waldvogel ; probably as P205. 6 Reported by A. Keller, Zeit. f. physiol. Chemie (1900), 29, p. 165. With the subject of Schreiber and Waldvogel, the excretion is not far from that of Sohn for the corresponding days of the fast. The more recent obser- vations on the fasting girl, Flora Tosca, show a relatively large excretion of phosphoric acid, assuming that her body-weight (which unfortunately was not given) is lower than that of the other fasting subjects. There is as a rule a tendency for the phosphoric acid excretion to increase for a few days after fasting begins and then subsequently to diminish. Some instances of an extremely low excretion are to be observed, namely, on the tenth 408 Influence of Inanition on Metabolism. day of the fast of Cetti and on the seventh day of the fast of Flora Tosca. The amount of phosphorus pentoxide eliminated per day is, in general, not far from 2.2 grams. The determinations of phosphorus in the Middletown experiments were made by two methods, as is shown in the detailed tables; one by titration with uranium salt, the other by fusion using the modified sodium peroxide method of Dubois. Since the titrations were possibly less accurate than the gravimetric deter- minations, the latter only are included in table 213. Table 213. — Ratios of nitrogen to phosphorus (P2OJ and amounts of phosphorus* excreted in urine in metabolism experiments without food. Exper- iment num- ber. Subject and duration of experiment. First day. Second day. Third day. Fourth Fifth day. day. Sixth day. Sev- enth day. 69.... A.L.L., Dec. 16 to 19, 1904.. •{ Oms. 10.78 0.936 Oms. 11.96 1.192 Oms. 14.19 1.060 Oms 12.44 1.04 Gms. 3 Gms. Oms 71 S.A.B., Jan. 7 to 10, 1905.. •{ 7.93 .736 S.33 1.326 6.55 2.000 5.27 2.03 3 .... 73 S.A.B., Jan. 28 to Feb. 1,1905 | 4.39 2.345 6.29 1 . 904 5.96 1.938 4.81 2.15 4.6S ? 2.134 75 S.A.B., Mar. 4 to 10, 1905.. •{ S.55 1.431 5.52 2.255 6.34 2.055 4.83 2.40 5.23 3 2.078 5.19 2.071 4.87 2.081 77 S.A.B., Apr. 8 to 11, 1905.. •{ 4.71 1.871 3.90 2.763 4.41 2.492 4.35 2.63- I 79 H.E.S., Oct. 13 to 14, 1905.. •{ 7.50 1.081 8.62 1.664 • • • 80 C.R.T., Oct. 27 to 28, 1005.. •{ 6.57 1.184 4.11 2.421 81.... A.H.M. , Nov. 21 to 22 1905.. { 6.19 1.472 5.37 2.430 82.... H.C.K. Nov. 24 to 25 1905.. •{ S.43 1.113 7.18 2.000 83.... H.R.D. , Dec. 5 to 6, 1905.. •{ 5.51 2.407 4.84 2.793 .... 85 N.M.P. , Dec. 9 to 10, 1905.. •{ S.79 2 1.257 9.55 1.189 ... 89 D.W., Jan. 10 toll, 1905.. •{ 4.24 2.354 4.61 3.134 Ave •{ 6.97 1.516 6.69 2.089 3 5.82 3 4.82 2.05f 4.96 ! 2.106 5.19 2.071 4.S7 1.9C 19 2. Of (1 1 By fusion method. 2 Does not Include possible amount in urine lost. 8 Omitting experiment No. 69. The total weight of phosphorus (expressed as P205) excreted on the first day of the different fasts varied from 0.736 gram, in experiment No. 71, to 2.407 grams in experiment No. 83, averaging for the first day 1.516 grams. Even with the same subject (S. A. B.) in different experiments, the variations range from 0.736 to 2.345 grams. On the first day of the fast, therefore, there was apparently no uniformity in the excretion of phosphoric acid. Phosphorus in Urine. 409 The variations on the second day are from 1.189 to 3.134 grams, the average for the 12 experiments being 2.089 grams. In general, there is a noticeable increase in the phosphoric acid elimination on the second da}' of the fasts, the only exceptions to this rule being in the case of experiments Nos. 73 and 85. In the 5 fasts which lasted 3 days or more, the phosphoric acid excretion on the third day ranges from 1.06 to 2.492, averaging for all experiments 1.909 grams. Considering the experiments with S. A. B., there is an increase on the third day over the second day in the first two and a decrease in the last two, the average amount for the four experiments being practically the same on the third as on the second day. In the experiments which lasted 4 days, the excretion on the fourth day varied from 1.043 to 2.634 grams. With the subject A. L. L., the phosphoric acid excretion practically reached a minimum, while with the subject S. A. B., there is a definite increase over the third day in all cases. The excretion on the fourth day of experiment No. 77 is not quite as large as on the second. The data secured with S. A. B., for the fifth, sixth, and seventh days of fasting show that the phosphoric acid elimination remains practically constant at a little over 2 grams, a distinct falling off from the average maximum amount on the fourth day. From the summary of the data given above it is evident that to draw conclusions concerning the relative phosphorus excretion during the successive days of fasting is most difficult, for even with the same subject there is no uniformity in the excretion. Thus, in considering the second, third, and fourth days of experiments Nos. 71 and 77, we find there was excreted 1.326, 2.000, and 2.038 grams in experiment No. 71, while in experiment No. 77, there was excreted during the same period 2.763, 2.492, and 2.634 grams. Compared with the earlier experiments the results here obtained show on the whole a noticeably lower phosphorus elimination. The unusually low results obtained in experiment No. 69 can possibly be partially explained by the fact that in the feces of the 3 days with food (experiment No. 70) there was an abnormally high ash and a large per cent of calcium soap, thus indicat- ing a marked disturbance of the ash metabolism. The results do not clearly show in just what manner the phosphorus metabolism was affected. Pending the determinations of the calcium and magnesium in the urine samples for these experiments,87 but few definite conclusions can be drawn. Organic phosphorus in urine. — The contention has been made by several writers w that varying amounts of phosphorus in organic combination exist in 87 It has thus far been impossible to complete these analyses as the determinations were taken up only recently. The results and a discussion of the data will be published later. ^Oertel, Zeit. f. physiol. Chemie (1898), 26, p. 123; A. Keller, Zeit. f. physiol. Chemie (1900), 29, p. 146; Bornstein, Archiv f. die ges. Physiol. (1904), 106, p. 66; Symmers, Journ. Path. Bact. (1905), 10, pp. 159 and 427. 410 Influence of Inanition on Metabolism. urine, which are not determined by titration with uranium salts. On the other hand, LeClerc and Dubois "9 were unable to detect measurable amounts of such organic phosphorus. In the Middletown experiments gravimetric determinations of phosphorus were made in the daily composite samples, while the determinations by titration with uranium acetate were made on the different periods as well as the composite samples for the day. Obviously, the sum of the amounts of phos- phorus found in the different periods should agree with that found in the composite sample. The determinations made in this way serve two purposes — they give an idea as to the proportional distribution of the excretion of phosphorus over the 24 hours, and they furnish a check on the samples for the daily composite and the periods. If the results obtained by the titration method are to be taken as representing the amount of phosphorus excreted in the urine of any given period, it is important that the absence of organic phosphorus in the urine be clearly proved as otherwise the results by titration have little value. Consequently, the com- posite sample for each day's urine was concentrated and ignited with sodium peroxide, and the phosphoric acid determined gravimetrically. The amount of phosphoric acid thus found in a total day's urine was compared with the amount found by titration, the results being recorded in the statistical tables along with the weight, composition, and heat of combustion of urine. An in- spection of these tables shows that, in general, the amount of phosphoric acid as found by titration was a trifle larger, if anything, than that found by fusion, which is contrary to what would be expected if organic phosphorus was present. It might further be properly contended that the very fact that the titration method gives on the whole a larger amount of phosphorus than the fusion method is of itself evidence that the titration method as carried out in these experiments is not sufficiently accurate to detect the presence of the small amounts of organic phosphorus usually found. Furthermore, Keller, whose experiment on himself is of especial interest here since it was made on a fast- ing man (4 days), found amounts of organically combined phosphorus pentox- ide amounting to 0.017, 0.0294, 0.0344, and 0.0573 gram, respectively. The increased organic phosphorus elimination as the fast progressed is, according to Keller, of especial significance in interpreting the role of organic phosphorus in metabolism. In the light of the experience of Folin and LeClerc & Dubois, it seems hardly probable that any appreciable amounts of organic phosphorus occur in the urine, and at least in the experiments here reported, the titration with uranium acetate may be taken as a measure of the total phosphorus. Periodic distribution of phosphorus. — By means of the simple titrations with uranium acetate, determinations of phosphorus in the samples of urine 80 Journ. Amer. Chem. Soc. (1904), 26, p. 1112. Phosphorus in Urine. 411 for the separate periods of the day were readily made. In the series of experi- ments, Nos. 73 to 76, inclusive, these determinations were carried out. The results are presented in table 214. For purposes of comparison, the experi- ments with food are also included. In comparing the results, it should be borne in mind that experiment No. 74 immediately followed experiment No. 73, and experiment No. 76 likewise followed immediately experiment No. 75. Table 214. -Periodic distribution of phosphorus 1 (as P2OJ in urine in metabolism experiments with and without food. Ex- peri- ment num- ber. Subject and date. 7 a. m. to 1 p. m. 1 p. m. to 7 p.m. Total for first 12 hours. Total for last 12 hours. Total for 24 hours. Am't. Pro- portion of total for 24 hours. Am't. Pro- portion of total for 24 hours. 73 75 74 76 Experiments without food. 3.A.B., Jan. 28 to 29, 1905. Jan. 29 to 30, 1905. Jan. 30 to 31, 1905. Jan. 31 to Feb. 1, 1905 Orams. 2 0^505 .490 .518 .575 Orams. 2 o'.6l8 .508 .690 .692 Grams. 2 1.123 .998 1.207 1.268 Per ct. 54^9 51.5 63.5 57.9 Orams, 2 .923 .940 .695 .920 Per ct. 45! i 48.5 36.5 42.1 Orams. 2 2.046 1.938 1.902 2.188 Feb. lto2, 1905... Total, 4 days 2.088 2.508 4.596 3.478 8.074 Average per day S. A. B., Mar. 4 to 5,1905 . Mar. 5 to 6, 1905 . Mar. 6 to 7, 1905 . Mar. 7 to 8, 1905 . Mar. 8 to 9, 1905 . Mar. 9 to 10, 1905 . Mar. 10 to 11, 1905 . Total, 7 days .522 .627 1.149 56.9 .870 43.1 2.018 .383 .591 .453 .549 .549 .561 .522 .394 .802 .635 .798 .665 .578 .618 .778 1.392 1.088 1.347 1.214 1.139 1.141 53.6 60.5 51.7 55.7 55.4 56.0 58.1 .674 .909 1.016 '.978 .896 46.4 39.5 48.3 44.3 44.6 44.0 41.9 1.452 2.301 2.104 3 2.416 2.192 2.035 3 1.963 3.608 4.490 8.099 44.473 14.463 Experiments with food. S.A.B., Feb. 2 to 3, 1905.. Feb. 3 to 4, 1905.. Feb. 4 to 5, 1905.. Total, 3 days .515 .641 1.157 56.0 4.895 544.0 2.066 .461 .315 .238 .392 .384 .308 .853 .699 .546 61.3 60.9 47.8 .537 38.7 39.1 52.2 1.390 3 1.148 3 1.141 1.014 1.084 2.098 .... 6.537 .... 3.679 S.A.B.,Mar. 11 to 12, 1905- Mar. 12 to 13, 1905. Mar. 13 to 14, 1905. Total, 3 days .338 .361 .699 57.0 .537 643.0 1.226 .375 .113 .053 .200 .109 .137 .575 .222 .190 70.0 59.1 24.1 .247 !598 30.0 40.9 75.9 .822 3 .375 .789 .541 .446 .987 7.845 .... 1.986 Average per day .180 .149 .329 49.7 .423 650.3 .662 'Titration method. 2 Not determined. 8 Total in composite urine. 4 For 5 days. 3 Percentage by 8 For 1 day. 7 For 2 days. difference. 412 Influence of Inanition on Metabolism. The proportion of the total phosphoric acid excreted in the first twelve hours is, on the average, not far from 57 per cent, although considerable differences, even in the fasting experiments, are found. For example, the variations in experiment No. 73 are from 51.5 to 63.5 per cent. In the food experiments, much larger variations appear, the most noticeable being that on the last day of experiment No. 76, where but 24.1 per cent of the total phosphoric acid was excreted during the period from 7 a. m. to 7 p. m. Thus it is seen that, in general, the larger proportion of the phosphorus is excreted in the day period and, indeed, on reference to table 198, it will be noticed that the proportions thus excreted correspond very closely, in general, to the proportions of total nitrogen excreted during this period. Ratio of nitrogen to phosphorus in the urine. — The disintegration of phos- phorized material in the body during fasting presumably follows the general course of protein katabolism, save for the drafts upon the phosphatic material of the bones, and hence the ratio of nitrogen to phosphorus in the excretion, when compared with the ratio of nitrogen to phosphorus in the various tissues of the body is of interest. According to Munk (7, p. 159) there are about 6.8 " parts of nitrogen to each part of phosphorus pentoxide in flesh, and 6.4 parts of nitrogen to one of phosphorus pentoxide in the liver. Accordingly it is to be expected that, during fasting, the ratio of nitrogen to phosphorus pent- oxide would be not far from 6.6. Eatios lower than this would indicate a draft upon phosphatic material of low nitrogen content such as bone. In experiments with Succi, the ratios were computed by Luciani (4), but for the Florence fast in table 212, given above, the values for total nitrogen as corrected by Munk are used in computing the ratios. The ratio on the first day in the experiment at Florence is 7.87 and there is a general tendency for the ratios to diminish as the fast progresses. In the fast at Naples the ratio is more nearly constant, and the initial ratio, 4.90, is much lower than that of the Florence fast. The Vienna fast shows singularly constant ratios after the first day, the minimum being 4.07 (second fasting day) and the maximum 4.85 (sixth fasting day). The excretion of so small an amount of phosphorus pentoxide on the tenth day of Cetti's fast resulted in a very high ratio (10.00). Aside from this day, the average ratio for the nine days of fasting is 4.4. In the other fasts the ratio is in practically all cases much below 6.6. The peculiarity in the ratios for the sixth and seventh days of the fast of Flora Tosca may possibly be explained by the fact that on the sixth day the subject defecated after taking an aperient, while on the seventh she had considerable muscular exercise. "Forster, Zeit. f. Biol. (1873), 9, p. 363, and ibid. (1876), 12, p. 466, gives the relation between the total nitrogen and phosphoric anhydride in dog flesh as 7.2. Chlorine in Urine. 413 In the Middletown experiments, wide variations in these ratios are observed. For example, in experiment No. 69, the first in which the ratio could be obtained, it rose to 14.19 on the third day. The lowest ratio observed on any day was that on the second day of experiment No. 77, 3.90. In experiment No. 69 there was an abnormally low phosphoric acid elimi- nation, and hence the ratio is unusually high. In averaging all of the experi- ments, these high ratios have been taken into consideration only on the first two days. While they do not have a great influence in the averages of the first two days of the fast, since they are there averaged with 11 other experiments, on the third and fourth days, they would very materially raise the average value of the ratio. There is, as is to be expected, a slight tendency for the ratio to fall off as the fast progresses, yet the ratios are on the whole con- siderably higher than those found with the other fasting experiments given in table 212." The ratio apparently is constant after the third day of fast. While ratios as low as are commonly observed during a fast point to the probable disintegration of the bones, it is only in those experiments where the calcium and magnesium output has been determined that the complete data for this deduction are present. Unfortunately the quantitative determination of the earthy bases in the urine of the Middletown experiments have not as yet been made. The ratios obtained by Brugsch (12) on the twenty-third to thirtieth days of fasting indicate that the draft upon the phosphorus of the skeleton may in prolonged fasting be very small. It is, moreover, still to be questioned whether data regarding the phosphorus excretion, even when supplemented by determi- nations of calcium and magnesium, will ever permit correct estimates of the apportionment of the phosphorus katabolism among skeleton, nucleins, and lecithins.02 CHLORINE. The elimination of chlorine in feces is normally very small, and hence dur- ing inanition it can properly be said that all chlorine is eliminated through the urine. The chlorine excreted during fasting may arise from the previous food, the soluble chlorides of which are rapidly excreted, the excess of chlorides in the fluids of the body, and the chlorine " combined " with the flesh katabolized.93 Since muscle contains normally but 0.04 per cent chlorine, the amounts of protein usually katabolized during inanition can result in the liberation of but a small amount of chlorine. 91 Brugsch (12) found an average ratio of 5.9: 1 during the last 8 days of the 30-day fast made by Succi in Hamburg. This ratio is considerably higher than those found on the average for the third to seventh days of the Middletown fasts. 93Edlefsen, Deutsches Arch. f. klin. Medizin. (1881), 29, p. 409. "Belli, Zeit. f. Biol. (1903), 45, p. 203. 414 Influence of Inanition on Metabolism. The method of determining chlorine in the urine has undergone no marked modifications in a number of years past and hence the determinations made by different observers may be compared with reasonable accuracy. While it has been the custom in many instances to report chlorine in terms of sodium chloride,94 for purposes of comparison the determinations in the earlier fasts are here expressed in terms of chlorine. Only the first 10 days of each of the three fasts by Succi are here given. Table 215. — Amounts of chlorine eliminated in urine daily by fasting subjects. Day of fast. Succi. At Flor- ence. At Naples. At Vienna. Cetti. Breit- haupt. J.1 Sohn. Flora Tosca. Last food day 1 2 3 4. 5 6 7 8 9 10 Grams. 2 6 . 322 1.350 .539 1.155 .848 .817 .840 .800 .736 .550 .513 Gramsfi 7.45 7.68 4.72 3.30 .78 .91 .62 .61 .42 .73 .77 Grams. 4 9.029 3.212 1.551 1.479 1.182 1.291 1.109 1.115 1.212 .873 Grams. 5.432 1.606 2.303 1.7 1.548 1.396 1.088 .95 .814 1.104 .62 Grams. 5.55 3.92 1.1 .85 .75 .44 .35 Grama. 624 011 49 01 Grams.4 7.472 4.143 2.341 2.402 1-53.608 1.513 1.466 1.997 Grams. 7.51 2.99 1.73 3.66 I 1.90 \ .38 .30 .32 1.15 1.32 1.07 1 Reported by Sadovyen. 2 Average of 6 days before fast began. s Reported as " chlorides," not converted to chlorine. 4 Given by the investigators as NaCl, but con\erted to chlorine for purposes of com- parison. 6 Amount for 2 days. In the fast of Succi at Naples, the authors (6) express the chlorine as " chlorides." The extremely small amounts of chlorine excreted after the first four days of the fast, however, make it appear questionable whether the determinations are not actually on a basis of chlorine rather than chlorides. The results as given in the second column of table 215 are, however, tran- scribed directly from the records of Ajello and Solaro. Obviously, if these results are in terms of sodium chloride, the chlorine corresponding to these amounts would be considerably less and the chlorine excretion much lower than in any other experiment with which they are compared, save on the last two days of the experiment with Breithaupt and the fifth, sixth, and seventh days of the experiment with Flora Tosca. 84 On the 3 single fasting days reported by Pettenkofer & Voit (Zeit. f. Biol. (1866), 2, p. 479), the sodium chloride in the urine amounted to 14.6, 13.2, and 8.56 grams, respectively. The determinations of sodium chloride in these experi- ments did not represent those during complete fasting since the subject consumed a considerable amount of salt in connection with a small amount of meat extract. Ranke (Archiv Anat. u. Physiol. (1862), p. 338) reports the sodium chloride excre- tion in two 24-hour fasting experiments as 11.0 and 5.3 grams, respectively. Chlorine in Urine. 415 The table above shows that the chlorine elimination on the last food day is invariably large and that on the first fasting day there is usually a marked diminution in the amount. On the first day of the Vienna fast, however, a very large quantity of chlorine, 9.029 grams, was excreted, while singularly enough, with the same subject, Succi, in the Florence fast, the smallest amount observed on the first day of any fast was recorded. In general, immediately after the first fasting day the chlorine output reaches a low level and there is a slight tendency for the amount to diminish as the fast progresses. No regularity, however, is observed in any of the experiments. Especially noticeable are the marked variations in the relative chlorine excretion in the three fasts of Succi. The amounts in the Vienna fasts are persistently higher than those in the Florence fast. Daiber* found Succi's urine almost chlorine-free on the twen- tieth day of the Zurich fast. It is, furthermore, to be noted that during Succi's fasts, he consumed at varying times different amounts of mineral water (Eiolo) containing a considerable percentage of chlorine; thus, on the third, sixth, and seventh fasting days of the Florence fast, this mineral water was used. Luciani, in discussing these results, points out that after the first few days of fasting, the body has lost such considerable amounts of stored sodium chloride that the amounts absorbed from the mineral water are not immediately excreted but retained, and subsequently eliminated. Table 216. — Chlorine excreted in urine in metabolism experiments without food. Exper- iment num- ber. 73... 75... 77... 79... 80... 81... 82... 83... 85... 89... Subject and duration of experiment. First day. S.A.B., S.A.B., S.A.B., H.E.S., C.R.T., A.H.M., H.C.K., H.R.D., N.M.P., D.W., Jan. 28 Mar. 4 Apr. 8 Oct. 13 Oct. 27 Nov. 21 Nov. 24 Dec. 5 Dec. 9 Jan. 10 to Feb to 10, to 11, to 14, to 28, to 22, to 25, to 6, to 10, to 11, . 1, 1905. 1905 1905 1905.... 1905 1905 1905 1905 1905 1906 Ave. of above experiments. Gms. 1.630 1.447 5.294 2.917 8.898 3.880 3.451 .517 24.582 5.857 3.847 Second day. Third day. Gms. 0.466 1.338 1.671 3.622 4.028 2.794 6.714 .628 1.461 1.805 Gms. 0.159 616 919 2.453 iO.898 Fourth day. Gms. 0.356 245 647 0.416 Fifth day. Gms. 0.408 0.408 Sixth day. Gms 0.387 Seventh day. Gms. 0A20 0.387 0.420 1 Not determined. 2 Does not include possible amount in urine lost. The data in table 215 show that the chlorine excretion during the fasting is markedly different for different individuals and, indeed, during different exper- iments with the same individual. The conditions which determine the varia- tions in this chlorine excretion are not clear. The chlorine excreted in the urine was determined in many of the Middle- town fasting experiments. The results are given in table 216. * Loc. cit. 416 Influence of Inanition on Metabolism. The data given above are of interest because they give some idea of the relation of the chlorine output to the length of the period of inanition. The variations in the amount of chlorine excreted on the first day of the several fasts are large, ranging from the very small excretion in experiment No. 83, namely, 0.517 gram to 8.898 grams in experiment No. 80. The average of all the experiments for the first day is 3.847 grams. These wide variations on the first day indicate clearly that the excretion must be influenced to a very great degree by the quantity of soluble chlorides taken with the food and drink of the preceding day. On the second day of the fast the variations are nearly as extreme, ranging from 0.466 gram in experiment No. 73, to 6.714 grams in experiment No. 82. The average for the second day of all the experiments is 2.453 grams. In three of the experiments, Nos. 79, 82, and 83, the excretion of chlorine on the second day was greater than that of the first day. Considering only the experiments with S. A. B., the second day invariably showed a diminished excretion as compared with the first, although there is no regularity in the per cent of decrease. Thus in experiment No. 73, there is a decrease of about 1.2 grams in the excretion, in experiment No. 75, 0.11 gram, and in experi- ment No. 77, in which there was an unusually large excretion on the first day, i. e., 5.294 grams, the excretion on the second day fell to 1.671 grams. Three experiments in which the chlorine was determined continued for three days or more. The excretion on the third day averaged 0.898 gram. A decrease in the first two experiments was observed, but there was an actual increase in the excretion on the third day of experiment No. 77 over that of the second day. The continued high output of chlorine in experiment No. 77 is difficult to understand except on the supposition that this subject must excrete soluble chlorides rather slowly. From the large excretion on the first day of the experiment, it is apparent that the body contained much more chlorine at the beginning of this fast than at the beginning of either of those preceding. The absolute rise on the third day of the experiment is difficult to explain. The abnormalities in the urinary constituents during experiment No. 77 have frequently been pointed out. Not until the fourth day of fasting is reached is there anything approxi- mating constancy in the excretion of chlorine for this subject. The variations here are from 0.245 to 0.647 gram, the average excretion being 0.416 gram. On the fifth day of experiment No. 75, the chlorine output was not determined. For experiment No. 73, it amounted to 0.408 gram, a little less than the average for the fourth day. In the one experiment in which the determinations were made for the sixth and seventh fasting days, the chlorine elimination is practically constant. Chlorine in Urine. 417 It thus appears that during fasting the excretion of chlorine may be con- siderable on the first two days, a smaller amount appearing usually on the second day. There is a marked decrease in the total excretion on the third day. On the fourth day the elimination decreases to about 0.4 gram and remains fairly constant for the three days following. The results obtained in these experiments are also of interest in discussing the question of the existence of an excessive amount of chlorine in the body over and above what is needed in the system. According to Munk's (7) con- ception, man has become accustomed to much larger amounts of sodium chloride than have actually been needed, and hence there is in the body an excess which is rapidly eliminated on fasting. A more recent statement of this view has been given by Magnus-Levy.85 An examination of the data given in table 215 shows that in nearly all the earlier experiments considerable amounts of chlorine were excreted on the first days of the fast. In general, the excretion diminished considerably after the second day. In practically all cases, the excretion remained above 0.7 gram. The marked exceptions to this in previously published work on fasting are the fifth and sixth days of the fast of Breithaupt and the fifth, sixth, and seventh days of the fast of Flora Tosca. On the basis of the results obtained on Cetti and Breithaupt, as well as the earlier results on Succi, Munk contends that there is a considerable accumulation of chlorine in the body which may amount to 10 or 15 grams. This is excreted during the first days of fasting, after which the excretion becomes constant. The data for the chlorine elimination on the later days of the fast with Succi indicate that about one-half a gram of sodium chloride was excreted per day. The data for the Middletown experiments, on the other hand, show markedly different results. While in the series of 2-day experiments, the chlorine excre- tion is perfectly comparable to that of the earlier experiments with one or two exceptions, in the long fasts with S. A. B., especially experiments Nos. 73 and 75, the rapid excretion of any chloride accumulated in the body is not observed. Indeed, even during the seven day fast, although to be sure the data for the fifth day are missing, the total elimination of chlorine is (allowing 0.450 as the elimination for the fifth day) less than 5 grams. Extremely low results, also, are observed in experiment No. 73. Singularly enough, in ex- periment No. 77, the chlorine excretion is more nearly in accord with the results of earlier observations. A possible explanation of the extremely small output of chlorine in experi- ments Nos. 73, 75, and 77 may be found in the fact that the subject of the experiments, S. A. B., used food largely of a vegetable nature, and practically no table salt. It was the opinion of the assistants who weighed and prepared 95 Physiologie des Stoffwechsels. Sonder-Abdruck aus; von Noorden, Handbuch der Pathologie des Stoffwechsels (1906), p. 451. 27 418 Influence of Inanition on Metabolism. his food, that in the latter part of the first nitrogen metabolism experiment, i. e., the days immediately preceding experiment No. 77, he consumed con- siderably more table salt than formerly. Unfortunately, weighings of the salt consumed were not made and hence this impression lacks scientific verification. An inspection of the menus of the food consumed during the nitrogen metabolism experiments would tend to verify this observation, since the subject used a little meat, which was contrary to his earlier custom. The subject of experiment No. 83 reports, on the other hand, that it is his custom to use average amounts of table salt, but the term " average " is at best vague. It is hardly probable that with the large amount of food eaten by the subject, S. A. B., during all the periods between the fasting experiments, there was not a considerable amount of sodium chloride consumed with the food, although a comparatively small amount of cooked food was commonly taken. Unfortunately, data regarding the retention of sodium chloride after the fast are lacking. Taking the data as presented in table 216, the results of experiments Nos. 73, 75 and 83, and indeed those of No. 77, certainly do not point toward the excretion of any considerable amount of excess chlorine accumulated in the body prior to the fast. A striking exception to the general trend of the experiments is seen in experiment No. 83, in which the chlorine excretion on the two days of the fast was 0.517 and 0.628 gram respectively. The analyses were repeated several times and the possibility of error seems to be eliminated. Proportion of sodium chloride in ash. — Since in all probability soluble chlo- rides consumed with the food on the day immediately preceding the fasting period are rapidly excreted and thereby appear in large measure on the first days of the fast, it is of interest to note the proportion of total ash which is represented by the chlorides. While Munk has shown (7) that the chlorine in the urine is to a certain extent combined with bases other than sodium, the amount of chloride other than sodium chloride is relatively small and for pur- poses of comparison we may assume that all the chlorine that is excreted is combined with sodium. The amounts of sodium chloride calculated as ex- creted in the urine are recorded in table 217, and in this table the proportion of the total ash represented by sodium chloride is likewise recorded. The largest excretion of sodium chloride is commonly found on the days when there was the largest excretion of ash. Thus, on the first day of experi- ment No. 80 when there were 18.93 grams of total ash, it is computed that, there were 14.69 grams of sodium chloride. Similarly on the first days of experiments Nos. 85 and 89, the large amounts of total ash are accompanied by large amounts of sodium chloride. The lowest percentages observed in the first two days of fasting are those found in experiment No. 83. Ash other than sodium chloride. — The very considerable fluctuations in the amounts of sodium chloride excreted in the urine from day to day make a comparison of the ash other than sodium chloride on different days of interest, Chlorine in Urine. 419 and while somewhat extraneous to the general matter under discussion in this particular section, the quantities of ash other than sodium chloride have been computed and placed in table 217. These amounts of ash are seen to be much more uniform than the total ash as determined, and from the regularity of the sulphur and phosphorus eliminations, it is to be expected that the ash other than sodium chloride would remain practically constant throughout the experi- ment. The second day of experiment No. 77 shows the largest amount of this portion of the total ash, amounting to 9.22 grams. In general, the excretion of ash other than sodium chloride is not far from 5 grams per day. Table 217. — Proportion of sodium chloride in ash of urine in metabolism experi- ments without food. Experi- ment number. 73. 75. 77. 79. 80. 81. 82. 83. 85. 89. Subject and duration of experiment. S. A. B.: Jan. Jan. Jan. Jan. Feb. S. A. B.: Mar. Mar. Mar. Mar. Mar. Mar. Mar. S. A. B.: Apr. Apr. Apr. Apr. H. E. S.: Oct. Oct. C. R. Y.: Oct. Oct. A. H. M.: Nov. Nov. H. C. K.: Nov. Nov. H. R. D.: Dec. Dec. N. M. P.: 28-29, 1905 29-30, 1905 30-31, 1905 31-Feb. 1, 1905. 1- 2, 1905 4- 5, 1905 5- 6, 1905 6- 7, 1905 7- 8, 1905 8- 9, 1005 9-10, 1905 10-11, 1905 8- 9, 1905 9-10, 1905 10-11, 1905 11-12, 1905 13-14, 14-15, 1905 1905 27-28, 1905 28-29, 1905 21-22, 1905 22-23, 1905 24-25, 1905 25-26, 1905 5- 6, 1905 6- 7, 1905 D. W. Dec. 9-10, 1905 Dec. 10-11, 1905 Jan. 9-10, 1906 Jan. 10-11, 1906 Ash. Grams. 9.96 6.23 5.74 6.09 6.02 6.03 7.54 6.44 7.80 7.48 5.85 5.27 13.88 11.98 10.55 9.14 8.73 11.17 18.93 12.52 12.61 11.22 9.67 16.33 8.19 8.73 20.36 12.47 5.57 13.34 7.45 Sodium chloride. Ash other Proportion than [ of sodium sodium chloride chloride. in ash. Grams. 2.69 0.77 0.26 0.59 0.67 2.39 2.21 1.02 0.41 i _ 0.64 0.69 8.74 2.76 3.17 1.07 4.82 5.98 14.69 6.65 6.40 4.61 5.69 11.08 0.85 1.04 '0.22 7.56 2.41 9.67 2.98 Grams. 7.27 5.46 5.48 5.50 5.35 3.64 33 42 39 5.21 4.58 5.14 9.22 7.38 8.07 3.91 5.19 4.24 5.87 6.21 6.61 3.98 5.25 7.34 7.69 0.14 4.91 3.16 3.67 4.47 Per cent. 27.0 12.4 4.5 9.7 11.1 39.6 29.3 15.8 5.3 10.9 13.1 63.0 23.0 30.0 11.7 55.2 53.5 77.6 53.1 50.8 41.1 58.8 67.9 10.4 11.9 61.1 60.6 43.3 72.5 40.0 1 Not determined. 'Calculated. See p. 243. 420 Influence of Inanition on Metabolism. WATER OUTPUT. No other compound exists in the body in such large proportions as does water. The fluctuations in the amount of water in the body during inanition, therefore, demand special study. During fasting experiments as ordinarily conducted, drinking-water is allowed, and hence the relations between the amounts of drinking-water and the amounts of water excreted either through the kidneys or lungs and skin are likewise of importance. In discussing the volume of urine during fasting, considerable emphasis was laid upon the relation between the amount of water consumed and the volume of urine, and specifically the water in urine.96 It was there pointed out that while, in general, the ingestion of large volumes of water was accompanied by large volumes of water in the urine, at the same time there were marked exceptions to this general rule. These exceptions imply that the water of urine is an excretory product governed by definite laws, the nature of which is as yet but little understood. Since, in nearly all experiments heretofore made with fasting men, an accurate measure of the total water excretion is lacking, the data obtained in these experiments will, it is hoped, be of value in explaining the nature of the apportionment to the kidneys, lungs, and skin of the water excreted by the body. Since drinking-water was allowed ad libitum in all the experiments here reported, the actual loss of water to the body was less than the loss occurring during complete abstinence from water as well as food. The amounts of drinking-water consumed by the various subjects were widely different in different experiments. In certain experiments, the subjects were especially requested to consume large quantities of water, but in the series of 2-day fasting experiments, the subjects as a rule consumed only as much as was actually desired, the amount taken rarely exceeding 1000 grams. On one day but 115.1 grams were consumed. The data for the amounts of water consumed are given in table 193. While under ordinary conditions when food is eaten, some water may be furnished the tissues by the oxidation of the organic hydrogen of the food, in fasting experiments where no food is ingested the water of oxidation of organic hydrogen must be formed from the organic hydrogen of the body material broken down. The proportions of water thus formed may be more properly studied after the consideration of the katabolism of body material. Thus, it is important to recognize that while the water output consists for the most part of preformed water, varying amounts resulting from the oxidation of organic hydrogen are present in the water excreted. Excretion of water from the body may take place in three ways ; through the lungs and skin, i. e., water of respiration and perspiration ; through the urine ; and through the feces. M For discussion of the relation of the water in urine to water consumed, see page 348 and table 193 (p. 355). Water Output. 421 water of feces. From the discussion of feces (see page 337), it can readily be seen that the isolation of so-called " fasting feces," namely, feces, the organic matter of which may be said to be derived from the disintegration of body material, is extremely difficult. The excretion of water in feces, however, can be readily determined. Unfortunately, careful determinations of water in feces were not made in some of the Middletown experiments, since it was apparent that the feces had resulted from the ingestion of food prior to the fast. Errors are thereby undoubtedly introduced, since the water of the feces was actually excreted. In the food experiments, it has been assumed that certain amounts of water were excreted in this way, and the preformed water lost has been calculated on this assumption. While normal feces contain not far from 70 to 80 per cent of water, during fasting, the feces are usually retained for a considerable period of time, become hard and pilular, and consequently have a much smaller water content. The exigencies of experimenting in one instance called for the removal of fecal matter by means of an enema, and hence in this connection it was difficult to determine the water actually excreted with the feces. WATER OF URINE. Large amounts of water are excreted in the urine during fasting. In a preceding section the ratio of water of urine to water of drink has been discussed at length (see p. 348). WATER OF RESPIRATION AND PERSPIRATION. Considerable quantities of water are evaporated from the lungs and skin and it becomes necessary in accurate metabolism experiments to measure the amounts lost. Since each gram of water vaporized requires the absorption of 0.592 calorie of heat, the influence of the water of respiration and perspiration on the heat output is very marked, and consequently for the accurate determina- tion of the heat production careful measurement of the water of respiration and perspiration is necessary. Furthermore, an accurate record of the water of respiration and perspiration is essential for determining the amount of pre- formed water excreted, and the water resulting from the oxidation of organic material in the body. The difficulties attending the collection and analysis of the respiratory gases have precluded, in practically all of the earlier experiments on fasting, a study of the amounts of water vaporized from the lungs and skin. Although the amount of carbon dioxide has been frequently determined, unusual difficulties attend the accurate measurement of loss of water from the body. The fluctua- tions in the relative humidity inside the respiration chamber, hygroscopic 422 Influence of Inanition on Metabolism. nature of clothing, furniture, etc., and difficulties attending the accurate record of body-weight, all tend to vitiate the accuracy of the indirect determination of water-vapor as heretofore made. To be sure, a fairly close approximation can be obtained by means of the so-called " insensible loss," which can be computed from the changes in body-weight from the beginning to the end of the day, the weights of urine and feces, and the weights of drinking-water, provided there is a determination of the amount of carbon dioxide exhaled. In the experiments on Cetti and Breithaupt reported by Munk (7), the respiratory quotient and consequently the carbon dioxide elimination was determined on different days but only for short periods each day. From these determinations, the authors calcu- lated the total carbon dioxide elimination for the 24 hours. Thus, the data were obtained for computing the insensible loss in these experiments. In the 10 days of Cetti's experiment, it was computed that there were 8016 grams of water of respiration and perspiration, an average of 802 grams per day.97 From the determinations of the amount of protein katabolized, and like- wise from the estimations of fat oxidized, on the basis of the respiratory ex- periments made with the Zuntz apparatus, the authors apportioned the amount of water vaporized in terms of water of oxidation of organic material and water lost from the body itself. Similar results were reported for Breithaupt in a 6 days' fast in which there were 4199 grams (700 grams per day) of water of respiration and perspiration. The errors involved in computing the daily absorption of oxygen and elimination of carbon dioxide from respiration experiments lasting but 10 to 20 minutes are obvious, and the errors which affect the measurement of the carbon dioxide elimination likewise affect the estimations of the water of respiration and perspiration. Nevertheless, the results obtained by this method are by no means as inaccurate as one might suppose. By means of the more accurate determination of the total elimination of carbon dioxide with the Tigerstedt apparatus, the water of respiration and perspiration was computed in the experiments on J. A. (9). The objections to the determination of carbon dioxide in short periods did not here obtain, and although the subject left the respiration chamber each day for a period of two hours, the estimation for the remainder of the day was unquestionably extremely accurate. When computed for the 24 hours of the day, the result represents the probable carbon dioxide output of a fasting man under the con- ditions of muscular activity obtaining while in the respiration chamber. An important factor in the indirect determination of the amount of water of respiration and perspiration is, however, the change in body-weight. Since the subject J. A. was outside the respiration chamber at least 2 hours of the " (7), p. 114. Water of Bespiration. 423 day and during that time his muscular activity was much greater than in the period inside the chamber, the losses in body-weight are in all probability too large for the accurate computation of the water vaporized. From the loss in body-weight and the carbon dioxide output, the water of respiration and per- spiration was computed. The amounts thus found were for the 5 days 1058, 931, 662, 1329, and 803 grams. A much more desirable method of obtaining the water of respiration and perspiration is that in which the subjects are inclosed in a respiration chamber and the water-vapor in the ventilating air-current accurately measured. The difficulties attending this determination are much greater than at first sight would appear. The bedding and other articles inside the chamber are prone to absorb or give off water and, indeed, in considerable amounts. To make a complete determination of the amount of water-vapor eliminated from the lungs and skin of the subject, therefore, necessitates, in addition to the measurement of the water-vapor in the air, a record of the changes in weight of bedding and other articles in the chamber. It is of special interest to note that the respiration chamber of Pettenkofer 88 was used first to study the metabolism of fasting man. The experiments were made by Eanke." From the carbon dioxide output and the changes in body- weight, Eanke computed the water of respiration and perspiration in three 24- hour fasting experiments to be 609, 1080, and 537 grams. The room tempera- tures were 19.5°, 25.4°, and 16.4°, respectively. On the last day no water was consumed. Direct determinations of the water-vapor output of fasting man were first reported by Pettenkofer & Voit.100 These writers early recognized the diffi- culties attending water determination, and the observations regarding changes in the weights of bedding, etc., are fully in accord with those of present day experiments. The experiments of these investigators lasted 24 hours. During this period the water determinations showed an elimination of 828.9 grams of water of respiration and perspiration in one experiment and 814.1 grams in another. In the third fasting experiment, during which, however, the subject engaged in considerable muscular work, the water of respiration and perspiration amounted to 1778.5 grams. While unquestionably the results obtained by Pettenkofer & Voit are the resultant of a number of errors which are more or less com- pensating, the fact remains that these observers recognized the desirability of determining the water-vapor directly instead of computing the insensible loss, and they also planned their experiments to cover 24 hours. The next successful attempt to determine directly the water-vapor eliminated by fasting man was that of Sadovyen (2). Two experiments were made, one "•Annalen der Chemie u. Pharm. (1862), n, Supplementband, p. 1. wArchiv Anat. u. Physiol. (1862), p. 340. 100Zeitsclir f. Physiol. (1866), 2, p. 478. 424 Influence of Inanition on Metabolism. of two days' duration in which no drinking-water was taken, and the other of four days during which water was consumed. The subject remained in the respiration chamber the larger part of each 24 hours. Usually the periods during which the experimental observations were recorded were of 9 to 10 hours' duration, and from the results obtained during these periods, the amounts for 24 hours were computed. On the 2 days of the first experiment the water of respiration and perspiration amounted to 961 and 831 grams, respectively, and for the 4 days of the second experiment, the amounts were 730, 709, 597, and 713 grams. Although the subject spent a portion of the day outside the respiration chamber with somewhat greater muscular activity, these results are not liable to the same criticism raised regarding the obser- vations made on J. A., since the loss of body-weight does not enter into the computation for 24 hours. The Pashutin respiration apparatus was likewise used in an experiment in which the water of respiration and perspiration was determined by Likhachev.101 In this experiment the subject remained 26 hours and 10 minutes inside the chamber, and from the observations thus made, the amounts for 24 hours were computed, thus marking a distinct advance in accuracy over the experiments of Sadovyen. In the 1-day fasting experiment, the water- vapor output was 664 grams, or 10.68 grams per kilo of body-weight. No records are given of changes in weight of the furniture and other articles inside the chamber. In the four earlier fasting experiments made in this laboratory and pre- viously reported,102 the water of respiration and perspiration was determined. No attempt was made in these experiments to allow for any changes in weight of the bedding and furniture. The amounts of water of respiration and perspiration for the different days were: experiment No. 36 (1 day), 768 grams; experiment No. 39 (1 day), 822 grams; experiment No. 42 (1 day), 842 grams; experiment No. 51 (2 days), 1018 and 895 grams, respectively. The particular type of respiration apparatus used in these experiments is especially adapted for the measurement of the quantity of water in the ventilating air-current. Furthermore, since in all but one (experiment No. 59) of the experiments here reported, special precautions were taken to secure reliable weighings of the bedding and other articles in the chamber of the apparatus, corrections for variations in the moisture content of these articles could be applied to the other measurements of water-vapor and the water of respiration and perspiration be accurately measured. Without the use of a respiration chamber and accurate balance for noting the gain or loss in weight of the articles in the chamber, accurate measurements of the water of respira- tion and perspiration are impossible. 101 The production of heat hy healthy man in a condition of comparative rest. A. A. Likhachev, Inaug. Dissertation (Russian), 1893, St. Petersburg. 102 U. S. Dept. Agr., Office of Expt. Sta. Bui. 136. Water of Eespiration. 425 The results obtained in the fasting experiments here reported are given for each day in table 218, the detailed determinations for the 2- and 3-hour periods being given in the statistical tables for the different experiments. Table 218. — Water of respiration and perspiration in without food. metabolism experiments Expe imen num ber. r- t Subject and duration of experiment. First day. Second Third day. i day. Fourth day. Fifth day. Sixth day. Seventh day. 59.. . B.F.D., Dec. 18-20, 1903 Chris. 982 Grns. Grns. 952 943 Grns. Gins. Grns. Grns. 68.. . A.L.L., Apr. 27-28, 1904 745 761 .... .... .... .... 69.. A.L.L., Dec. 16-19, 1904 738 898 795 728 .... .... ■ • . . 71.. . S.A.B., Jan. 7-10, 1905 745 665 568 518 .... .... «... 73.. . S.A.B., Jan. 28-Feb.l, 1905. 684 636 602 569 543 .... .... 75.. . S.A.B., Mar. 4-10, 1905 650 642 658 596 579 542 543 77.. . S.A.B., Apr. 8-11, 1905 670 618 638 627 .... .... 79.. 667 704 . • . .... • • • . .... 80.. . C.R.T., Oct. 27-28, 1905 927 1061 ... . . . .... .... .... 81.. . A.H.M., Nov. 21-22, 1905 609 671 ■ • ■ ■ • . .... 82.. . H.C.K., Nov. 24-25, 1905 842 940 ... ■ . . 1 .... .... 83.. . H.R.D., Dec. 5- 6, 1905 685 672 ... . . . • • • . .... .... 85.. . N.M.P., Dec. 9-10, 1905 776 813 . . . ... .... .... .... 89.. . D.W., Jan. 10-11, 1906 820 803 .... 753 774 701 607 561 542 543 The figures in table 218 show that the variations between different days with different subjects are very considerable, ranging from 518 grams on the fourth day of experiment No. 71 to 1061 grams on the second day of experiment No. 80. It is to be noted that these subjects were performing no muscular work other than that incident to the ordinary habits of life inside the chamber. Considering all the experiments, the variations on the first day of the fast range from 609 to 982 grams, the average being 753 grams. On the second day of the fast, the range is from 618 to 1061 grams, the average for all the experiments being a little more than that on the first day, namely, 774 grams. The smaller number of experiments which lasted three or more days show a range on the third day from 568 to 943 grams, the average being 701 grams. As the fast progresses, the average amount of water of respiration and perspi- ration per 24 hours gradually diminishes, but for the last 3 days of fast, as here recorded, the amount is nearly constant. Considering the experiments with the same subject, the agreement between Nos. 68 and 69 is fairly close, the second day of experiment No. 69 being an exception. On this day there were 160 grams more water given off than on the day preceding. The extended series of experiments with S. A. B. show that for the most part on the first day without food, the amount of water was fairly constant, ranging 426 Influence of Inanition on Metabolism. from 650 to 745 grams. On the second day there is much less variation, the minimum and maximum being 618 and 665 grams. The difference between the lowest and the highest amounts on the third day is 90 grams, and on the fourth day about 100 grams. In general, as the fast progresses, there is a diminution in the amount of water vaporized. This is especially noticeable in the longer fasts with S. A. B. The absolute amounts of water excreted in the respiration and perspiration are on the whole considerably smaller than those usually found with resting subjects. For example, in the numerous rest experiments with food made in this laboratory and previously reported, the lowest amount of water of respira- tion and perspiration recorded is 697 grams 103 with E. 0. and the highest amount is 1212 grams with the same subject. The average of all the rest experiments gives a water excretion from lungs and skin of 935 grams.1M The possible factors affecting the elimination of water-vapor are the absolute amount of water in the body, including the drinking-water, the relative humidity of the air in the respiration chamber, and the muscular activity of the subject. Ratios between the water of respiration and perspiration and amounts of drinking-water. — The ratios between the water in the urine and the water con- sumed have received special discussion on page 348. It was there seen that while, in general, large amounts of drinking-water were accompanied by large volumes of urine, there is no definite ratio between the volume of urine and the volume of water consumed. Especially is this the case in experiments in which very small volumes of water were taken. The effect of the ingestion of large quantities of water on the elimination of water-vapor can be studied by means of the data obtained in these experiments, since the amount of water consumed each day, as well as the water of respiration and perspiration, was accurately measured. The ratios between the water of respiration and perspiration (table 218) and the amounts of drinking-water, as recorded in column a of table 193, have, therefore, been computed. The results show widely varying ratios, the lowest being that for the third day of experiment No. 73, namely, 0.218, i. e., for every 1000 grams of water consumed there were but 218 grams of water of respiration and perspira- tion. The largest ratio is that of the first day of experiment No. 89, namely, 7.121. On this day the subject consumed an unusually small amount of water, i. e., 115.1 grams, while the water of respiration and perspiration was 820 grams. The ratios in the experiments with S. A. B. are more nearly constant. In experiment No. 71, the ratios for the 4 days are 0.618, 0.337, 0.245, and 0.348, 10S A special experiment, in which abnormal ventilation conditions obtained, is reported in U. S. Dept. Agr., Office of Expt. Sta. Bui. 175. In this experiment the water vaporized from the body was 267 grams. 104 U. S. Dept. Agr., Office of Expt. Sta. Bui. 136, p. 137. \Vater of Eespiration. 427 averaging 0.358. In experiment No. 73, the ratios are 0.329, 0.232, 0.218, 0.291, and 0.505, averaging 0.286. In experiment No. 75, the ratios are remarkably constant, i. e., 0.330, 0.371, 0.311, 0.312, 0.366, 0.318, and 0.321, averaging 0.331. The ratios for experiment No. 77 are somewhat larger, i. e., 0.327, 0.400, 0.606, and 0.651, averaging 0.454. With the series of experi- ments with S. A. B., therefore, it would appear that there is a possible ratio between the water of respiration and perspiration and the amount of drinking- water. Nevertheless, it must be borne in mind that only on one day in all the experiments with S. A. B. did the drinking-water consumed fall below 1000 grams. That there can be no definite ratio can readily be seen from the fact that on the first day of experiment No. 73 when the subject drank 2082 grams of water, the water of respiration and perspiration was 684 grams, while on the second day, with a very considerable increase in drinking-water (total 2747 grams), the water from the lungs and skin was actually lower, i. e., 636 grams. Furthermore, in experiment No. 71 on the third day, the amount of water consumed was nearly twice that on the first day, while the amount of water in respiration and perspiration was nearly 200 grams less. In the shorter experiments, Nos. 79, 81, and 85, there was also in each instance a decrease in the amount of water consumed on the second day over the first, accompanied by an actual increase in the water of respiration and perspiration. Considering all the experiments seven show an increase in the water of vapor- ization on the second day. In general, the water of respiration and perspiration remains fairly constant on the different days of the same experiment, there being a slight though per- sistent diminution as the experiment proceeds. Frequently there is a parallel diminution in the amount of drinking-water, but in certain instances, especi- ally in the two cited above, the contrary is true. Since, therefore, there is such relative constancy in the water of respiration and perspiration and such wide variations in the amount of water consumed, it is obvious that the ratio between the water vaporized from the body and the water consumed must show marked fluctuations. It is difficult, therefore, to trace any relationship between the amount of drinking-water consumed and the elimination of water-vapor. As a matter of fact, since all of the experiments here compared were made under condi- tions of rest, what is recorded as water of respiration and perspiration is practi- cally confined to the water-vapor leaving the lungs and skin. The absence of any apparent influence of the amount of drinking-water on the water of respiration and perspiration in these fasting experiments is in accord with the observations of Laschtschenko,105 who made a number of experi- ments in a Pettenkofer respiration apparatus to study the influence of the in- 105Archiv f. Hygiene (1898), 33, p. 145. 428 Influence of Inanition on Metabolism. gestion of large amounts of water on the water-vapor output of man. The experiments did not include observations during a period of inanition. The figures in tables 193 and 218 permit a comparison of the water of respiration and perspiration with the water in the urine, although no ratios between these two factors have been computed and tabulated. As is the case with the water of respiration and perspiration, the water in the urine tends to diminish as the fast progresses, and yet in experiments Nos. 69, 71, and 75 the contrary is true. Furthermore, on the second day of experiment No. 73, on which the largest amount of water in the urine (2928 grams) was voided, the total water-vapor output was but 636 grams, while with the same subject on the first day of experiment No. 71, where the water in the urine was about one-third of that on the second day of experiment No. 73, the water of respira- tion and perspiration was more than 100 grams larger. No relation, therefore, appears between the amount of water in the urine and the amount of water- vapor eliminated. Influence of the water content of the body. — As will be seen from the dis- cussion on page 467, there is material loss of preformed water from the body as the fast progresses, hence it might be contended that the restricted elimina- tion of water-vapor may partly be accounted for by the lowering of the absolute amount of water present in the body. On the other hand, the actual amount of preformed water lost from the body is but a small proportion of the total water in the body and the percentage of water present in the body is hardly affected by the losses in fasts as short as are these under discussion. It does not seem reasonable, therefore, to conclude that the relatively small losses of preformed water from day to day can influence appreciably the diminution in the output of water-vapor noted as the fast progresses. Influence of variations in relative humidity. — As the air comes in contact with the moistened mucous membrane of the mouth, nose, throat, and lungs, water is rapidly evaporated from these tracts and the exhaled air has com- monly been assumed to be saturated with water-vapor at the temperature of the body. Air is inspired at the relative humidity of the air in the chamber. The lower the humidity, the greater the amount of water-vapor taken up by the air, as it passes through the lungs. Similarly, as the air comes in contact with the surface of the body, the lower the humidity the greater the amount of water-vapor taken up. We should then, naturally, expect to find that the amounts of water given off by the body per day would in the majority of instances be affected by the relative humidity of the air, although the influence of the relative humidity of the air might or might not be so great as to obliterate any other factors. An examination of the total amount of water remaining in the chamber at the end of each period, as given in the detailed tables in connection with these Water of Kespiratiox. 429 experiments, shows that as a rule the relative humidity of the air inside the chamber becomes lower and lower as the fast progresses. The percentage relative humidity is determined in large measure by the rate of ventilation of the chamber. The more rapid the rate of ventilation, the greater the amount of air withdrawn and completely deprived of its moisture by passing through concentrated sulphuric acid, and hence the larger the volume of dry air returned to the chamber. While in the 2-hour periods it is more than probable that the rate of speed of the electric motor and consequently the volume of air passing through the chamber may vary considerably, it is highly improbable that in the course of 24 hours any marked variations in the amounts of air passing through the chamber are to be observed. In the earlier series of experiments made with the apparatus in a very different form, a mechanical air pump was used and a large number of records were obtained to show the total ventilation per 24 hours. Inasmuch as the same electric motor, shafting, fittings, etc., aside from substituting a rotary blower for the mechanical air pump, are now used, the conditions are not different now than they were before. At that time all the air passing through the chamber was first caused to pass through a large gas meter and thus a record of the total ventilation was obtained. The total ventilation for a large number of days rarely altered more than 3 or 4 per cent from day to day.106 It seems probable that the rate of ventilation stays reasonably constant. Under these circumstances, then, any fall in the relative humidity must be ascribed to a diminished loss of water from the body through the lungs and skin and the figures given in table 218 show that this is actually the case. Since, then, there is a diminishing loss of water of respiration and perspiration from the body, even with a markedly diminished relative humidity, it seems clear that at least in experiments with fasting men some factor other than relative humidity determines the loss of water of respiration and perspiration. Influence of muscular activity. — The only remaining factor that seems in any way connected with the formation of water of respiration and perspiration is muscular activity. The frequent observations that excessive muscular activity results in profuse sensible perspiration justify the assumption that even minor muscular activity would cause an increase in the insensible perspi- ration over that commonly occurring during rest. It might be expected, furthermore, that the muscular activity would determine to a considerable extent the degree of insensible as well as sensible perspiration. The muscular activity in experiments of this nature may be measured relatively by three methods: (1) The record of the muscular movements of the subject obtained from the diary, notes of assistants, etc.; (2) the carbon dioxide production, which is an approximate index of muscular activity; and 106 See table 120, pp. 273-300, U. S. Dept. Agr., Office of Expt. Sta. Bui. 136, in which the ventilation per 24 hours is given for experiments from 35 to 55 inclusive. 430 Influence of Inanition on Metabolism. (3) the heat production, which varies directly with the muscular activity. By comparing the estimated amount of muscular activity (derived by the above methods) with the quantities of water-vapor in respiration and perspira- tion, any relationships existing between them may readily be observed. As the result of observations in a large number of experiments on muscular work,107 it has been found that excessive muscular work produces an enormous increase in the water vaporized from the lungs and skin. It has been definitely shown that when sensible perspiration appears, there is a noticeable increase in the water-vapor output accompanying muscular work. On the other hand, when no extraneous muscular exercise is taken, it is difficult to demonstrate the influence of minor muscular activity on the water-vapor output. In all the experiments here reported, the subjects were practically at rest, with the single exception of the second period of experiment No. 71 when the subject rode a bicycle ergometer for 10 minutes. There was no visible perspiration, however, in this short period of exercise. The marked differences in the elimi- nation of water-vapor, therefore, must be explained by some factors other than marked muscular activity, for the daily routine of the subjects was not unlike, as is seen by comparing the records of body movements. However, that there were marked differences in the sum total of bodily activities, not only in experiments with different subjects but also on different days of the same experiment, is shown in the discussion regarding muscular activity. A com- parison of the relative amounts of muscular activity there estimated with the amounts of water-vapor leaving the lungs and skin shows that there is a dis- tinct relation between even the minor differences in muscular activity as es- timated and the actual water-vapor output. The problem is, however, not as simple as it first appears, for on the assumption that even minor differences in muscular activity produce wide variations in the total quantity of water vaporized, we should expect to find a much smaller vaporization of water during the night period of all the experi- ments, when the subjects were asleep, than during the day period. This is not invariably the case as is shown in the discussion in the next section. The periodic elimination of water-vapor. — Opportunity was offered in the experiments here reported to study the periodicity of the output of water-vapor during the experimental day. The experimental periods were all of 2 hours' duration. Since unusual care was taken in the measurement of the water-vapor and comparatively small changes in the weight of the bed, bedding, etc., occurred in a majority of the experiments, the results give an accurate measure of the water-vapor output for the different periods of the day. The methods of applying the corrections for the changes in weight of the bed and bedding have been explained previously. The results for both the fasting and the food experiments are given in detail in table 219. In this table is recorded the 107 U. S. Dept. Agr., Office of Expt. Sta. Bui. 136. Water of Eespiration. 431 Table 219. — Elimination of water of respiration and perspiration during different periods of the day in metabolism experiments with and without food. Fasting experiment*. No. 69, Dec. 16-17, 1904 Dec. 17-18, 1904 Dec. 18-19,1904 Dec. 19-20, 1904 No. 71, Jan. 7- 8,1905 Jan. 8- 9, 1905 Jan. 9-10, 1905 Jan. 10-11, 1905 No. 73, Jan. 28-29, 1905 Jan. 29-30, 1905 Jan. 30-31, 1905 Jan. 31-Feb. 1,1905. Feb. 1- 2, 1905 No. 75, Mar. Mar. Mar. Mar. Mar. Mar. 4- 5, 1905. 5- 6. 1905. 6- 7, 1905. 7- 8, 1905. 8- 9, 1905. 9-10, 19115. Mar. 10-11, 1905. No. 77, Apr. 8- 9, 1905. Apr. 9-10, 1905. Apr. 10-11, 1905. Apr. 11-12, 1905. No. 79, Oct. 13-14, 1905. Oct. 14-15, 1905. No. 80, Oct. 27-28, 1905. Oct. 28-29, 1905. No. 81, Nov. 21-32, 1905. Nov. 22-23, 1905. No. 82, Nov. 24-25, 1905. Nov. 25-26, 1905. No. 83, Dec. 5- 6, 1905. Dec. 6- 7, 1905. No. 85, Dec. 9-10. 1905. Dec. 10-11, 1905. No. 89, Jan. 10-11,1906. Jan. 11-12, 1906. Average of above : First day of fast.. Second day Third day Fourth day Fifth day Sixth day Seventh day All days Food experinu No. 70, Dec. 20-21, Dec. 21-22, Dec. 22-23, No. 72, Jan. 11-12, No. 74, Feb. 2- 3, Feb. 3- 4, Feb. 4- 5, No. 76, Mar. 11-12. Mar. 12-13. Mar. 13-14, A v. of above food First day Second day. . Third day.... ■nts. 1904.... 1904.... 1904.... 1905. . . . 1905. . . . 1905. . . . 1905. . . . 1905.... 1905. . . . 1905. . . . exp'ts : 7a.m. to 9a.m 9 to 11 a.m. All days Gins. 75.2 84.9 80.2 70.8 81.6 82.5 53.0 51.6 72.9 59.0 59.0 52.5 51.0 72.6 74.2 68.0 52.8 56.4 54.1 55.1 74.0 77.8 68.4 50.2 53.0 55.2 67.2 92.3 56.1 56.5 99.4 110.3 68.3 78.8 76.8 73.7 68.0 65.7 75.0 75.7 86.8 53.0 49.9 40.5 48.7 57.3 51.0 58.7 Gms. 59.8 68.9 67.4 55.6 81.3 67.9 51.0 42.6 67.6 67.3 54.8 49.7 47.2 62.4 58.8 58.2 47.6 45.9 48.5 44.3 60.4 48.5 47.5 51.4 55.4 57.7 47.5 77.4 63.4 64.3 87.0 85.8 57.4 52.5 6S.4 63.4 64.3 61.2 11 a.m. tol p.m. Gms. 61.0 64.2 69.4 61.8 69.9 54.5 45.0 43.5 59.0 54.9 53.4 53.3 44.1 55.9 61.7 45.4 51.1 50.4 46.8 46.5 56.6 49.7 50.3 49.1 55.6 63.9 84.3 88.0 54.6 55.1 59.8 78.3 57.9 59.9 66.5 65.6 66.1 68.5 1 to 3 3 to 5 p.m. p.m. Gms. 61.0 66.9 63.9 59.3 64.3 58.3 48.7 43.8 56.4 63.3 52.2 48.1 45.4 51.2 60.6 58.1 52.1 49.4 44.8 47.3 57.7 50.1 54.7 50.8 55.6 57.9 74.3 95.3 51.8 50.9 72.8 75.8 53.2 52.3 59.6 61.2 71.0 64.5 Gms. 58.6 69.3 65.3 62.0 70.1 50.5 48.3 43.3 54.6 62.6 52.4 45.8 40.2 47.4 50.0 53.4 48.3 47.2 39.0 44.2 54.7 47.2 49.1 47.1 54.8 58.8 75.7 89.5 51.6 49.7 73.8 74.4 51.7 54.6 59.6 59.9 64.6 61.9 5 to 7 7 to 9 p.m. p.m. Gms. 64.9 71.0 67.1 61.9 66.7 65.7 52.1 48.4 58.5 55.2 51.6 46.6 46.3 47.5 49.9 63.8 51.2 45.7 43.0 43.1 54.3 48.3 60.1 59.4 65.7 59.8 76.1 91.9 53.6 58.2 67.7 74.0 60.5 54.0 65.4 68.7 71.1 73.7 Gms. 68.4 67.7 65.1 61.2 70.5 52.7 46.9 42.9 63.9 53.5 52.7 50.3 48.2 48.9 50.4 54.2 48.8 48.3 44.6 38.0 53.7 54.8 51.3 50.8 52.9 61.2 70.1 83.0 48.3 56.4 70.5 84.2 5S.6 58.1 73.6 70.1 72.5 79.9 9toll p.m. Gms. 55.6 65.2 69.7 59.1 61.1 49.2 48.1 39.7 51.4 51.4 46.6 46.2 49.9 54.0 52.6 54.3 62.9 49.3 44.7 45.1 45.4 68.3 52.2 57.2 54.4 61.5 71.0 83.2 43.6 57.6 05. 3 80.2 52.8 53.6 61.3 68.9 73.3 69.3 lto 3 a.m. Gms. 60.6 85.0 60.3 60.5 55.3 49.1 44.3 41.6 54.3 48.1 46.6 46.6 41.7 51.3 49.7 51.2 50.3 45.4 43.4 45.4 63.5 63.1 43.6 63.0 54.1 59.8 73.3 95.3 46>6 55.3 64.9 73.9 53.3 55.4 63.8 68.0 69.2 69.9 3 to 5 a.m. Gms. 60.3 89.2 70.0 58.0 50.7 48.9 44.7 43.3 53.6 50.2 42.7 44.2 43.3 50.6 53.6 54.4 46.4 47.4 45.3 42.4 47.7 50.9 53.6 54.7 63.0 61.0 95.3 93.8 45.6 55.8 50.4 57.8 61.2 51.8 60.0 72.9 63.5 61.7 Gms. 67.2 88.7 73.8 61.5 33.4 44.6 43.5 40.6 51.3 47.6 47.6 43.9 41.4 53.7 48.2 54.1 48.9 49.4 43.6 47.1 57.5 61.3 45.0 50.9 54.2 54.2 88.6 85.0 45.3 54.7 63.6 69.5 53.4 48.1 59.9 67.9 68.0 64.2 5 to 7 a.m. Gms. 65.8 77.4 52.6 56.4 50.0 61.4 43.1 37.5 61.7 44.0 44.0 43.1 44.7 54.6 52.4 52.8 45.8 44.6 44.4 44.4 55.3 47.9 52.9 52.0 58.2 63.2 103.7 86.9 49.1 56.5 67.0 77.4 66.3 52.9 61.9 73.2 68.2 62.4 72.1 64.6 62.3 60.7 69.8 60.9 60.9 56.6 58.2 58.5 58.0 75.9 64.5 62.9 61.3 59.9 63.4 64.3 63.6 as. 4 63.3 60.3 65.5 55.8 52 5 55.5 53.7 54.9 54.0 63.2 49.2 53.9 52.8 55.6 49.4 51.3 50.8 49.3 53.5 50.8 51.0 50.4 49.3 49.2 53.7 46.5 47.3 47.4 43.7 46.0 48.3 49.6 43.6 45.3 45.4 54.1 48.5 46.8 44.8 39.0 43.0 44.5 44.7 43.4 45.3 43.6 55.1 44.3 46.5 58.1 47.3 44.2 43.1 38.0 45.1 45.4 42.4 47.1 68.4 59.5 57.5 55.8 58.3 58.1 66.2 56.1 56.3 55.6 61.8 61.3 49.1 47.0 44.7 44.4 44.4 56.3 66.8 73.8 79.6 67.0 71.3 76.9 65.1 73.1 70.5 64.6 81.9 69.9 71.0 90.2 97.9 65.9 101.1 105.0 66.7 SS.3 86.9 72.5 95.2 103.9 90.9 95.8 110.5 70.6 83.9 90.1 51.9 44.0 53.8 46.3 42.0 46.2 40.2 42.3 43.6 39.5 53.3 48.4 45.3 50.6 43.3 42.3 59.0 49.8 44.9 51.3 45.4 44.7 52.6 49.9 49.4 49.6 41.7 44.1 51.2 42.8 43.2 39.7 41.7 40.8 45.8 41.7 41.6 39.8 40.3 40.6 50.1 46.5 46.0 65.4 49.6 53.5 55.9 50.0 53.6 53.0 50.4 54.6 48.6 50.5 51.7 54.7 50.8 56.7 63.1 52.5 50.1 48.6 46.1 48.1 47.5 46.5 46.8 46.8 43.4 47.6 64.2 73.6 81.1 43.3 43.2 43.2 39.5 46.9 45.0 45.0 58.8 55.7 64.7 55.3 55.9 57.0 54.3 54.4 57.5 58.3 57.6 66.0 53.5 59.2 66.4 53.5 63.5 66.3 54.1 64.5 68.6 52.8 61.2 60.1 50.8 61.0 64.3 57.0 61.3 66.3 49.2 55.9 59.4 69.6 56.0 55.3 57.4 56.1 60.4 61.6 57.5 57.9 61.1 64.3 49.4 54.0 55.3 63.5 432 Influence of Inanition on Metabolism. elimination of water-vapor for each period on 38 days of fasting experiments. The larger number of observations were, naturally, on the first and second days. The average excretion of water-vapor for each period on the different days of the fasts is given at the foot of each column and the average excretion for the period in all the fasts is given as a grand average beneath the column for each period. The results for food experiments are also shown in order that any possible effect on this output of water-vapor due to the ingestion of food may be considered. The first period of the day from 7 a. m. to 9 a. m. is characterized by the largest water-vapor output, and during this period the fluctuations as the fast progresses are especially noticeable. On the second day of fasting the highest average water-vapor output is noted, namely, 75.9 grams, while on the fourth, fifth, sixth, and seventh days, the average amounts are practically constant. During the period from 9 a. m. to 11 a. m. there is a decrease of 18 grams between the average water-vapor output on the first and second days of fast and the average amounts on the sixth and seventh days. The decrease as the fast progresses is quite regular after the second day, although the difference between the second and third days is rather greater than that between the third and fourth. Considering the period from 11 a. m. to 1 p. m., it will be seen that the elimination is slightly higher on the second day than on the first, but that on the third day it is reduced by about 10 grams. There is then a gradual falling off after the second day. Similar differences between the first and second days of the fast are observed with practically all the remaining periods of the day, the most noticeable exceptions being the periods from 9 to 11 p. m. and 11 p. m. to 1 a. m., in which there is an increase on the second day of about 5 grams. In general, with all the periods there is a diminution of about 10 grams between the second and third days of fasting. The largest individual water-vapor output noted in any experiment is that of the second day of experiment No. 82 from 7 a. m. to 9 a. m., 110.2 grams, and the lowest is on the first day of experiment No. 71 from 3 a. m. to 5 a. m., 33.4 grams. This corresponds to about 17 grams per hour. The average general distribution of the water elimination throughout the day as shown by the grand averages in table 219 indicates that the greatest output of water-vapor, 68.4 grams, occurs during the period from 7 to 9 in the morning. The lowest output occurs between 3 a. m. and 5 a. m., namely, 55.6 grams. The excessive output during the first period of the day may be explained in two ways: first, the extra muscular effort attendant upon the work of rising in the morning, arranging the furniture, weighing of subject and bedding, and the general activity at the beginning of the experimental day; second, the moisture condensed in the bed clothing over night may be given off during Water of Kespiration. 433 this particular period in greater amounts than in any other period. This latter factor may likewise partially explain the tendency for the water output to be less during the night, since there may be an accumulation of water in the bedding. Excluding the period from 7 a. m. to 9 a. m., the average amounts of water- vapor eliminated from the lungs and skin per 2-hour period remain singularly constant, varying from 59.5 in the period 9 a. m. to 11 a. m. to 55.6 from 3 a. m. to 5 a. m., a difference of less than 7 per cent. On the whole, the greatest water elimination takes place during the waking hours and the smallest during sleep. In the recent study of the periodic elimination of water-vapor by Wolpert & Peters,108 a much greater variation in the water vaporized is shown per hour than in the averages of these experiments. This, however, is to be expected since their observations were confined to but three experiments. In their experiments the attempt was made to obtain the correction for the changes m weight of the clothing and bedding at the end of each experimental period, which was of 4 hours' duration. To accomplish this, the subject of the experiment removed the clothes and weighed the bed, bedding, and clothing at the end of each period. While, theoretically, this procedure is well designed to give accurate data regarding the water-vapor output, practically, the extra- neous muscular activity of dressing and undressing and weighing the clothing is an abnormal one in experiments in which the water-vapor output during rest is to be studied. Furthermore, during the period when the clothing is removed, water is rapidly vaporized from the body as well as from the clothing, and hence there is a vaporization of water much greater than would normally occur with the subject clothed and resting quietly. It is thus seen that the two series of experiments can hardly be compared, although it may readily be contended that the results of the experiments here reported represent more nearly what would be expected to be the water-vapor output of the resting man. This is especially true of the first day of fasting. The factors entering into the later days of fasting obviously would affect the elimination of water-vapor to such a degree that the averages of all these experiments would be distinctly lower than what could be considered a normal for the resting man. Consider- ing only the first day of fasting, the water-vapor output is still noticeably greater during the first period of the day and, aside from this period, it is somewhat larger per hour during the day time than that during the night. Eecognizing the well-known effect of activity on the water-vapor output, it is reasonable to suppose that in these fasting experiments, the large output during the first period may in part at least be accounted for by the extraneous muscular effort. Experiments for the study of the true water-vapor output of resting man should be so designed as to insure regularity of muscular 1MArchiv f. Hygiene (1906), 25, p. 299. 28 434 INFLUENCE OF INANITION ON METABOLISM. activity ;md conditions which would otherwise obtain during the ordinary rest experiment. The differences between the output during the day time and that of the night are, it is true, not sufficiently great to warrant an assertion that there is a positive increase in the water vaporized from the actual body surface of the man during the day, but making due allowance for the vaporization from the bed clothing, which was in these experiments weighed but once each 24 hours, the evidence still points towards a slightly greater elimination of water- vapor during the day period. The increased respiration rate of the day, thereby causing a greater ventilation of the lungs and a greater vaporization of water, would certainly imply an increased water- vapor output during this period. Influence of the ingestion of food. — In the few food experiments which are here reported, the data are also given for the comparison of the output of water-vapor during the different periods of the day. While the number of individual experimental days from which the data are drawn is less than half of those of the fasting experiments, the number is sufficient to average and give general information regarding the distribution of the water-vapor output for the day. The differences are much less than in the fasting experi- ments. A close examination of the table shows that the water of respiration and perspiration on the second and third days of experiment No. 70 was abnormally high in practically all periods. On these two days the subject was evidently in a slightly febrile condition as an examination of the body tempera- ture recorded on page 314 will show. While the data are too meager to draw definite conclusions, they may indicate that slight increases in body tempera- ture affect in a marked degree the elimination of water-vapor, and from the results upon these two days, it would appear that the greatest factor, other than perhaps excessive muscular activity, in determining the water-vapor output of resting man is body temperature. It must be borne in mind, that in no instance in these experiments were the quantities of food appreciably more than enough for maintenance. The food was, generally, in the form of milk and similar products given in small amounts and at frequent periods through- out the day. A comparison between the food experiments and the fasting experiments may also be made by noting the variations in the water-vapor elimination in the food experiment immediately following a fast. Thus, in comparing experi- ments Nos. 71 and 72, 73 and 74, and 75 and 76, such an inspection shows that, although the water-vapor output is by no means as large on the first day with food as on the first day of fast, it is in general somewhat greater than on tbe last day of fast. Thus, on the seventh day of experiment No. 75, the water-vapor output during the period from 7 a. m. to 9 a. m. was 55.1 grams and on the next day, namely, the first day of experiment No. 76, the output was 57.2 grams. On the seventh day of fasting for the second period it was Water of Kespiration. 435 44.3 grams, and for the same period on the first day in experiment No. 76 it was 50.1 grams. A similar comparison holds for practically all the periods of the experiment, so that there is a slight positive increase in the elimination of water-vapor accompanying the ingestion of these small amounts of food. This may well be ascribed to the slight additional internal muscular activity called for on the ingestion of food.109 A portion of the increase may also result from the rise in the respiration rate and the increased muscular activity as a whole indicated by the change in pulse rate. Proportions of water-vapor eliminated from the lungs and shin. — The two chief opportunities for the vaporization of water are from the lungs and from the skin. Air taken into the lungs containing a varying amount of water- vapor is expelled saturated with water-vapor at the temperature of the body. The volume of air thus taken into the lungs and its relative humidity thus determines the total amount of water-vapor leaving the body in the expired air. The evaporation of water from the skin is not so directly measurable, since the quantity thus vaporized may show wide variations. The air in con- tact with the skin is certainly not saturated with water-vapor at the external temperature of the body. Furthermore, the clothing and the interlying air- spaces produce, so to speak, an artificial atmospheric environment. These air-spaces may contain widely varying amounts of water-vapor according to the thickness of the layer of air, the movements of the subject and the tempera- ture of the air as well as the external temperature of the body. Many attempts have been made to measure directly the water-vapor expelled from the lungs. Such measurements, save for experiments of short duration, are impracticable. The data obtained in the Middletown experiments give the total water of respiration and perspiration for each 2-hour period and for the day. By an indirect method of calculation, it is possible to apportion these amounts in such a manner as to estimate the amount exhaled from the lungs and thus show how the vaporization of water is divided between the lungs and skin. With this form of respiration apparatus, no direct measure of the actual venti- lation of the lungs is at hand. Since, however, the amount of oxygen absorbed is accurately determined, it is possible to form a reasonably accurate estimate of the total ventilation of the lungs from the knowledge of the oxygen con- sumption. Zuntz uo and his associates have found in a brilliant series of experiments that for every cubic centimeter of oxygen consumed by a man at rest, 21 cc. of air are inspired. Using this ratio, therefore, it is possible to compute the total ventilation of the lungs by multiplying the total oxygen consumption by 21. 109 The marked difference in muscular activity in experiments Nos. 69 and 70 explains the one exception to the above comparisons. u0H6henklima und Bergwanderungen in ihrer Wirkung auf den Menschen, N. Zuntz, A. Loewy, Franz Muller, W. Caspari (1906), Berlin, p. 380. 436 Influence of Inanition on Metabolism. From the data recording the water determinations of the air residual in the chamber and the temperature of the air, the relative humidity is obtained and it is therefore possible to compute the total amount of water-vapor in the air taken into the lungs. Assuming that the air leaves the lungs saturated with water-vapor at the body temperature of 37°, the total amount of vapor in the expired air may thus be computed. Deducting the amount in the air inspired from that in the air expired, the water-vapor taken up by the air as it passes through the lungs is readily obtained. The data are presented in table 220 herewith, in which the first column shows the total ventilation of the lungs, the second, the water in the inspired air, the third, the water in expired air, and the fourth the water eliminated from the lungs. If this last amount be deducted from the total water of respiration and perspiration, the water eliminated from the skin is obtained. This is recorded in column e. In the last two columns of the table the pro- portions of total water vaporized from the lungs and from the skin are given. It will be seen that this method of computation involves two assumptions — first, that the total ventilation of the lungs is 21 times the volume of oxygen absorbed, and second, that the air leaving the lungs is saturated with water- vapor. Eegarding the first of these assumptions, it is probably true that with certain individuals this factor may be wrong, but it is highly probable that in the large number of experiments here averaged the factor is reasonably correct. Eegarding the second factor Eubner m has shown that with forced respiration during muscular work, the air leaving the mouth and nose may not be satu- rated with water-vapor at the temperature of the body. Eecently Lesageus has, by means of a respiratory hygrometer, computed the water-content of the expired air from the dew point obtained by means of his apparatus. With ordinary respiration the tension of the water-vapor is found to be 36.9 mm. Since the expired air has a temperature of 36.5°, corresponding to a maximum tension of 43.1 mm., the author contends that the air is not saturated with water-vapor. While, therefore, this second assumption may be erroneous, for the want of more accurate data, it seems undesirable to make any correction for these values. If Lesage is correct, the result will be to lower the estimated amounts of water-vapor eliminated from the lungs. The amount of water-vapor eliminated from the lungs is in most instances somewhat less than that from the skin. In some experiments, notably in experiment No. 80, it is but 30 per cent of the total water elimination, while in other experiments, for example, food experiment No. 74, it is over 50 per cent. Marked differences are noted between different subjects, especially in the longer experiments. In the series of 2-day experiments, these differences, aside from experiment No. 80, practically disappear. 111 See discussion by Magnus-Levy, Physiologie des Stoffwechsels (1905), p. 426. luCompt. rend. 136, 1097. Water of Respiration. 437 While with food experiment No. 70, the proportion of water eliminated from the lungs is somewhat less than during fasting, with experiments Nos. 72, 74, and 76, very little, if any, difference is noted. On the average, about 44 per cent of the water vaporized from the body of a resting man during fasting is from the lungs. Table 220. — Proportion of total water of respiration and perspiration eliminated from the lungs and skin, respectively, in metabolism experiments with and without food. Experi- ment number. 59. 68. 69. 70. 71. 72. 73. 74. 75. Subject and date. (a) Total venti- lation of lungs. A B. F. O Dec. Dec. Dec. A. L. L Apr. Apr. A. L. L Dec. Dec. Dec. Dec. L. L Dee. Dec. Dec. S. A. B. Jan. Jan. Jan. Jan. S. A. B. Jan. S. A. B. Jan. Jan. Jan Jan. Feb. S. A. B. Feb. Feb. Feb. S. A. B. Mar. Mar. Mar. Mar. Mar. Mar. Mar. 18-19, 1903. 19-20, 1903. 20-21, 1903. 27-28, 1904. 28-29, 1904. 16-17, 1904. 17-18, 1904. 18-19, 1904. 19-20, 1904. 20-21, 1904. 21-22, 1904. 22-23, 1904. 7- 8, 1905. 8- 9, 1905., 9-10, 1905., 10-11, 1905. Liters. 9,253 : 9,250 ' 9,498 9,408 9,446 8,588 9,493 9,106 8,838 9,149 9,867 10,773 8,660 8,146 7,908 7,244 11-12, 1905 ' 7,598 (6) Wa- ter in air in- spir ed. Oms. 74.30 70.30 70.76 79.03 78.87 66.13 83.73 74.85 66.11 70.45 1.11 93.08 66.68 51.08 45.31 38.18 41.64 (c) Water in air ex- pired. (d) Water elimi- nated from lungs (c-b). 28-29, 1905 29-30, 1905 30-3], 1905 31-Feb. 1, 1905 1- 2, 1905.... 2- 3, 1905. 3- 4, 1905. 4- 5, 1905. 4- 5, 1905. 5- 6, 1905. 6- 7, 1905 7- 8, 1905. 8- 9, 1905. 9-10, 1905. 10-11, 1905. 8,000 8,054 7,836 7,389 7,137 7,529 7,191 7,275 49, 43. 11, 36. 34. 38. 35. 36. 43. 38. 39. 7,844 7,854 7,875 7,637j35. 7,218j32. 6,85l|28. 6,856'29. 68 33 45 65 11 70 31 88 38 96 61 59 63 64 is Oms. 402.18 402.05 412.83 408.92 410.57 373.28 412.61 395.79 384.14 397.66 428.87 468.25 376.41 354.07 343.72 314.86 330.25 (e) Water elimi- nated from skin included determinations of the carbon dioxide. During experiments Nos. 36, 39, and 42, each of 24 hours' duration, the production was 711, 649, and 620 grams, respectively. The amounts eliminated for the 2 days of experiment No. 51 were 703 and 698 grams, respectively. Since the production of carbon dioxide and the metabolic activity are, in general, largely proportional to the body-weight, these older experiments may perhaps be better compared by noting the amounts of carbon dioxide per kilo- gram of body-weight and per square meter of body surface. The total carbon dioxide production for the older experiments is given in table 221, together with the amounts per kilo of body-weight and per square meter of body surface. 12TZeit. f. Biologie (1866), 2, p. 478. 128 Dissertation (Russian), 1893, St. Petersburg. 129 U. S. Dept. of Agr., Office of Expt. Sta. Bui. 136. 442 Influence of Inanition on Metabolism. Table 221. — Carbon dioxide production of fasting men, at rest. Investigators. Day of fast. Body weight. Carbon dioxide per 24 hours. Total. Per kilo of body- weight. Per square meter of body surface.1 Kilos. 71.9 70.0 70.0 Grams. 2 663 738 729 Gravis. 9.2 10.5 10.4 Grams. 311 353 349 Pettenkofer 183 5 1 362 181 1 7.1 9.0 16.1 8.1 178 2 183 r 361 ;» 180 '.» 153.3 152.5 305 . 8 152.9 7.7 10.8 18.5 9.2 209.8 220.0 429.8 214.9 9.7 12.8 175 170 2 7 22.5 11.3 345 173 9 o! 7.7 8.2 15.9 8.0 197.7 204.4 402.1 201.0 8.4 9.9 205.4 202.3 18.3 9.1 407.7 203.9 9.6 174.3 7.5 3.5 5.6 4.6 4.0 4.9 4.5 4.9 6.2 5.6 3.7 4.9 4.3 5.6 7.5 6.5 3.9 4.0 4.0 4.1 4.9 4.5 5.5 446 Influence of Inanition on Metabolism. The errors involved in the determination of carbon in the urine have been discussed previously, and even if a gross error of 5 per cent be assumed, the effect on the total carbon excretion is seen to be extremely small, and hence it is obvious that considerable errors in the determination of the carbon in the urine are practically without effect on the total carbon elimination in ordi- nary metabolism experiments. Carbon in the urine, feces, and respiratory products was determined in the 5-day fasting experiment with J. A. (9). There were found 8.0, 8.3, 9.9, 10.3, and 9.3 grams of carbon in the urine, while the corresponding amounts of carbon in the respiration were 188.5, 179.4, 172.2, 169.4, and 165.8 grams. Computed on the basis of the total carbon elimination, the percentages of carbon in the urine for the 5 days were 4.1, 4.5, 5.4, 5.7, and 5.3, respectively. The general conclusion can, therefore, be made that with fasting man at rest, the carbon of the urine is about 5 per cent of the total carbon excretion. OXYGEN CONSUMPTION. Since the classic experiments of Lavoisier showing the relation between oxygen consumption and vital processes, the direct measurement of oxygen in the respiratory gases has not been attempted, save in one or two cases. Although as Lusk 1S0 points out there is no record of the method employed by Lavoisier in obtaining his results, it is remarkable how closely his statements of the quantities of oxygen consumed by resting man agree with the measure- ments by modern methods. In the earlier experiments with the Pettenkofer respiration apparatus, in which the attempt was made to determine accurately the water vaporized, estimations of the amount of oxygen were attempted by computations based on the gains or losses of body weight, carbon dioxide excretion, and water-vapor elimination. When the errors involved in the determination of water-vapor are taken into consideration, it is seen that whatever approach to accuracy the oxygen estimations appear to have, it must have been the resultant of a num- ber of more or less compensating errors. An attempt was made in the more accurate experiments of Sadovyen (2), in which the amounts of carbon dioxide and water-vapor were determined for a great part of each day, to measure by the indirect method the amount of oxygen consumed. On the 2 days of the fasting experiment with man in which no water was consumed, the oxygen intake was estimated to be 1081 and 1010 grams, respectively. The oxygen consumption during the 4-day experiment, in which water but no food was taken, was estimated to be 746, 921, 698, and 943 grams for the different days. Similarly Likhachev m estimated indirectly the amount of oxygen consumed during a 1-day fast to be 596 grams, or 11.9 grams per kilo of body-weight, 50 Elements of the Science of Nutrition, Philadelphia (1906), p. 18. ulLoc. cit. Oxygen Consumption. 417 With the development of the type of respiratory apparatus brought to such signal success by Zuntz and associates, innumerable experiments have been made in which the respiratory quotient has been determined during brief fasts. It is true of the oxygen as of the carbon dioxide, that the determinations of the fasting values are of importance in the interpretation of the respiratory quotient in the large majority of experiments. This type of apparatus was used in determining the oxygen intake of the fasting subjects Cetti and Breithaupt (7). Although results were obtained for but a few minutes each day and during the respiration experiment the subject was lying on a sofa, the authors computed the total oxygen consumption for the 24 hours on the basis of these short experiments. The amounts of oxygen consumed for the 10 days of Cetti's fast were 567, 501, 482, 478, 470, 467, 502, 526, 489, and 449 grams, respectively. These values are obtained by multiplying the quantity of oxygen absorbed per minute during rest by the total number of minutes in the day and no allowance is made for the variations in muscular activity, so that the values can at best be only approximate. In the experiments on Breithaupt, the complications of a cold and other disturbances were so notice- able that the authors did not consider the computed results for the 24 hours as of general value. Hanriot & Kichet,132 using a different form of respiration apparatus, deter- mined the oxygen consumption of a fasting man after 46 hours of fasting. The amounts consumed per hour after the seventeenth, twenty-fourth, twenty- ninth, and forty-sixth hours of the fast were 17.04, 16.85, 16.05, and 16.90 liters, respectively. The subject of the experiment weighed 50 kilograms. These observations were likewise made only during short periods. The first direct determinations of oxygen consumed by man during 24 hours are those made with the Hoppe-Seyler 133 modification of the Kegnault and Eeiset apparatus, although none of the experiments reported as made in this apparatus were fasting experiments. Zuntz has recently devised an apparatus on the Kegnault and Keiset plan that can be used for small animals or infants.134 No experiments with infants have, however, as yet been reported. Direct determinations of the amount of oxygen consumed by man have been made in a number of experiments in this laboratory during the past 3 years. A brief preliminary report of one of these experiments 13° has been followed by a more detailed discussion of one day of an experiment with this 132Compt. rend, de l'Academie des Sciences (1888), 106, p. 496. 133Zeit. f. physiol. Chemie (1894), 19, p. 574. 1S4Archiv f. Physiol. (Physiol. Abth. d. Archiv f. Anat. u. Physiol.) (1905), Supplement, Band, p. 431. 135 A respiration calorimeter with appliances for the direct determination of oxy- gen, 24 pages. Wesleyan University, Middletown, Connecticut. Printed, not pub- lished, Aug., 1903, W. O. Atwater & F. G. Benedict. 448 Influence of Inanition on Metabolism. apparatus/86 and the results of a series of experiments covering 24 days have been recently published.137 So far as we are aware, the first determinations of the oxygen consumption of fasting man are reported herewith. The method has been considered in brief in the introductory section of this report and in detail elsewhere.188 The accuracy of these measurements has also received special treatment. In brief, it may be said that for experiments of 24 hours or longer the results are extremely satisfactory. For 2-hour periods the oxygen determinations are satisfactory only when there is like muscular activity at the beginning and end of the period, since constancy in the average temperature of the air residual in the chamber is of the utmost importance in securing accurate oxygen measurements.139 The detailed data regarding oxygen during the 2-hour periods of the fasting experiments are given in the statistical data in the preceding section. The total amounts of oxygen consumed during each 24 hours are shown in the following table : T able 224. — Oxygen consumed in metabolism experiments without food. Exper- iment num- ber. Subject and duration of experiment. First day. Second day. Third day. Fourth day. Fifth day. Sixth 'Seventh day. day. 59... B.F.D., Dec. 18 to 20, 1903 Oms. 629.4 Oms. 629.2 Oms. 646.1 Oms. Oms. Gms. Oms. 68... A.L.L., Apr. 27 to 28, 1904 640.0 642.6 .... .... .... . . 69... 71... A.L.L., Dec. 16 to 19, 1904 589.1 645.8 554.1 619.4 538.0 601.2 492.7 .... .... 73... S.A.B., Jan. 28 to Feb. 1, 1905. 544.2 547.9 533 . 0 502.7 485.5 .... . . 75... S.A.B., Mar. 4 to 10, 1905 533.6 534.3 535.7 519.5 491.0 466.1 466 4 77... S.A.B., Apr. 8 to 11, 1905 556.0 571.6 530.7 531.6 .... . . 79... H.E.S., Oct. 13 to 14, 1905 575.9 605.3 .... .... .... . . 80... C.R.T., Oct. 27 to 28, 1905 576 . 2 628.6 .... .... . . 81... A.H.M., Nov. 21 to 22, 1905 516.8 527.1 . . .... .... 82... H.C.K., Nov. 24 to 25, 1905 663.3 733.8 . . .... . . . . . . 83... H.R.D., Dec. 5 to 6, 1905 585.2 554.4 .... .... 85... N.M.P., Dec. 9 to 10, 1905 627.5 675.6 . . .... .... . . 89... D.W., Jan. 10 to 11, 1906 645.4 681.3 .... 609.4 567.2 529.6 488.3 466.1 466 4 Table 224 shows that the total oxygen consumption during fasting may vary widely with different individuals. Thus, on the second day of experiment No. 82, 734 grams were absorbed, while on the sixth and seventh days of experi- ment No. 75, only 466 grams were consumed. The average for the first 2 days of fast is nearly the same, i. e., 591 and 609 grams. The amounts are 184 Carnegie Institution of Washington Publication No. 42, p. 178. 1,7 U. S. Dept. Agr., Office of Expt. Sta. Bui. 175 (1907). ^Carnegie Institution of Washington, Publication No. 42; U. S. Dept. Agr., Office of Expt. Sta. Bui. 175. 130 For discussion of the errors affecting the oxygen determination see Office of Expt. Sta. Bui. 175, pp. 24-32. Eespiratory Quotient. 449 larger on the second day of fasting than on the first day with but three excep- tions, the increase being, in one instance, as much as 70 grams, and in another about 61 grams. In practically all cases the oxygen intake diminishes after the second day, reaching a constant, 466 grams, on the sixth and seventh days. Considering experiments with the same individual, S. A. B., it is to be noted that even on the first day of fast, there are noticeable differences in the amounts consumed, which range from 534 grams in experiment No. 75, to 589 grams in experiment No. 71. The excessive amount in the latter case may, however, be explained, in part at least, by the fact that a 10-minute period of hard muscular exercise on the bicycle ergometer was taken on this day. A similar variation appears, however, between the amounts for the first day of experiments Nos. 68 and 69, made with the subject A. L. L. Obviously, as was the case with the carbon dioxide elimination, differences in the bodily activity and body-weight of the subjects may explain in large part the variations in the amounts of oxygen consumed during the experiments, and indeed during different experiments with the same subject. The rela- tions between the oxygen consumption, carbon dioxide excretion, heat pro- duction, and muscular activity are discussed in detail in a subsequent section. The discussion of the relations between the body-weight and the oxygen intake are also considered elsewhere. RESPIRATORY QUOTIENT. The ratio between the volumes of carbon dioxide excreted and oxygen con- sumed is of great value in indicating the nature of the material undergoing katabolism in the body. The combustion of the carbohydrate is accompanied by an absorption of oxygen which in the course of the oxidation results in the production of an equal volume of carbon dioxide. On the other hand, when fat is oxidized, the oxygen absorbed is used to oxidize not only the carbon of the fat, but also the organic hydrogen. While the combustion of the carbon results in the formation of an aeriform combustion product, carbon dioxide, oxidation of the hydrogen results in the formation of water-vapor which is not measured in the ordinary methods of studying the respiratory gases. Consequently, the volume of carbon dioxide excreted is less than that of the oxygen consumed. The ratio between the volumes of carbon dioxide and the oxygen is called the respiratory quotient. It is commonly expressed by the CO fraction ^-? . In the case of the carbohydrate, this respiratory quotient is equal to 1 since equal volumes of oxygen and carbon dioxide are involved in the respiratory exchange. The combustion of fats, on the other hand, results in a respiratory quotient which is always less than 1, and for the ordinary animal fats, it has been computed to be about 0.711.140 140 For a more detailed discussion of the theoretical respiratory quotients to be derived from the combustion of different kinds of fats and carbohydrates, see U. S. Dept. Agr., Office of Expt. Sta. Bui. 136. 29 450 Influence of Inanition on Metabolism. The computation of the theoretical respiratory quotient from the combustion of carbohydrate or fat is relatively simple, but the nature of the cleavage of the protein molecule and the excretion of partially oxidized material in the urine complicates the computation of the quotient resulting from the oxidation of protein. The value commonly accepted for protein is 0.809. For further discussion of the theoretical factors involved in the computation of the theoretical respiratory quotients resulting from the combustion of protein see Magnus-Levy.141 While the theoretical consideration of the respiratory quotients to be derived from the combustion of the various nutrients in the body is capable of mathe- matical expression, considerable difficulty is experienced in interpreting the respiratory quotients actually determined on man. For, instead of the com- bustion of any given one of these ingredients of the body, protein, fat, or carbohydrate, there is in practically all cases a simultaneous combustion of the three. It has commonly been believed that in fasting experiments of short duration, the combustion is chiefly that of fat with a small amount of protein, and it has been maintained that the combustion of protein is relatively con- stant from hour to hour. Hence, in the studies made by Zuntz, Loewy, Magnus-Levy, and their associates, the quotient obtained during fasting has been taken as a measure of the respiratory quotient derived from the com- bustion of fat with a constant amount of protein. The various factors influencing the quotients, such as the ingestion of food, muscular exercise, etc., have been studied in the majority of the experiments made by these investi- gators, and the results obtained are, for the specific purpose for which they were designed, all that could be desired. On the other hand, when the respiratory quotient is determined not only for the total 24 hours, but also for 2-hour periods, there may be, during the periods, a lack of uniformity in the amount and rapidity of the disintegration, not only of protein but likewise of carbohydrate, and, therefore, it is readily seen that the interpretation of quotients for 2-hour periods, even when cor- rectly determined, is a matter of considerable complexity. Indeed, the use commonly made of the respiratory quotient is of but little value in complete metabolism experiments, and recourse must be had to the more complete apportionment of the oxidation among the three principal ingredients of the body — protein, fat, and carbohydrate — according to the method elaborated and discussed in connection with the statistical data of experiment No. 59 (p. 36). The respiratory quotients for each day, as determined in these experiments, are of decided general interest judged from the standpoint of the observations of other investigators, and accordingly they are here presented in table 225. 141 Physiologie des Stoffwechsels (1905), p. 217; also Zeit. f. klin. med. (1906), 60, Heft. 3, p. 1. Eespieatory Quotient. 451 Table 225. — Respiratory quotients for 2Jf hours in metabolism experiments without food. Exper- 1 iment num- ber. Subject and date. First day. Second Third Fourth Fifth day. day. day. day. Sixth day. Seventh day. 59... B.F.D., Dec. 18 to 20, 1903 0.78 0.76 0.73 1 68... A.L.L., Apr. 27 to 28, 1904 .79 .77 . , . .... .... .... • * . . 69... A.L.L., Dec. 16 to 19, 1904 .79 .75 .75 0.74 * . * • .... .... 71... S.A.B., Jan. 7 to 10, 1905 .. . .83 .75 .75 .75 . • • . .... 73. .. 8.A.B., Jan. 28 to Feb. 1, 1905. .81 .74 .74 .75 0.72 .... . . . 75... S.A.B., Mar. 4 to 10, 1905 .78 .75 .74 J .75 .74 0.75 0.74 77. .. iS.A.B., Apr. 8 to 11, 1905 .78 .73 .76 j .75 .... .... 79... H.E.S., Oct. 13 to 14, 1905 .80 .76 . . .... • . . • 80... C.R.Y., Oct. 27 to 28, 1905. . . .79 .74 . . i • . . .... .... 81... A.H.M., Nov. 21 to 22, 1905 .75 .72 .... 82... H.C.K., Nov. 24 to 25, 1905 .81 .76 . , * . • ■ .... 83... .75 .76 , . .... . . . • 85... .81 .77 . . .... . . * . 89... D.W., Jan. 10 to 11, 190(5 . . .81 .75 .... .... 0.79 0 . 75 0.7- 1 0.74 0.73 0.75 0.74 The respiratory quotients for the 2-hour periods are given with the detailed statistics of the experiments. Unfortunately, but little reliance can be placed on these determinations for short periods save in those instances when the muscular activity of the subject was the same at the beginning and end of the experimental period. As has been pointed out previously, the exact determi- nation of the average temperature of the large volume of air residual inside the chamber (4900 liters) is of the utmost importance to the accurate measure- ment of the oxygen consumption. Under like conditions of muscular activity at the beginning and end of each period, the average temperatures may readily be secured. Experience has shown that even minor differences between the muscular activity at the beginning and at the end of a period result in abnor- mal temperature observations for the residual air and consequently erroneous oxygen determinations and respiratory quotients. On the contrary, since the experimental day ends at 7 a. in. and the subject is quietly resting in bed, the respiratory quotients for 24 hours we believe to be as accurate as can be determined with the type of apparatus used in these experiments. The marked acidosis accompanying certain fasting experiments and which we have reason to believe was not absent in some of those here reported, would have an effect upon the respiratory quotient. The conversion of partially oxidized fat into /? oxybutyric acid, for example, would mean a taking up of oxygen unaccompanied by a corresponding liberation of carbon dioxide, and hence the oxygen consumption would be too large and the respiratory quotient too small. The exact effect of this acidosis on the quotient has not been com- puted since the respiratory quotients are not used in this discussion in their ordinary sense. The influence of such an absorption of oxygen unaccompanied 452 Influence of Inanition on Metabolism. by a carbon dioxide liberation on the computation of the protein, fat, and carbohydrate by means of the method of simultaneous equations might be very complex. One serious objection to this method of computation is that the possi- bilities of intermediary metabolism are not considered in any way, since only the end products of metabolism are used for obtaining the data. It is believed, however, that acidosis did not proceed to any such degree in these experiments as to depreciate materially the accuracy of the computations of the protein, fat, and glycogen katabolized. BODY MATERIALS KATABOLIZED. The measurement of the end products of katabolism appearing in the respira- tory gases and the urine furnishes the necessary data for determining the nature of the total katabolism. Aside from the interpretations of the nitrogen output and more specifically the recent emphasis laid upon the interpretation of the partition of the nitrogen in the urine, deductions from the respiratory quotient have been of greater value in indicating the nature of the total katabolism than have those from any other measured factor. But while the respiratory quotient is an admirable index of katabolism in short experiments in which the body is at absolute rest, provided that the body material has not previously been heavily drawn upon as a result of a protracted period of fast- ing, knowledge of the katabolic processes during inanition has been so defi- cient that the usual method of employing the respiratory exchange as an index of katabolism during a prolonged fast is hardly justified. In nearly all the studies of the respiratory exchange that have thus far been made, the so-called " michternwert " is of fundamental importance. It is necessary, however, in considering experiments of the nature of these here reported, to bear in mind that, even after 12 hours' fast, there may still be a considerable absorption from the intestine of food or at least of partially digested material. So long as this absorption is uniform, it introduces no material error into the determina- tion of the fasting value, since its effect is measured along with the effect of the continued protein katabolism. The ingestion of even small amounts of food produces immediately very considerable changes in the respiratory ex- change, and since in all experiments of short duration, the differential method is emplo}red, it is important in studying factors influencing metabolism such as the ingestion of food and muscular work, that the respiratory exchange be measured prior to the ingestion of food or the beginning of muscular exercise. Hence, the value of the respiratory exchange, when the body is at absolute rest after a period of 12 hours without food, is usually taken as the basis for comparison. The differences, then, between the respiratory exchange during rest and that of the changed condition of the experiment indicate the effect of the change. The respiratory quotients obtained in the fasting experiments here reported, on the other hand, represent the resultant value of all the oxy- gen absorption and the total carbon dioxide production of each 24 hours. Katabolism of Protein. 453 As commonly used, the respiratory quotient excludes completely any con- sideration of the total nitrogen excretion, on the assumption that for the short period of the respiration experiment the katabolism of protein is constant. In the experiments made in this laboratory the protein broken down is as accurately and regularly measured as is the carbon dioxide production, and hence the total carbon dioxide excretion can be resolved into certain portions representing the katabolism of the various ingredients of the body. The determination in metabolism experiments of the complete carbon dioxide and nitrogenous output alone has served heretofore as the basis for the compu- tation of the proportions of protein and fat katabolized. In this computation it has been necessary to assume that the store of glycogen in the body remained constant during the experiment. Such an assumption may not be grossly erro- neous during food experiments in which a maintenance ration is employed, though for periods of inanition where there may be very considerable drafts upon the store of glycogen, this method of computation, admittedly but approx- imate for experiments with food, may be entirely inadequate. It was early recognized in this laboratory that a fundamental study of metabolism during inanition is necessary for a proper understanding of many physiological processes. But until the means were at hand for determining directly the oxygen consumption, it was apparent that such studies were of but little value. With the added data regarding the total amount of oxygen absorbed per day, a much more accurate apportionment of the total katabolism among protein, fat, and carbohydrate is possible. The apportionment is based upon the chemical determinations showing the losses of material to the body in terms of chemical elements, and for this reason especial care has been taken in making the elementary analyses. The losses to the body of nitrogen, carbon, hydrogen of organic matter, oxygen, water, and ash have all been determined as accurately as present methods of analysis will permit. The method of obtaining the gains and losses of chemical elements has been discussed in detail in connection with experiment No. 59 (see p. 36). From these gains or losses of elements it is possible by means of the method of simultaneous equations to compute the proportions of protein, fat, and carbohydrate katabolized. The detailed treatment of this subject is likewise given in connection with the discussion of the same experiment. It is only necessary here, therefore, to compare the total amounts of protein, fat, and carbohydrate katabolized on different days of the different fasting experiments. KATABOLISM OF PROTEIN. Since the protein katabolized is computed from the nitrogen excretion by multiplying the amount of nitrogen by the factor 6.0, the discussion which has been accorded the total excretion of nitrogen during fasting applies in general in this connection. For while, as is well known, the nitrogen excreted 454 Influence of Inanition on Metabolism. in the urine may have originated either in katabolized protein, purin bodies, creatine and creatinine, or preformed urea and other crystallized end products of katabolism, the only index of the total protein katabolism commonly used is the total excretion of nitrogen. Contrary to the natural supposition, the problem is most complicated in fasting experiments ; for although under ordi- nary conditions with food the excretion of phosphorus and sulphur may be significant of the breaking down of protein, during inanition the excretions of nitrogen, sulphur, and phosphorus indicate profound disturbances in the katabolism. A number of writers have attempted to compute the protein katabolized from the sulphur elimination, by assuming that the sulphur content of protein is relatively constant. The ratio of nitrogen to sulphur has been used in this connection. But the wide variations in the amounts of sulphur present in the various proteins existing in the body make it difficult to utilize with any degree of satisfaction the data obtained from the sulphur and nitrogen determinations and especially the ratio between them. Sherman142 has computed the ratio of nitrogen to sulphur in a large number of vegetable and animal proteins. With the proteins of the body, the ratios range from 44.6 : 1 in oxy-hemoglobin to 5:1 in tendon mucin and osseo-mucoid ; in myosin, serum globulin, and fibrinogen, the ratios are more constant, namely, 13.1 : 1, 14.3 : 1, and 13.3 : 1, respectively. It is commonly assumed that body protein contains from 16 to 17 per cent of nitrogen, and since this percentage is not markedly different in the different animal proteins, the assumption is reasonably well grounded; but as has been shown above, the ratios of nitrogen to sulphur vary so widely as to practically preclude any scientific deductions from them regarding the nature and amount of the protein katabolized. The probability that there are at least two kinds of protein katabolism occurring in the body, namely, exogenous and endo- genous, justifies the belief that there is a noticeable difference in the sulphur content of the two kinds of protein katabolized. While the nitrogen-sulphur ratio in fibrinogen, serum globulin, and myosin is fairly constant and this could properly be used if these were the only proteins katabolized, an examina- tion of the ratios found in the experiments here reported (see table 209) shows that on only one day was the nitrogen-sulphur ratio as low as 14.12, namely, the fifth day of experiment No. 73. The ratios on all the other days were considerably higher, averaging not far from 16.8. This fact clearly indicates that material amounts of some form of protein with a lower sulphur content than that of myosin, serum globulin, or fibrinogen, must have been katabolized. Obviously, this ratio would be very much increased by an excretion of the sulphur-free extractives. The output of total creatinine, from the results obtained in these experiments (see table 203, p. 388) remains singularly con- U. S. Dept. Agr., Office of Expt. Sta. Bui. No. 121, p. 10. Katabolism of Protein. 455 stant throughout the fast, and according to the results of Brugsch (12), the purin bodies remain constant, at least on the last 8 days of a 30-day fast. This would imply that there was no excess in the excretion of extractive nitrogen. The sulphur determinations are of value in suggesting the nature rather than the total amount of the protein katabolized. But little satisfaction attends the attempt to designate the kinds and amount of protein katabolized from the phosphorus output, for as has been shown in discussing the phosphorus excretion, the ratio of nitrogen to phosphorus is very much smaller than that occurring in the ordinary nucleo-proteins, which signi- fies that in all probability phosphatic material of the bones has been drawn upon and excreted. An attempt has been made to measure protein katabolism by means of the chlorine excretion, assuming that the chlorine content of muscle is constant. From the discussion of the chlorine output during fasting, it is apparent that here again the results of the experiments are such as to preclude an accurate comparison between the elimination of chlorine and the amount of protein katabolized. Becent advances in the study of the significance of the partition of nitrogen would imply that a knowledge of the purin bodies, amino-acids, ammonia, and uric acid of the urine would furnish a much more accurate basis for the computation of the total protein katabolism than would perhaps the total nitrogen excretion. In the absence of more definite information regard- ing the nature of the nitrogenous material excreted during fasting, the only alternative is to follow the usual custom and to assume that the total nitrogen output indicates the total protein katabolism. In table 226 the amounts of protein katabolized per day, computed by multiplying the total nitrogen excre- tion by 6.0 are recorded. There are considerable differences in the body- weights of the subjects of these experiments, and consequently the amounts per kilo of body-weight are also recorded for purposes of comparison. Since there is a mass of evidence to show that muscular activity, at least when not excessive, does not influence the nitrogen output, the uniformity or lack of uniformity, between the total amounts of protein katabolized by different subjects and on different days by the same subject can be better compared than the katabolism of either carbohydrate or fat. An examination of the figures in the table shows that during inanition, the protein katabolized on the first day of fasting varies from 35.0 grams to 79.5 grams. The average for the first day for the 14 experiments is 60.2 grams. Aside from the extremely low amount on the first day of experiment No. 71, the lowest result for the first day is 46.7 grams. On the basis of per kilo of body-weight, the fluctuations for the first day of the fast range from 0.61 to 1.44 grams, averaging 0.94 gram. The variations from the average protein katabolism per kilogram of body-weight indicate that there is no approach to uniformity in the results obtained on the first day without food. 456 Influence of Inanition on Metabolism. The unusually low nitrogen output on the first day of experiment No. 71 has been the subject of much study. Unfortunately, since this was the first day of experimenting with this subject, no data are at hand regarding his protein katabolism prior to the fast nor, indeed, have we that accurate record of food consumed and nitrogen excreted during the several weeks before this fast that is available for the interpretation of the protein disintegration in one of the subsequent fasts. Table 226. — Protein katabolized in metabolism experiments without food. Experi- ment num- ber. Subject and duration of experiment. First day. Second day. Third day. Fourth day. r-5 o H Cm 5.3> £ © •§ ' 13 -a n S ©.Q Pw "3 o 5 s« U © O f Oh "3 o H S tm TO -^ U © m> Ol — >> Ph "5 o 5 ^ C3-.H t< © ,2 ' Ph 59 68 .. 69.... 71 73 75 77.... 79 80.... 81 82 83 85 89 B. F. D., Dec. 18 to 20, 1903 A. L. L., Apr. 27 to 28, 1904 A. L. L., Dec. 16 to 19, 1904 . . . S. A. B., Jan. 7 to 10, 1905 S. A. B., Jan. 28 to Feb. 1, 1905. S. A. B., Mar. 4 to 10, 1905 S. A. B., Apr. 8 to 11, 1905 H. E. S., Oct. 13 to 14, 1905 C. R. T., Oct. 27 to 28, 1905 H. R. D., Dec. 5 to 6, 1905 D. W., Jan. 10 to 11, 1906 Oms. 71.0 73.6 60.5 35.0 61.7 73.4 52.9 48.7 46.7 79.5 68.2 59.9 Gnis. 1.06 1.02 .82 .61 1.05 1.23 .86 .86 .68 .89 .79 1.44 1.02 .77 Oms. 84.7 78.2 85.6 66.2 71.8 74.7 64.7 86.1 59.7 78.3 86.2 81.2 68.1 86.8 Oms. 1.28 1.11 1.18 1.17 1.25 1.26 1.07 1.55 .90 1.30 1.23 1.49 1.04 1.13 Oms. 88.9 90.2 78.6 69.2 78.1 65.9 Oms. 1.36 1.25 1.39 1.22 1.34 1.12 (7 ms. 77.8 64.4 62.3 69.8 68.7 0)ns. 1.09 1.16 1.11 1.21 1.19 60.2 .94 76.6 1.21 78.5 1.28 68.6 1.15 Experi- ment num- ber. Subject and duration of experiment. Fifth day. Sixth day. Seventhday. o H If JH © bD£ OT n2 >> *8 ©.o Ph -M o 2 *» Sh © h ° Ph "3 O <> XV £.o Ph 73.... 75 S A. B Jan. 28 to Feb 1 1905 Oms. 59 9 Oms. 1.08 1.15 Oms. 64.4 Gms. 1.14 Gms. 60.8 Gms. 1.08 S. A. B., Mar. 4 to 10, 1905. . 65.2 62.6 1.11 64.4 1.14 60.8 1.08 The amounts of protein katabolized in the different experiments are much more nearly uniform on the second day of fasting, the minimum being 59.7 grams and the maximum 86.8 grams. The average for the 14 experiments is Katabolism of Protein. 457 76.6 grams, a material increase over the average for the first day, thus sub- stantiating the contention made by Prausnitz "* that the protein katabolism on the first day of fasting is regulated in large measure by the amount of glycogen in the body. Furthermore, reference to the quantities of glycogen katabolized (see table 228) on the second days of these experiments shows that the amounts of katabolized glycogen in all but one instance were greatest on the first day. Computed on the basis of per kilogram of body-weight, the protein kata- bolism on the second day of fasting is reasonably constant, experiments Nos. 79 and 80 being the marked exceptions. Omitting these experiments, the widest variation from the average (1.21 grams) is 0.28 gram. The uniformity of the protein katabolism is still further supported by the results on the third day of fasting, the maximum amount being 90.2 grams, the minimum 65.9 grams, and the average 78.5 grams, somewhat more than the average for the second day. The average per kilogram of body-weight is 1.28 grams, and the fluctuations from this average in the different experiments are very small. Similarly, constancy in the results from day to day is observed on the fourth day of fasting, the amounts on the basis of per kilogram of body-weight being practically uniform after the third day. In comparing the results of protein katabolism as indicated in this table, it is important to bear in mind that the factor for the computation of protein here employed is 6.0, while the factors 6.3 and 6.25 are commonly used to express the ratio between nitrogen and protein. Obviously, the use of either of these latter figures, would indicate a somewhat greater protein katabolism than is recorded in the table. The results show that on the first day of fasting the amounts are extremely irregular and that the fasting protein katabolism can not be said to be estab- lished before the second day. The bearing of the protein katabolism on the protein requirement, the influence of the previous store of glycogen,144 fat and protein, and the relation of the protein katabolism to the total active mass of protoplasmic tissue are problems all closely related, but discussion of them in this place would be beyond the confines of this report. With the publication of experiments made subsequent to these here reported, it is hoped that these problems may be more fully dealt with. Certain of the data accumulated in this series of experiments, while admit- tedly too meager to serve as the basis for the enunciation of a theory of the effect of inanition upon protein katabolism, are sufficiently suggestive to warrant more complete examination. 143Zeit. f. Biologie (1892), 29, p. 151. 144 The interesting discussion of the relation between the store of glycogen in the body and the proteid katabolism by Landergren (Skan. Archiv f. Physiol. (1903), 14, p. 169), is supplemented by the observations on the actual glycogen katabolism observed in these experiments. 4.58 [nfluenoe of Inanition ox Metabolism. In tlio discussion of the creatinine output (see p. 389) the interpretation of the results obtained in these experiments viewed in the light of the Folin theory of protein metabolism was pointed out. The attempts to harmonize the results of the fasting experiments with this theory showed that, while the body during fasting may protect reserve tissue protein to a greater extent than during ordinary conditions of nutrition, there is also the possibility that fasting may result in a greater encroachment upon stored organized body material. The fact that the total creatinine output remained constant even during a 7-day fast was taken as an indication that no greater drafts were made upon organ- ized protein than during complete nutrition. The presence of preformed creatine in the urine of fasting man was con- sidered the result of a decrease in the power of the body to dehydrate the creatine resulting from protein katabolism. In support of this view is the remarkable fact that the total creatinine elimination, i. e., preformed creatinine plus preformed creatine (expressed in terms of creatinine), remains constant throughout the fast. Without losing sight of the possibility of this expla- nation, it is of interest to consider another possible correlation of these data. The exact nature of the endogenous protein katabolism is not known. That it is essentially different from the exogenous is by no means inconceivable, and if the assumption is made that tissue protoplasm is broken down, the marked decrease in the total protein katabolism during prolonged fasting would imply that the katabolism of tissue protein would likewise decrease. The total creatinine output considered in the light of the foregoing discussion then would indicate that the tissue katabolism was supplemented by some other source of creatinine. As a matter of fact the preformed creatinine elimination decreases as the fast progresses and the preformed creatine increases. The presence of con- siderable amounts of creatine in flesh would suggest that in the katabolism of body flesh during inanition this creatine was liberated and that it was excreted by the body as such. This involves no assumption regarding the source of the preformed creatinine excreted during inanition. This latter may be derived from the katabolism of protein or it may be taken from the preformed creatine in the muscle, dehydrated to creatinine and so excreted, but, unfortunately, no evidence is at hand to show clearly this phase of intermediary metabolism. The suggestion above, therefore, assumes that the so-called endogenous protein katabolism during inanition decreases as the fast progresses, using the preformed creatinine excretion as the measure of endogenous katabolism. The excretion of preformed creatine indicates the katabolism of flesh. According to the results of Van Hoogenhuyze and Verploegh (11), each kilogram of flesh contains not far from 4.2 grams of creatine.14*3 In the later '"aGrindley: Jour, of Biol. Chem. (1907), 2, p. 4, has recently found 4.1 grams of creatine in one kilogram of lean beef. Katabolism of Protein. 459 days of fasting in the Middletown experiments the total amount of protein katabolized was not far from 60 grams, corresponding to about 300 grams of flesh. Assuming the whole protein katabolism to have resulted from katabol- ized flesh about 1.2 grams of creatine would have been set free. The amounts actually found are much less than this theoretical quantity. The absence of uniformity in the nitrogen-sulphur ratio points strongly to a protein katabolism other than that of muscle. As the fast progresses this ratio gradually approaches that of muscle protein. It is suggested, then, that the true measure of flesh or muscle katabolized may in fasting man be the amount of preformed creatine excreted in the urine. According to this hypothesis the total protein katabolism on the first few days of fasting involves little muscle protein, but as the fast progresses the muscle protein becomes dis- integrated and releases creatine which is excreted unchanged. The muscle katabolism does not comprise the total protein katabolism, for the creatine excretion is practically constant on the fifth, sixth, and seventh days. The nitrogen-sulphur ratio likewise becomes constant during these periods, and the total amount of creatine excreted would correspond to that in but 125 grams of katabolized flesh. There must be a continuous protein katabolism other than that of flesh, which has reached a minimum on the fifth day. This suggestion of the use of preformed creatine eliminated in the urine as an index of flesh katabolized has at least the value of indicating many possible research problems. Our experimental data are far too limited to bring the discussion out of the field of speculation. Folin 145 has shown that, contrary to the prevailing opinion, creatine ingested, especially after a period with food in which the protein content of the body has been somewhat lowered, i. e., after a low protein diet, is not excreted unchanged, nor indeed is it excreted in the form of creatinine. There is, moreover, no corresponding increase in the total nitrogen elimination, and hence it would appear that under these conditions the creatine is retained by the body. On the other hand, his experiments show that when the body is surcharged with protein, as after a high protein diet, the creatine ingested is in large part excreted as such. These results are difficult to harmonize with those obtained in the fasting experiments here reported unless some such assumption as is suggested above is made. During a low protein diet the creatine of the muscles is apparently drawn upon and whether it is excreted as creatine or as creatinine we as yet do not know. Under such conditions, then, the ingestion of creatine supplies the drafts upon the body creatine and there is no loss through the urine. It would be interesting to see how soon the store of creatine in the body would be replenished and the excess excreted in the urine with the subject remaining on a low protein diet. 145 Festschrift fur Olof Hammarsten (1906). 4G0 Influence of Inanition on Metabolism. The excretion of neutral sulphur parallels in a marked manner that of pre- formed creatinine. While Folin is inclined, according to recent statements,140 to regard the excretion of neutral sulphur in a somewhat different light than when he enunciated his theory of protein metabolism, it is significant that, in fasting experiments, the excretion of preformed creatinine and neutral sulphur should decrease as the fast progresses at practically the same rate. But one explanation for this phenomenon appears at present. Possibly dur- ing the earlier stages of fasting, the fluid proteins are first drawn upon. This represents a condition of katabolism not unlike that during the ordinary digestion of food. It may be measured by the total creatinine and the neutral sulphur output. The body becomes depleted of its fluid protein as the fast continues, and then the muscle is disintegrated and the creatine in the flesh katabolized is set free. Under ordinary conditions the body may cleave the protein molecule to creatinine during the process of katabolism, but be unable to dehydrate preformed creatine to creatinine. On this assumption it would appear that the preformed creatine existing in the muscles does not undergo any katabolism, that it is a relatively constant quantity and only when actual muscle substance is drawn upon, as during inanition, is it excreted as such. The remarkable constancy of the total creatinine excretion as the fast pro- gresses does not justify final conclusions regarding the excretion of creatine in flesh katabolized during the later clays of fasting. It would seem more than a coincidence that the amount of creatinine resulting from endogenous protein katabolism plus the amount of creatine in flesh katabolized should remain constant during the whole of the 7-day fast. Eecently Lichtenfelt,147 observing the influence of inanition upon the com- position of fish muscle, found that the muscle became richer in water when due allowance was made for the loss of fat and protein. This observation, likewise, is of interest in connection with the older assumption that when flesh was katabolized the water in it was liberated and excreted. According to Lichtenfelt the water is not necessarily eliminated, but the muscle has a higher water content. While Lichtenfelt's results indicate that there may be a loss of protein from muscle, Abderhalden, Bergell & Dorpinghaus 148 found on investigating the proteids of the blood and the body, by the esterfication method, that there was no difference to be observed when comparing the determinations on animals in health and animals after inanition, thus showing that the residual protein had not been altered as a result of inanition. While undoubtedly a portion of the katabolized protein is that of muscle, a significant fraction is in all probability derived from some of the larger organs. 14a Extract from a personal letter cited by Shaffer, Amer. Journ. of Physiol. (1906), 16, p. 274. u7Archiv f. d. ges. Physiol. (1904), 103, p. 402. lttZeit. f. physiol. Chem. (1904), 41, p. 153. Katabolism or Fat. 461 It appears clearly established that there is a marked diminution in the size of the liver149 and pancreas as well as other glandular organs, and in all proba- bility these organs are drawn uporj to a considerable extent to furnish the protein for katabolism. Since they rapidly regain their original size on the ingestion of food, it is probable that the fluid of the cell rather than the organized cell proteid is the portion drawn upon. KATABOLISM OF FAT. It has been customary in all experiments in which the katabolism of fasting man has been measured, to assume that the total fasting metabolism was sus- tained by the katabolism of protein and fat. The amounts of these compounds katabolized by a man at rest were computed from analyses of the respiratory products and the urinary nitrogen. The total protein katabolism was com- puted from the nitrogen excreted in the urine and the amount of carbon in the protein was calculated from the weight of protein katabolized. The carbon from protein was deducted from that of the carbon dioxide and the carbon in the urine, and the remainder was considered to have resulted from the kata- bolism of fat. This procedure has been common in the calculation of all meta- bolism experiments up to the publication of the direct determination of oxygen in some of the Middletown experiments.150 With the direct determination of oxygen, the data are available for computing by means of simultaneous equa- tions (see p. 38) the quantities of protein, fat, and carbohydrate katabolized. The amounts of fat katabolized per day and per kilogram of body-weight in fasting experiments have been computed and are recorded in table 227. In all cases there was a material draft upon body fat which was never less than 106.6 grams per day. On one day 203.6 grams were katabolized. The amounts for the first day of fasting ranged from 106.6 to 156.2 grams, and averaged 135.1 grams. The katabolism per kilogram of body-weight averaged 2.10 grams. In all but one instance, more fat was katabolized on the second day of the fast, the average amount for the 14 experiments being 165.9 grams or 2.61 grams per kilogram of body-weight. The amounts for the third day were by no means as regular as might be expected, the quantities ranging from 183.4 grams in experiment No. 59 to 137.7 grams in experiment No. 77. On the basis of per kilogram of body-weight, the range was from 2.80 to 2.25 grams. The average fat katabolism of the 6 experiments was 155.2 grams, somewhat less than the average for the second day of all the experiments. The average amount on the fourth and fifth days was about 147 grams, and on the sixth and seventh days about 15 grams less per day. The largest averages, per kilogram of body-weight, are found on the second and fifth days, while on the first and sixth days, the smallest amounts appear. The minimum katabolism per kilo 148 Possibly due to depletion of the store of glycogen. 150 U. S. Dept. Agr., Office of Expt. Sta. Bui. 175 (1907). 462 Influence of Inanition on Metabolism. of body-weight was observed on the first day of the fasr. A proper interpre- tation of these fluctuations in the katabolism of fat can not be made without a comparison of the amounts of glycogen which were simultaneously katabolized. The large amount of fat drawn upon by the body to support its vital func- tions is of special interest in connection with the question of acidosis during fasts. This observation is wholly in accord with the modern conceptions regarding the formation of organic acids from fat. Table 227. — Fat katabolized in metabolism experiments without food. Experi- ment num- ber. Subject and duration of experiment. First day. o 5 *» U CS Ph Second day o § be Ph Third day. o I to CO -.-t Mrs 4).Q Ph Fourth day o § to TO -h t. 4) 5 ' b O Ph 59.. 68.. 69.. 71.. 73.. 75.. 77.. 79.. 80.. 81.. 82.. 83.. 85.. 89.. B. F. A. L. A. L. S. A. S. A. S. A. S. A. H. E. C. R. A. H. H. C. H. R. N. M. D. W. D., Dec. L., Apr. L., Dec. B., Jan. B., Jan. B., Mar. B., Apr. S., Oct. T., Oct. M., Nov. K., Nov. D., Dec. P., Dec. , Jan. 18 to 27 to 16 to 7 to 28 to 4 to 8 to 13 to 27 to 21 to 24 to 5 to 9 to 10 to 20, 1903. 28, 1904. 19, 1904. 10, 1905. Feb. 1,1905 10, 1905.. 11, 1905 . 14, 1905.. 28, 1905. . 22, 1905 . 25, 1905.. 6, 1905.. 10, 1905 . 11, 1906.. Gms. 150.7 145.1 134.9 116.5 106.6 126.4 135.0 132.6 141.6 146.9 140.1 156.2 127.4 131.8 Gins. 2.25 2.01 1.84 2.02 1.82 2.12 2.20 2.34 2.07 2.39 1.97 2.82 1.91 1.68 Gms. 156.6 160.6 174.3 152.3 151.7 147 5 171.9 158.2 190.1 161.2 203.6 143.9 168.0 182.6 Gms. 2.37 2.27 2.40 2.68 2.63 2.49 2.85 2.85 2.85 2.67 2.90 2.63 2.56 2.37 Gms. 183.4 161.7 142.9 152.6 153.0 137.7 Gm 2.80 Average . 135.1 2.10 165.9 2.61 155.2 2.54 Gms. Gms. 169.2 133.0 139 . 2 144.7 149.9 2.37 2.39 2.48 2.51 2.60 147.2 2.47 Experi- ment num- ber. Subject and duration of experiment. Fifth day. ■H O+S S to U 4) bti> o T « >> "oS Mrs +s *- 8 O 43 & H Ph Sixth day. Seventh day, CO o H £ to 03-^ - ID .2 ' M-Z3 >*8 Ph o 5 to U 4) of — >. Mrs Ph 73... 75 . . . S.A.B., Jan. 28 to Feb. 1, 1905 S.A.B., Mar. 4 to 10, 1905 . . . Average Gms. 148.1 144.7 146.4 Gms. 2.68 2.54 2.61 Gms. 129.8 Gms. 2.30 129.8 2.30 Gms. 132.5 Gms. 2.36 132.5 2.36 There is good reason to suppose that the slight muscular activity exhibited by the subjects of these experiments could not have had any marked influence on the protein katabolism. Even slight variations in muscular activity, how- ever, have considerable effect upon the amount of fat katabolized, hence it is Katabolism of Glycogen. 463 necessary to consider the fat katabolism along with the variations in muscular activity in order to obtain an intelligent conception of the causes for the variations in the results for fat. In a subsequent section the relation between the muscular activity and protein, fat, and carbohydrate katabolism, along with a number of other factors of metabolic activity, is considered. KATABOLISM OF GLYCOGEN. In all respiration experiments thus far made, in which the total income and outgo for 24 hours has been determined, it has been assumed that the store of glycogen in the body remains constant during the experiment, or if there are fluctuations during the day, that the amount in the body at the beginning and end of each experimental day remains unchanged. For experiments with normal diets in which a maintenance ration is supplied, this assumption may not be entirely erroneous. The study of fasting katabolism, on the other hand, involves such abnormal conditions that the assumption that the glycogen con- tent of the body remains constant is hardly justified. With the development of the present form of apparatus in use in the laboratory of Wesleyan Univer- sity, it was possible to obtain direct evidence regarding the absolute amount of oxygen absorbed by man per day. It has recently become possible to compute the loss of body material, not only the amounts of protein and fat, but also the amount of glycogen, and in these experiments the quantities of glycogen katabolized per day, have been computed. All earlier experimenters assumed that the total carbon elimination minus the carbon of the katabolized protein, was derived from the katabolism of fat, but accurate measurements of the oxy- gen intake permit an apportionment between the fat and the glycogen of the carbon eliminated other than carbon of katabolized protein. Food was administered in the first series of experiments with which this apportionment was attempted,101 although in some instances in amounts con- siderably less than that required for maintenance. The first observations regarding the glycogen katabolism of fasting man made with this apparatus are here recorded. The method of computing the amount of glycogen has been elaborated in connection with experiment No. 59, and the possible errors of the method (and they certainly exist) have been pointed out in a discussion elsewhere.152 Eecognizing, then, the possibilities of error in this determination, the results are given as representing the closest approximation to the true glycogen katabolism that, so far as we are aware, has yet been made. The respiratory quotient commonly used for determining the apportionment of the combustion between fat and glycogen is not, at least in its ordinary sense, here used. 151 U. S. Dept. Agr., Office of Expt. Sta. Bui. 175. 152 Carnegie Institution of Washington Publication No. 42; U. S. Dept. Agr., Office of Expt. Sta. Bui. 175. 4G4 Influence of Inanition on Metabolism. The computations are based on the loss to the body of nitrogen, carbon, hydro- gen, and oxygen, the elementary analyses furnishing the data whereby these losses are determined. The amounts of glycogen katabolized per day together with the quantities per kilogram of body-weight are recorded in table 228. Table 228. — Glycogen katabolized in metabolism experiments without food. Ex- peri- ment num- ber. Subject and duration of experiment. First day. Total. Per kilo- gram of body- wt. Second day. Total. Per kilo- gram of body- weight. Third day. Total. Per kilo- gram of body- wt. Fourth day. Total. Per kilo- gram of body- wt. 59.. 68.. 69.. 71.. 73.. 75.. 77.. 79.. 80.. 81.. 82.. 83.. 85.. 89.. B. F. D., Dec. 18-20, 1903. . . A. L. L., Apr. 27-28, 1904. . . A. L . L. , Dec. 16-19, 1904 . . . S.A.B., Jan. 7-10, 1905 S. A. B., Jan. 28-Feb. 1, 1905 S. A. B., Mar. 4-10, 1905.... S.A.B., Apr. 8-11,1905.... H.E.S.,Oct. 13-14, 1905... C.R.Y., Oct. 27-28, 1905... A. H. M., Nov. 21-22, 1905. . H. C. K., Nov. 24-25, 1905. . H. R.D., Dec. 5-6, 1905. ... N.M.P., Dec. 9-10, 1905.... D.W., Jan. 10-11,1906 Gms. 89.2 112.5 103.8 181.6 135.3 64.9 92.7 117.6 103.6 28.7 165.6 32.8 146.0 165.6 Average 110 . 0 Gms. 33 56 41 15 31 09 1.51 2.08 1.51 .47 2.33 .59 2.18 2.12 Gms. 59.4 72.6 31.5 29.7 18.1 23.1 14.9 40.0 17.1 '25.7 44.7 41.6 91.6 39.6 1.69 240. 3 Gm s. 0.90 1.03 .43 .52 .31 .39 .25 .72 .26 1 .43 .64 .76 1.40 .51 Gms 4.2 20.62 32.7 22.0 7.4 5.4 58.9 21.8 Gms 0.06 Gms. Gms. .46 .39 .13 .09 1.00 15.30.21 25.3 .46 21.6 .39 25.2 29.2 .44 .51 0.36 23.30.40 Ex- peri- ment num- ber. Subject and duration of experiment. Fifth day. Total. Per kilo- gram of body- weight. Sixth day. Total. Per kilo- gram of body- weight. Seventh day. Total. Per kilo- gram of body- weight. 73... 75... S. A. B., Jan. 28-Feb. 1, 1905 S.A.B., Mar. 4-10, 1905.... Average Gms. 40.8 8.2 8.2 Gms. H).20 .14 20.14 Gms. 21.' 7 21.7 Gms. 6.38 Gms. 18. '7 0.38 18.7 Gms. 0^33 0.33 1 Glycogen gained. 2 Omitting glycogen gained. The figures above show in a striking manner the great possibilities of error involved in the assumption made in all earlier fasting experiments with men, that the store of glycogen in the body is not materially drawn upon during the earlier days of fasting. On the first day of fast the amounts katabolized varied in these 14 experiments from 28.7 grams to 181.6 grams, and on the IvATABOLISM OF GLYCOGEN. 465 average for 14 experiments there were 110 grams of glycogen katabolized on the first day. The quantities per kilogram of body-weight on the first day show marked differences since the variations in body-weight of the different subjects were by no means as great as the fluctuations in the actual amount of glycogen katabolized. On the second day of fasting, which, as has been seen in the previous discussion, appears to represent more nearly than the first day, the true fasting metabolism, the quantity of glycogen is considerably less, ranging from 14.9 to 91.6 grams. It is to be noted that in one of these experiments, No. 81, the records indicate a gain of glycogen amounting to 25.7 grams. Similarly a gain is recorded on the fifth day of experiment No. 73. Deferring for the moment the discussion of these gains of glycogen, it is seen that the amount of glycogen katabolized in the average of 13 experi- ments on the second day was 40.3 grams. On the basis of per kilogram of body-weight 0.62 gram was katabolized. The large amounts on the first day of fasting are thus very materially reduced on the second day. Unfortunately, but few experiments continued after the second day, the average amount of glycogen for the third day of 6 experiments being 21.8 grams. Wide fluctu- tions appear, however, even on this day, ranging from 4.2 to 58.9 grams. On the fourth day the glycogen katabolized was nearly constant in the 5 experi- ments, averaging 23.3 grams. The quantities katabolized on the fifth, sixth, and seventh days are somewhat lower, the amounts showing a general tendency to decrease as the fast progresses. The lowest result is noted on the third day of the longest experiment, while on the fourth, sixth, and seventh days, the total amounts katabolized are nearly equal to those of the second. Marked irregularities are to be noted in the katabolism as the fast progresses in practically all the experiments. Storage of glycogen. — On 2 of the 43 fasting days here reported, the figures indicate a storage of glycogen. It was hoped that the data would perhaps indicate the exact source of the stored glycogen, but, unfortunately, the instances of such storage are too few and the quantities stored too small to justify definite conclusions from the results as recorded, and, accordingly, these experiments fail to throw important light on the much discussed question regarding the source of glycogen. It may be questioned whether the analytical methods and the computations used in these experiments are sufficiently accurate to warrant the belief that there is an actual storage of glycogen in these two instances. But a close examination of the figures will show that the oxygen consumption and carbon dioxide output may well include a cleavage of protein or oxidation of fat to form glycogen.153 It is much to be regretted that definite experiments to study this problem with this apparatus have not as yet been reported, although at the 153 See discussion of probable error in oxygen determination on one of these days, p. 514. 30 4f>6 Influence of Inanition on Metabolism. moment of writing experiments are in progress which will, it is hoped, con- tribute to the discussion of this matter. Total glycogen content of the body. — The assumption has commonly been made that the total glycogen content of the body is not far from 400 grams. By means of phloridzin, the glycogen of the body of a dog has been rapidly driven out in the urine, and the data thus obtained furnish information regarding the glycogen content of the animal under experimentation. With man the estimations are necessarily founded on a much less scientific basis. The data here presented give some evidence regarding the quantities of glycogen in the body since the total amounts katabolized during varying periods of inanition have been computed. The greatest amount katabolized during the first 24 hours without food was, as has previously been stated, 181.6 grams. The greatest output measured during a 2-day fast is that of experiment No. 85, namely, 238 grams. The largest amount recorded at the end of 3 days of fasting is that in experiment No. 71, 233 grams, a little less than that of the 2 days of experiment No. 85. For 4 days the largest recorded amount (259 grams) is in experiment No. 71. This amount is, as a matter of fact, not even exceeded by the total glycogen katabolized in the 7 days of experiment No. 75. Since it is highly probable that only a moderate portion of the total glycogen of the body is oxidized in a fasting man during a period of inanition no longer than 4 days, it would appear that the estimate of 400 grams of glycogen for the content of the body is, if anything, too small rather than too large. WATER. The factors involved in a complete study of the income and outgo of water during inanition are the drinking-water and the water of urine, respiration and perspiration, and feces.154 In striking a water balance, the income, namely, water of drink, is deducted from the outgo, i. e., water of respiration and perspiration, urine and feces. There is one factor affecting this balance, however, which must be taken into consideration, namely, the portion of the water of outgo which is not preformed water but represents the water of oxidation of the amounts of protein, fat, and glycogen katabolized. As the result of the chemical transformations in the body, there is an actual formation of water from the oxidation of organic hydro- gen of body material katabolized, and hence the output of water may be said to consist of two fractions : first, preformed water, i. e., that taken in the drink or abstracted from the body tissues and fluids; and second, the water of oxi- dation of organic hydrogen. In experiments of the nature of these here re- ported, where a complete balance of intake and outgo is attempted, the data are available for computing not only the loss from the body of preformed water, but also the amount of water resulting from the oxidation of organic hydrogen. 1M Special treatment has been accorded the feces in the fasting experiments and therefore the water of feces is not discussed here. See p. 120. Water. 467 The wide variations in the amounts of drinking-water consumed by the different subjects of these experiments have been discussed in detail in an earlier section (see p. 3-18). The actual amounts consumed per 24 hours have also been recorded in table 193. Similarly, the output of water of respiration and perspiration has received special discussion in a preceding section, as has also the amount of water in the urine. It remains, therefore, to consider in detail the amount of water of oxidation of organic hydrogen and the loss of preformed water. Water of oxidation of organic hydrogen. — The water resulting from the oxidation of organic hydrogen may be directly computed from the data for the quantities of protein, fat, and glycogen katabolized and the analysis of the urine. The percentages of hydrogen in these compounds are known (see p. 37), and the total quantity of organic hydrogen contained in them may be computed. By deducting the organic hydrogen in the solid matter of the urine, the total hydrogen oxidized is obtained. From these data in the different experiments, the results for the water of oxidation of organic hydrogen as recorded in table 229 may be obtained. The computation actually used in obtaining these results was slightly different in form. In the calculations of the quantities of protein, fat, carbohydrate, and water katabolized, according to the method of simultaneous equations (see p. 38), the values found for water represent only the preformed water involved in the katabolism, for the organic hydrogen of the protein, fat, and glycogen had already been appor- tioned among the various quantities of these three compounds. Deducting the preformed water, as computed by the formulae, from the total water output, gives the values for the water of oxidation of organic hydrogen. The water of oxidation of organic hydrogen is dependent upon the amounts of protein, fat, and glycogen katabolized, and in nearly every case the amounts decrease as the fast progresses. The largest amount in the fasting experiments occurred on the second day of experiment No. 82, and the smallest amount on the seventh day of experiment No. 75. The average water of oxidation for all the fasting experiments is 209 grams, corresponding to the oxidation of 23.4 grams of organic hydrogen. The factors affecting the amounts of protein, fat, and glycogen katabolized, therefore, obviously affect the amounts of organic hydrogen oxidized. Each gram of organic hydrogen oxidized in food or body material is accom- panied by a total liberation of about 70 calories of energy.155 The comparison of the organic hydrogen oxidized in these experiments with the total heat output is deferred to the discussion of the heat elimination. Loss of preformed water. — While the water of oxidation of organic hydrogen is directly proportional to the amounts of hydrogen in the protein, fat, and glycogen katabolized, and is, therefore, not properly to be considered in a water 155 See the computations made by Magnus-Levy. Physiologie des Stoffwechsels (1905), p. 424. Thus during fasting 32 grams of hydrogen corresponds to a total heat output of not far from 2300 calories. 468 Influence of Inanition on Metabolism. balance, an entirely distinct set of factors determines the loss of preformed water. The computation of the output of preformed (katabolized) water has already been discussed. Deducting from this the water of drink, the loss of preformed water is obtained. The amounts of this loss from the body during inanition are given in table 229. In one instance there is an actual gain of preformed water to the body noted, namely, on the first day of experiment No. 75. The most striking feature of these results is the very large amount of water lost during inanition. It is to be remembered that all these subjects were supplied with drinking-water, and hence it is fair to assume that the needs of the body, so far as thirst would dictate, could be fully met. It is furthermore to be noted that in some instances, the subjects actually drank enormous volumes of water, so it is probable that the body was liberally supplied, and yet in a majority of instances the subjects continually lost preformed water from the body. The source of this loss is difficult to determine. It has generally been con- sidered that the katabolism of protein results in the breaking down of flesh which contains a considerable amount of water. Assuming that each gram of protein is combined with water to form 4.9 grams of flesh, the total amount of water resulting from the katabolism of the flesh on each day of the experi- ments may be determined by multiplying the weight of protein by the factor 3.9. The amounts of water thus computed are recorded in column a of the table. If it be assumed that in the katabolism of the various amounts of flesh, the water was liberated and excreted, it is seen that in a majority of experi- ments the water from the katabolism of flesh is but a small part of the total loss of preformed water. Especially is this true in a number of the shorter experiments. The figures show, however, no relation whatever between the quantity of protein and flesh katabolized and the loss of preformed water, and we are forced to the conclusion that if the water of the flesh is excreted as preformed water, it forms but a small portion of the total, especially in the first days of inanition. Thus it would appear that, at least in short experi- ments during inanition, there was a gross error involved in the calculations of the loss of water frequently made on the basis of flesh katabolized. It is clear, therefore, that in the earlier days of fasting, the body may lose very much larger amounts of water than are represented by the water of flesh katabolized. A comparison of the amounts of fat and carbohydrate katabolized with the loss of preformed water on the different days is likewise inconclusive. It is conceivable that the glycogen in the liver and muscles holds a certain amount of water as water of hydration,158 but there is as yet no evidence to warrant this assumption. In certain of the experiments it may appear that the large amounts of glycogen katabolized were coincident with the large losses of preformed water, but there are a number of marked exceptions to this parallelism and no definite deduction can be drawn. 180 See Magnus-Levy, loc. cit. Water. 469 Table 229. — Distribution of water in metabolism experiments with and without food. Experi- ment number. Subject and date. (a) Water in flesh (protein X3.9). (6) Loss of preformed water. (c) Loss of preformed water greater + or less — than water in flesh (fi-a). (d) Water of oxidation of organic hydrogen. 59 Experiments without food. B.F.D. • Dec. 18-19, 1903 Grams. 277 Grams. 905 Grams. + 628 Grams. 229 Dec. 19-20, 1903 330 296 - 34 223 347 404 + 57 222 68 A.L.L. Apr. 27-28, 1904 287 1192 + 905 241 Apr. 28-29, 1904 305 703 + 398 237 69 A.L.L. . Dec. 16-17, 1904 236 286 + 50 221 Dec. 17-18, 1904 334 659 + 325 231 Dec. 18-19, 1904 352 203 — 149 221 Dec. 19-20, 1904 304 386 + 82 213 71 S.A.B. 137 440 -1- SOS 235 Jan. 8- 9, 1905 258 476 + 218 200 Jan. 9-10, 1905 307 564 + 257 191 251 306 + 55 176 73 S.A.B. Jan. 28-29, 1905 241 618 + 377 210 Jan. 29-30, 1905 280 621 + 341 196 270 349 + 79 188 Jan. 31-Feb. 1, 1905.. 243 362 + 119 180 234 300 + 66 170 75 S.A.B. 286 - 48 — 334 195 Mar. 5- 6, 1905 291 564 + 273 189 Mar. 6- 7, 1905 305 529 + 224 187 272 453 + 181 185 Mar. 8- 9, 1905 254 216 - 38 173 Mar. 9-10, 1905 251 255 + 4 166 Mar. 10-11, 1905 237 239 + 2 164 77 S.A.B.: Apr. 8- 9, 1905 206 942 + 736 208 252 965 + 713 202 Apr. 10-11, 1905 257 927 + 670 192 Apr. 11-12, 1005 268 578 + 310 188 79 H.E.S. : Oct. 13-14, 1905 190 661 + 471 220 Oct. 14-15, 1905 336 959 + 623 215 80 C.R.Y. Oct. 27-28, 1905 182 1682 + 1500 217 Oct. 28-29, 1905 233 1365 + 1132 224 81 A.H.M. : Nov. 21-22, 1905 213 753 + 540 186 Nov. 22-23, 1905 305 1082 + 777 179 82 H.C.K. : Nov. 24-25, 1905 220 267 + 47 256 336 1327 + 991 261 83 H.R.D. : Dec. 5- 6, 1905 310 168 — 142 209 Dec. 6- 7, 1905 317 603 + 286 197 85 N.M.P. 266 1010 + 744 240 Dec. 10-11, 1905 266 497 + 231 252 89 D.W. : Jan. 10-11, 1906 234 1056 + 822 248 Jan. 11-12, 1906 338 694 + 356 244 70 Experiments with food. A.L.L.: Dec. 20-21, 1904 305 160 - 145 226 Dec. 21-22, 1904 230 204 - 26 244 Dec. 22-23, 1904 238 -150 - 388 275 72 S.A.B.: Jan. 11-12, 1905 249 38 — 211 184 74 S.A.B.: 251 — 52 - 303 184 Feb. 3- 4, 1905 193 -106 — 299 185 159 — 127 — 286 187 76 S.A.B.: Mar. 11-12, 1905 238 179 — 59 191 Mar. 12-13, 1905 167 -285 — 452 194 Mar. 13-14, 1905 , 183 10 — 173 206 470 I XFLUENCE OF INANITION ON METABOLISM. Knowing that the fat in the body does not hold material amounts of water, it is hardly conceivable that the excretion of preformed water should have any relation to the fat katabolized and an inspection of the figures here given shows that no such relation can be observed. With the enormous amounts of drinking-water consumed bv some of these subjects, it may appear that the loss of preformed water was in a measure influenced by the quantities of drinking-water consumed. Here again com- parisons fail to show any clear relationship. For example, it might be expected that with large amounts of drinking-water, the body would be supercharged with it and thus a minimum loss of preformed water occur, and yet in some of the experiments, where the largest loss occurred, the subjects consumed very liberal amounts of drinking-water. Although no relation can be clearly seen between the quantities of protein, fat, and glycogen katabolized and the loss of preformed water, there is one factor which has previously been treated in this report, which apparently has some relation to this loss. That factor is the ratio between the amount of water in urine and the water in drink, discussed on page 348. This ratio has been given in column e of table 193. Comparing the ratios in this column with the losses of preformed water, it is noted that when there is a high ratio, there is a large loss of preformed water. This is to be expected since the quantities of water of respiration and perspiration remain relatively constant throughout the fasts. An examination of the figures for the longer fasting experiments shows that, in general, the loss of preformed water becomes less and less as the fast pro- gresses, and from a comparison of the amounts of water lost with the water of flesh katabolized in the case of experiment No. 75, it might appear that during the earlier days of fasting, there is a very considerable loss of pre- formed water to the body other than that of the protein katabolized, and that as the fast progresses this extra loss of water diminishes until, on the sixth and seventh days of fasting, the water of flesh katabolized corresponds to the preformed water lost. Apparently, at the beginning of the fast there is no connection whatever between the preformed water lost and the other factors of katabolism. It seems, therefore, that the body must have a large residuum of water other than that in muscle and glands. In the muscles, the ratio of the organic matter to the water is presumed to be very fixed. Indeed, until recently I5r it was believed to be impossible to separate the juice from fresh muscle material by even the highest pressures. Evidently there is a large amount of preformed water in the body aside from that in the protein of flesh or gland. It must be borne in mind, how- ever, that these conclusions are drawn from only one experiment, although it 157 O. v. Piirth, Beitrage zur chem. Physiol, u. Pathol. (1903), 3, p. 543; Schmidt- Nielson, Beitrage zur chem. Physiol, u. Pathol. (1903), 4, p. 182. Water. 471 should also be added that the results on the fifth day of experiment No. 73 indicate the gradual approach of the loss of water to that of the water in the flesh. The results of the fifth day of experiment No. 75 show a loss of pre- formed water less than that of preformed water in the flesh katabolized, so that the extremely close figures of the sixth and seventh days of the experiment may have been a coincidence. Nevertheless, the general trend of the experi- ments is to indicate an approaching equilibrium toward the end of a prolonged fast. It was thus seen that the excess of water in the body was practically all eliminated by about the fifth day of fasting, and that, thereafter, the loss of water was determined in large measure by the flesh katabolized. It would be interesting to compute the water balance of experiments in which the period of inanition was much longer than any of these here reported. Unfortunately the exact data for such comparison are lacking. As a result of these observations, and specifically the results obtained in experiment No. 75, it may be contended that the preformed water in the body exists in two forms, first, that which has a more or less fixed relation to the quantity of protein, and, second, a residuum of water retained in a manner not as yet clear. However definite the apparent proportion of protein and water in flesh during normal nutrition, Lichtenf elt 158 has shown that, at least with fish, during inanition there is a relatively increased amount of water in the flesh, and consequently the proportions of protein and water in flesh are by no means as fixed as the factors commonly used would imply. These experi- ments with fasting men imply that in addition to the water held as water of flesh, there is a very considerable residuum of water in the body. This resid- uum may be drawn upon during the early days of inanition. The ingestion of large amounts of water does not prevent its loss, although in numerous instances during a long period of inanition, when the amount of drinking-water was greatly increased, there was a marked retention of water by the body. Several experiments point toward this view. For example, on the tenth and thirteenth days of Succi's fast in Naples, there were large quantities of water consumed with a very small elimination of urine.159 It is reasonable to sup- pose that the muscular activity on these days was not greatly in excess of that on the preceding days, and he must have stored considerable amounts of water in the body. This observation has likewise been noted in a number of instances in experiments immediately following a fast. It is unfortunate that the exact data have been recorded in no instances and the comparison is at best unsatisfactory. If water is stored in the body during 158 Loc. cit. 159 In striking contrast to these results is the observation of Brugsch (12) that, while the drinking-water and urine excretion of Succi during the Hamburg fast had been constant at 750 cc. and 600 cc, respectively, on the thirteenth day the water of drink was increased to over 1000 cc. and simultaneously the urine increased by about a corresponding amount. 472 Influence of Inanition on Metabolism. a fast it is clear that the tissues of the body have the power of absorbing and retaining water other than as water of flesh. Contrary to this view is the fact observed that there are no material variations in the density of the blood, and the exact place where this excess water accumulates is as yet unexplained. If the deductions based upon experiment No. 75 are correct, it is especially interesting to note that, of all the factors of katabolism during inanition, the excretion of residuum water, other than that combined with the flesh, alone ceases on the sixth day. The katabolism of protein as shown by the experi- ments with Succi may continue for 30 or more days. Certainly there is a large amount of fat left in the body, even after prolonged inanition, and ex- periments on animals would imply that there are also small amounts of glyco- gen remaining in the body after prolonged inanition. Similarly, the skeleton may be drawn upon persistently throughout the whole of the fast, but the re- sults of this 7-day experiment suggest that the residuum or excess water in the body is wholly eliminated by the end of a 5-day fast. Effect of inanition on the proportion of water in the body. — Since the total katabolism is measured in these experiments, it is of interest to compare the losses of solid matter with those of water. From the data regarding the amounts of protein, fat, glycogen, and ash katabolized and preformed water lost from the body, the proportion of water to total solid matter leaving the body during inanition may be computed. In table 230 are recorded the weights of solids katabolized, i. e., the amounts of protein, fat, glycogen, and ash, the loss of preformed water, and the per cent of the total loss due to water. The data are given only for the longer fasts. The total weight of solid material katabolized may vary from 217 to 341 grams, while the loss of preformed water in these experiments ranges from 203 to 965 grams. In one instance, namely, on the first day of experiment No. 75, there was an absolute gain of preformed water to the body. The total loss may be as high as 1236 grams. An examination of the figures in the last column of the table shows that the per cent of total loss due to water varies within considerable limits, the lowest, 41.1 per cent, being on the third day of experiment No. 69, and the highest, 78.6, on the second day of experiment No. 77. The daily averages for the 5 experiments given in the table are 54.4, 62.0, 63.2, 59.6, and 75.3 per cent, respectively. Since it is commonly assumed that the body contains approximately 60 per cent of water, it is seen that the proportions of water to solid matter in the material lost are not widely different from those occurring in the body. In experiment No. 69, the per cent of water lost is much lower than the average (60 per cent) and in experiment No. 77 the loss is much larger (75.3 per cent) . The fact that there was an actual gain of preformed water on the first day of experiment No. 75 would lower very considerably the proportion of the total loss due to water. Water. 473 If the weight of the skeleton be deducted from that of the body the pro- portion of water in the flesh and other tissues is considerably greater than 60 per cent. Assuming such an increased percentage, from the relative amounts of water lost during inanition it would appear that the observations Table 230. — Proportion of total loss of flesh due to preformed water. Experiment number. Subject and day of experiment. Weight of solids kata- bolized. Loss of pre- Totai formed loss. water. Per cent of total loss due to water. 69. 71. 73. 75. 77. A.L.L., First day. . . Second day. . Third day.. . Fourth day. . Average. S.A.B., First day Second day. . Third day . . . Fourth day. . Average. S.A.B., First day Second day. . Third day... Fourth day. . Fifth day . . . Average, S.A.B., First day Second day. . Third day. .. Fourth day. . Fifth day. . . Sixth day. . . Seventh day. Average S.A.B., First day Second day. Third day . . Fourth day. , Average Grams. 306 298 291 268 291 Grams. Grams. 286 592 659 957 203 494 386 654 383 674 Per cent. 48.3 68.9 41.1 59.1 54.4 341 254 249 229 268 M40 780 1476 730 1564 813 306 534 446 714 56.4 65.2 69.3 57.3 62.0 314 248 235 229 225 250 1618 932 1621 869 1349 584 362 591 300 525 450 700 66.3 71.5 59.8 61.2 57.3 63.2 271 263 243 247 226 222 217 240 2-48 564 816 529 771 453 700 216 442 255 476 239 456 s376 3 610 69.1 68.6 64.7 48.9 53.6 52.4 3 59.6 294 263 273 257 272 1942 1965 927 578 853 1236 1228 1200 835 1125 76.2 78.6 77.3 69.2 75 . 3 1 Does not include water of feces. 2 Gain. Water of feces not taken into account. 8 Average of 6 days. of Lichtenfelt 16° on the flesh of fish during inanition are likewise true for the flesh of fasting man, i. e., inanition results in an increase in the proportion of water in the body. 160 Loc. cit. 474 Influence of Inanition on Metabolism. ENERGY. In common with all living organisms, fasting man is constantly producing and giving off heat as a result of katabolism. The amounts of heat produced, the source of the energy, the factors influencing heat production, and all allied topics have an unusual interest when studied with fasting man. Likhachev 161 experimented on a man fasting for 24 hours in the Pashutin respiration apparatus modified so as to permit the direct measurements of heat. It has been necessary in all other inanition experiments to compute the energy transformations from the data for the total katabolism, and since there was no means of ascertaining whether the katabolized material other than protein consisted of fat or glycogen, it was assumed that it was all fat. Hence, the computations of the energy transformations were subject to all the errors incidental to the determination of the amounts of katabolized protein and fat. The measure of the protein katabolism was essentially that used in the present day, but we have seen from the foregoing discussions that the amounts of fat katabolized as computed from the total carbon output and the carbon of pro- tein are materially different from those determined by the present method, in which the direct measurement of the oxygen consumption permits an apportion- ment of the total katabolism as protein, fat, and glycogen. The oxidation of one gram of carbon in the form of glycogen gives rise to much less heat than the oxidation of one gram of carbon in the form of fat, and hence the energy transformations computed on the assumption that only fat and protein were burned in the body must of necessity be erroneous. In the experiments here reported, not only were the total amounts of katabolized protein, fat, and glycogen computed, but the type of apparatus also permitted direct determinations of the amounts of heat eliminated. Since there were varying amounts of heat residual in the body at the end of the different days of the different experiments, corrections were applied to the heat elimination to obtain the actual heat production. (See discussion, p. 46.) For the purpose of comparing the estimated energy derived from material oxidized in the body, the heat production rather than the heat elimination must be used. Heat elimination will first be considered in the following discussion, and then the heat production. Aside from the kinetic energy leaving the body there are considerable amounts of potential energy excreted in the unoxidized material of the urine. For this reason, a particular section of the report is reserved for the discussion of the energy of the urine. HEAT ELIMINATION. The calorimetric features of the respiration calorimeter were devised for the special purpose of measuring directly the heat eliminated by man. The tests of the accuracy of the apparatus have been numerous and severe, and in practically every instance, the results obtained have been all that could 181 Dissertation, Russian (1893), St. Petersburg. Energy. 475 be desired, indicating an error of somewhat less than 1 per cent. Difficulties attending the control of the temperature of the room in which the experiments have been made and minor discrepancies in the manipulation of the apparatus by assistants with varying degrees of skill have undoubtedly introduced errors of slight magnitude that are not compensated and are not absolutely known. It is believed that in a large majority of the experiments here reported, the accuracy of the heat measurement is well within 1 per cent. In some instances the error may be greater than this but in no case could it have amounted to 5 per cent. Heat can leave the body in a number of different ways. The larger portion of the heat is lost by radiation and conduction; a certain amount is used to warm the inspired air; the urine and feces carry away measurable amounts of heat; finally large quantities are necessary to vaporize the water eliminated from the lungs and surface of the body. Aside from the measurement of these factors, there are the numerous corrections which have to deal with the actual heat production during the period of the study, corrections which receive spe- cial subsequent treatment. The heat lost by radiation and conduction, together with that required to warm the inspired air is measured by the amount of water passing through the heat absorbers and the temperature through which this water is raised. The amount of heat thus measured, however, includes also the heat lost from the urine and feces that are allowed to cool to the temperature of the calorimeter before being removed. From the weights and specific heats of urine and feces and the difference between the body temperature and the temperature of the calorimeter, the amount of heat lost by this means can be computed. It is recorded for the experiments here published in table 231, column c. The amount of heat required to warm the inspired air from the temperature of the calorimeter to that of the body may be obtained by the use of four factors: the total ventilation of the lungs, the weight of a liter of air, the specific heat of air, and the difference between the temperature of the calorimeter and that of the body. The volume of the inspired air has been computed and the total ventilation of the lungs is given in column a of table 220. Since these results are recorded in liters reduced to standard conditions of temperature and pressure, the weight of air warmed is obtained by multiplying the total venti- lation by 1.293, the weight of a liter of air at standard conditions of tempera- ture and pressure. From the weight of air, its specific heat16a and the difference between the body temperature and that of the chamber, the amount of heat actually required to warm this inspired air may readily be computed. Deducting from the total heat brought away by the water current the heat from the feces and urine and that required to warm the inspired air, leaves the heat of radiation and conduction, which is recorded in the first column of table 231. 182 The specific heat of air has been taken as 0.237. ■A76 Influence of Inanition on Metabolism. o e OB S o •*» © o v. ■a s <3 ■«§ a S 49 5 §§ Sts O (S OS £ eo § 8 « s <» •St o rO ■a <» •*» e s to CO a u a o JS as u o o a o o ft o >> •a u o h 3 —•£00 >o a .2 00 03 * *a "^ Cj_. 83 O a ?° £5 a fl^ ® M 3 83 » 1 o « . CD • a1 ^ p,83 to I 1 o ^.Sg-a 3 rj o o +3 03 -a •a a 83 +» O » .O 3 02 asa x a s w a ■►."coco-* eoco coiocbcs "#ci5ff}Oi moi-*r-it— oo-*cocot-mio| ~ C-J II Cl j ci oi ri e-i »i »j _°^ •COr-l-* t-CO ©X(M5-^»Ct CO-^COt' -^4--^-«#10IO -^^^--tS 0 »0 . 5"» ■*** Y) o HO CO CO L- CO ^HOICOO Ot-OKC"! X '/. xiC-4"T!:i -^lO^JH^ -^COCOCO ^CCCCCOCO Ct CC- CO CC CC CO CO .00 CO lO tfjXCO tr CO CO CO 6 CO'* »Ol.«HM OOlOOiCO XlOOXX OJHOCOt-HN HWWN C\t C-l 1— 1 1— 1 O-i f- ( -— 11— 1 1— ( (MCINHHHH a t- 10 t-o»« a CO 10 to 05 lO tj OO ^1— I H 1—1 o CO-* too tot- t-CO iO C-"! CO CO C© 1— ( ■^XWjO Oi CO o to X '^COCO'M CO CO CO Cv» 1— I to »— I to >— < ^h CO iO ^^)CiOiOXX CO CO CO CO U <— ( r-i — • CO CO CO C:ooo *C5 OS OS 00 05 05 »r> o us 10 0000 ~- ~. ~- ~ JCkOlO o o o T OS 05 Oi • o* »OiOiOiD»OiO>0 O O — ' — ■ o o o CT> OS OS OS OS 05 OS rC: 1 1 i •WXOSO COOS (MCI tLoo C-JCM XCSOrt t- coos O CD O <-H rZ^ CQ CI CO CO hH , 1/ 1 I 1 cc CO o -— 1 •— 1 C-2 CM CO CO lO tO t- CC OS O rH fll— 1 ^ iO tO t- CO OS O 42 © © a) a. C5 ft ft 0000 H O CD Q PQQQ - 1-3 a a a a 83 cS o3 83 C9 02- a a a a-o 03 o3 03 03 P i-si-aT-si* 02 u u u u u u t-> 83 03 03 03 83 83 03 ea Energy. 477 W rl •a IN 0) CO a OS CJ) CS a o j - s Influence of Inanition on Metabolism. The heat of vaporization of water is taken as 0.592 calorie per gram, and by means of this factor and the weight of water vaporized from the body, the energy absorbed by the water is computed. The weights of water vaporized from the lungs and skin are recorded in columns d and e of table 220, and the energy absorbed in their vaporization is recorded in columns d and e, of table 231, the total heat required to vaporize all the water being recorded in column /. The total heat elimination for the different days of the experiment is recorded in column g. It is to be noted that these values are not identical with those given elsewhere for the total heat production, and it must be borne in mind that heat elimination, as distinguished from the heat production, is here under discussion. The proportions of the total heat eliminated from the body in the various ways have been computed and recorded in columns h to m inclusive. From these data, it may be seen that not far from 75 per cent of the total heat eliminated leaves the body by radiation and conduction; 2.3 per cent is utilized in warming the inspired air; as a rule less than 2 per cent is given up by the excreta; about 10 per cent is required to vaporize the water from the lungs; and 12 per cent is required to vaporize the water from the skin. The individual fluctuations from these averages are noticeable chiefly in the heat of urine and feces, which is to be expected from the marked variations in the volume of urine excreted. All the other factors, while varying somewhat, are relatively constant, and the average of these experiments with fasting men at rest shows that 1440 calories are eliminated by radiation and conduction, 44 calories are required to warm the inspired air, 22 calories are given off by the urine and feces, 424 calories appear in the water vaporized from the lungs and skin, the total average heat elimination being 1931 calories. No noticeable abnormalities are to be observed in the similar data for the experiments with food. HEAT PRODUCTION In the katabolism of protein, fat, and glycogen during inanition, heat is produced. A comparison of the heat production with the katabolism can not be made by simply measuring the heat lost from the body through radiation, conduction, and vaporization of water. The principles involved in the compu- tation of this production as distinguished from the heat elimination have already been discussed in considerable detail in connection with experiment No. 59 (see p. 46). It is only necessary to call to mind here that changes in body-weight and body temperature are theoretically at least of great importance in determining the heat production as distinguished from the heat elimination. As the fast progresses, the body loses weight. The material lost is cooled from the temperature of the body to the temperature of the calorimeter, but the Energy. 479 heat given off by the cooling of this material does not represent heat actually produced from katabolism. The discussion which follows deals specifically with heat production and takes into account, therefore, the heat elimination corrected for energy gained or lost because of changes of body- weight and body temperature. The results for the measured heat production are recorded for all experiments without food in table 232, together with the distribution of rates and proportions over the main periods of the day. By a comparison of the first column of table 232, which gives the total heat production in 24 hours, with column g of table 231, which gives the total heat elimination, some conception can be had of the variations due to the corrections for changes in body temperature, body-weight, etc. It is surprising how slight is the variation between the different experiments. Thus, the average heat production for all the fasting experiments is 1924 calories, while the heat elimination for the same series is 1931 calories. It might seem that the heat elimination may be taken as the measure of the heat production, and indeed in fasting experiments the correctness of this assumption is fairly well estab- lished, since the corrections for the heat production are more or less compen- sating. When the data for shorter periods are desired, however, it is of extreme importance that the heat production rather than the heat elimination be used. The heat production and heat elimination in experiment No. 59 were determined not only for the 24 hours but also for the 3-hour periods (see table 17). The differences between the heat production and heat elimination for 3-hour periods are much more marked than for the day. The distribution of the heat production over the 6-hour periods has been computed and given in table 232. The proportion of the total heat in 24 hours produced for each period has likewise been calculated and recorded in the table. As is to be expected the greater amount is produced during the day period. This is in harmony with the conception that muscular activity determines in large measure the heat production, for even though these are distinctively rest experiments, the muscular activity is obviously greater in the day time than at night. The fact that in some of the experiments the subjects spent not a little time sleeping during the day would, however, tend to more nearly equalize the heat production of the first 12 hours of the day with the last twelve. Eeference to the detailed records of the body movements will show during which hours the subject was asleep. The results show that the heat during the last period of the night, i. e., from 1 a. m. to 7 a. m. is more nearly uniform on the different days of the fasting experiments than during any other period. However, there is, as a rule, a decrease in heat production during this period of the night as the fast progresses. The irregularities in muscular activity in the day time render any comparison of the different experiments, and indeed different days of the same experiment, extremely difficult. It is only by taking into account the total sum of body -1SII Influence of Inanition on Metabolism. Table 232. — Heat production of body, amounts per day and per hour, and propor- tions for different periods — Metabolism experiments without food. Ex- peri ment num ber. Subject and date. Total am'nt in 24 hours. Rate per hour. Day periods. 7 a.m. to 1 p. m. 1 to 7 p.m. Night periods. 7 p.m. to la. m. 1 to 7 a. m. Aver- age for 24 hours Proportion of total for 24 hours. Day periods. 7 a. m. to 1 p. m. 1 to 7 p. m. Night periods. 7 p. m. to 1 a.m. 1 to 7 a.m. 59. 68. 69. 71. 75. B.F.D., 1903: Dec. 18-19 Dec. 19-20 Dec. 20-21 Total Av. per day. A.L.L., 1904: Apr. 27-28 Apr. 28-29 Total Av. per day. A.L.L., 1904: Dec. 16-17 Dec. 17-18 Dec. 18-19 Dec. 19-20 Total Av. per day. S.A.B., 1905: Jan. 7- 8 . . . . Jan. 8- 9 Jan. 9-10 Jan. 10-11 Total Av. per day. S.A.B., 1905: Jan. 28-29 Jan. 29-30 Jan. 30-31 Jan. 31-Feb.l. Feb. 1- 2 Total Av. per day. S.A.B., 1905: Mar. 4- 5 . . . . Mar. 5- 6 Mar. 6- 7 Mar. 7- 8 Mar. 8- 9 Mar. 9-10 Mar. 10-11 Total Av. per day. Cals. 2080 2107 2102 6289 2096 Cals. Cals. 102 95 98 91 97 91 297 277 99 92 2167 2217 4384 2192 1951 2163 2035 1958 8107 2027 1970 1844 1746 1606 3166 1792 1866 1791 1739 1663 1548 8607 1721 Cals. 79 87 83 Cals. 71 75 79 Cals. 87 88 88 Per ct, 29.4 27.9 27.7 Per ct, 27.4 25.9 26.0 249 83 225 75 263 88 28.4 26.4 Per ct. 22.8 24.8 23.7 23.7 Per ct, 20.4 21.4 22.6 21.5 100 96 93 72 90 27 7 26 6 25.8 98 99 91 82 92 26 5 26 8 24.6 198 195 184 154 182 ,. t .. m 1 99 98 92 77 91 27 1 26 8 25.1 19.9 22.1 21.0 78 93 78 76 91 24.0 28.6 24.0 91 105 89 76 90 25.2 29.1 24.7 92 95 89 63 85 27.1 28.0 26.3 86 89 81 70 81 26.4 27.3 24.8 347 382 337 285 337 • • ■ a 87 96 84 71 84 25.7 28.4 24.9 23.4 21.0 18.6 21.5 21.0 115 85 69 59 82 35.1 25.9 21.0 98 76 68 65 77 31.9 24.8 22.1 83 81 67 60 73 28.6 27.8 23.0 79 71 59 58 67 29.6 26.6 22.1 375 313 263 242 299 • • • • • • • • 94 78 66 61 75 31.4 26.1 22.1 18.0 21.2 20.6 21.7 20.4 94 82 68 66 78 30.3 20. 5 21.9 89 81 66 62 75 29.9 27.2 22.1 85 83 64 58 72 29.3 28.6 22.1 81 75 61 59 69 29.3 27.2 22.1 73 66 62 56 65 28.4 25 . 7 24.1 422 387 321 301 359 ■ a 84 77 64 60 72 29.5 27.0 22.5 21.3 20.8 20.0 21.4 21.8 21.0 1765 89 72 66 67 74 30.3 24.5 22.4 1768 89 76 66 63 74 30.3 25.9 22.4 1797 87 85 63 64 75 29.1 28.4 21.1 1775 87 81 65 63 74 29.4 27.3 22.0 1649 80 76 60 59 69 29.1 27.6 21.8 1553 77 68 58 56 65 29.7 26.3 22 .4 1568 77 71 57 56 65 29.5 27.2 21.8 11875 586 529 435 428 496 • ■ ■ • 1696 84 76 62 61 71 29.7 26.9 21.9 22.8 21.4 21.4 21.3 21.5 21.6 21.5 21.5 Energy. 481 Table 232. — Heat production of body, amounts per day and per hour, and propor- tions for different periods — Metabolism experiments without food — Continued. Ex- peri- ment num- ber. Subject and date. Total am'nt in 24 hours. Rate per hour. Day periods. "a. m. to lp.m. 1 to 7 p.m, Night periods. 7Ptomi lto lam.|7a-m Aver- age for 24 hours Proportion of total for 24 hours. Day periods. 7 a. m. to lp. m. 1 to 7 p. m. Night periods. 7 p.m. to 1 a. m. 1 to 7 a.m. 77. 79. 80. 81. 82.. 83. 85. 89. S.A.B., 1905: Apr. 8- 9 Apr. 9-10 Apr. 10-11 Apr. 11-12 Total Av. per day. . H.E.S., 1905: Oct. 13-14 Oct. 14-15 Total Av. per day. . C.R.Y., 1905: Oct. 27-28 Oct. 28-29 Total Av. per day. . A.H.M., 1905: Nov. 21-22 Nov. 22-23 Total Av. per day. . H.C.K., 1905: Nov. 24-25 Nov. 25-26 Total Av. per day. . H.R.D., 1905: Dec. 5-6 Dec. 6-7 Total Av. per day. . N.M.P., 1905: Dec. 9-10 Dec. 10-11 Total Av. per day. . D.W., 1906: Jan. 10-11 Jan. 11-12 Total Av. per day. . Av. of experi ments with out food. . 31 Cals. 1874 1880 1840 1807 7401 1850 1951 2047 3998 1999 1954 2099 4053 2027 1729 1781 3510 1755 2222 2477 4699 2350 1914 1907 3821 1911 2109 2305 4414 2207 1924 Cals. 90 82 79 75 Cals. 84 85 80 79 326 82 328 82 Cals. 67 72 70 71 280 70 Cals. 71 75 78 76 Cals. 78 78 77 75 300 75 308 77 Per ct. Per ct. Per ct. 28.8 26.9 21.5 26.1 27.1 22.9 25.7 26.1 22.8 24.9 26.2 23.6 • • • • • • • ■ • • • ■ 26.5 26.5 22.7 Per ct. 22.8 23.9 25.4 25.3 24.3 88 79 82 77 81 27 0 24.2 25.2 89 87 84 81 85 26 1 25.5 24.6 177 166 166 158 166 89 83 83 79 83 26 6 24.9 24.9 23.6 23.8 23.6 88 99 75 89 82 81 81 82 81 87 27.0 28.2 23.0 25.4 25.2 23.1 187 94 164 82 163 82 163 82 168 84 27.7 24*1 24.1 24.8 23.3 24.1 81 90 82 69 66 71 59 67 72 74 28.1 30.3 28.5 23.2 22.9 23.9 171 86 151 76 137 69 126 63 146 73 29.3 25.8 23.5 20.5 22.6 21.4 107 lis 88 101 97 106 78 88 93 103 28.9 28.6 23.8 24.4 26.2 25.7 225 113 189 95 203 102 166 83 196 98 28.8 24.2 25.9 21.1 21.3 21 .1 94 98 64 68 89 80 71 72 80 79 29.6 30.8 20.1 21.4 28.0 25.2 192 96 132 66 169 85 143 72 159 80 30.1 20\7 26.6 22.3 22.6 22.6 99 106 84 93 89 99 79 86 88 96 28.2 27.6 23.9 24.2 25.4 25.8 205 103 177 89 188 94 165 83 184 92 27^9 24.1 25.5 22.5 22.4 22.5 2150 2254 80 88 101 105 95 104 82 78 90 94 22.4 23.5 28.2 2S.0 26.5 27.7 4404 2202 168 84 206 103 199 100 160 80 184 22.9 28.1 27.2 22.9 20.8 21.8 90 84 77 80 28.0 26.2 24.0 21.8 482 Influence of Inanition on Metabolism. movements that any intelligent comparison can be made. In fasting experi- ments such as those here reported, in which the subjects were distinctly inactive, the proportions of the total heat for 24 hours are about evenly divided among the periods. The average of all the fasting experiments shows that 28 per cent of the heat was produced between 7 a. m. and 1 p. m., 26.2 per cent between 1 and 7 p. m., 24 per cent from 7 p. m. to 1 a. m., and 21.8 per cent from 1 to 7 a. m. When these men were asleep the heat production per hour averaged 70 calories. Considerable variations in the quantities produced per hour may be observed, not only with different subjects, but with the same subjects in different experiments. The unusually large production of 115 calories of heat per hour during the first period of the first day of experiment No. 71 is due to the fact that during this period the subject rode for 10 minutes on the bicycle ergometer. In no other period with this subject did the heat pro- duction rise to over 98 calories per hour. The lowest heat production recorded in any experiment was 56 calories per hour. This small amount was produced in the last period of the last day of the two longest experiments, Nos. 73 and 75. In explanation of these irregularities reference must be made to the notes in the diaries of the subjects and records of body movements previously recorded and to the estimates of muscular activity given beyond. Heat production per kilogram of body-weight and per square meter of body surface. — The irregularities in the heat production noted in the fasting experi- ments reported herewith may be accounted for in several ways: First, there may be marked differences in muscular activity. This factor receives special discussion elsewhere (see p. 484). Second, there may be a difference in size of the different subjects and consequently the total heat production per kilo- gram of body-weight is of value in comparing the different experiments. For other comparisons commonly made by physiologists, the production per square meter of body surface is likewise of value. The heat produced per kilogram of body-weight and per square meter of body surface has been computed for all of these fasting experiments and is recorded in table 233. The results for the food experiments Nos. 70, 72, 74, and 76 are also included in the table. The average heat production per kilogram of body-weight for the first day of the 18 experiments with and without food was 30.7 calories. The range is from 26.6 to 34.6 calories, but the results in the large majority of the experi- ments were very close to the general average. On the second day the average is over 1 calorie greater, i. e., 31.8. The fluctuations range from 28.6 to 36.9 calories. The average heat production per kilogram of body-weight on the third day is 31.0 calories and on the fourth, fifth, sixth, and seventh days, the averages are 29.6, 28.5, 27.5, and 28.0 calories, respectively. There is, then, a distinct tendency for the heat (per kilogram of body-weight) to decrease after the second day. Energy. 483 Table 233. — Heat produced per kilogram of oody-iveigM and per square meter of body surface in metabolism experiments ivith and without food. Ex per iment num- ber. Subject and date. First day. Second day. Third day. Fourth day. Fifth day. 59. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77.. 79. 80. 81. 82., 83., 85.. 89.. B.F.D., Dec. 18 to 19, 1903. . A.L.L., Apr. 27 to 28, 1904. . A.L.L., Dec. 16 to 19, 1904. . A.L.L., Dec. 20 to 22, 1904. . S.A.B., Jan. 7 to 10, 1905.. S.A.B., Jan. 11, 1905 Per kilo Per sq. meter. . Per kilo Per sq. meter. . Per kilo Per sq. meter. . Per kilo Per sq. meter. . Per kilo Per sq. meter. . (Per kilo ] Per sq. meter. . S. A.B., Jan. 28 to Feb. 1, 1905 3 Per kilo ( Per sq. meter. . S.A.B., Feb. 2 to 4, 1905. .. 3 Per kilo j Per sq. meter. . S.A.B., Mar. 4 to 10, 1905. . J Per kilo1 j Per sq. meter3. 8. A.B., Mar. 11 to 13, 1905 ... J Per kilo I Per sq. meter. . S.A.B., Apr. 8 to 11, 1905... 3 Per kilo ] Per sq. meter. . H.E.S., Oct. 13 to 14, 1905... j Per kilo \ Per sq. meter. . C.R.T., Oct. 27 to 28, 1905... J Per kilo I Persq. meter. . A.H.M.,Nov. 21 to 22, 1905... 3 Per kilo j Persq. meter. . H.C.K.,Nov. 24 to 25, 1905... 3 Per kilo j Persq. meter. . H.R.D., Dec. 5 to 6, 1905. . . 3 per kilo ( Persq. meter. . N.M.P., Dec. 9 to 10, 1905... 3 Per kilo j Per sq. meter. . D.W., Jan. 10 to 11, 1906. . . j Per kilo Per sq. meter. . Average.... iler kil° body-weight ( Per sq. meter body sui Cals. 31.9 1046 31.3 1052 29.8 1009 31.4 1056 32.4 1023 Cals. 31.0 1023 80.1 1017 26.6 903 29.7 9S8 34.2 1072 80.4 940 31.9 1005 30.7 949 29.7 941 31.7 9S4 30.6 980 34.5 1075 28.5 948 2S.1 SOI 31.2 1050 1183 34.6; 34.9 1070 1075 31.1 975 28.6 SS6 29.9 946 31.0 961 31.2 993 36.9 1143 31.5 1037 29.5 939 35.3 Cals. 32.1 1050 28.3 956 34.8 1168 31.0 964 30.6 955 28.9 895 30.8 969 31.2 971 31.2 987 31.5 1039 27.5 945 rface. 30.7 992 35.1 1149 29.3 1011 31.8 102S 81.0 991 Cals 27 924 28 S95 29 922 30 966 31 982 29.6 Cals. 28 866 29 985 28.5 93S 885 1 Values for the sixth and seventh days are 27.5 and 28.0 calories respectively. 2 Values for the sixth and seventh days are 857 and 869 calories respectively. Since the formula for computing body surface is dependent upon body- weight, the fluctuations observed in the heat production per kilogram of body-weight likewise appear when computed on the basis of per square meter of body surface. The average production per square meter of body surface on 48-t Influence of Inanition on Metabolism. the first day of all the experiments was 992 calories. On the second day the average was somewhat higher, 1028 calories, while on the third day, it was noticeably less, 991 calories. The averages for the fourth, fifth, sixth, and seventh days were 938, 885, 857, and 869 calories, respectively. These results show in general a tendency for the heat production per square meter of body surface to diminish after the second day of fasting. The presence of results for food experiments in the calculations (aside from those for experiment No. 70) has no material influence on the trend of the averages. While the con- stancy of the heat production per square meter of body surface in all classes of animals has been emphasized in many discussions, it may be observed that even with fasting man at rest, there are marked fluctuations in this factor, and here again variations in internal and external muscular activity may account for these seeming discrepancies in the heat production per square meter of body surface. MUSCULAR ACTIVITY. This series of experiments was made upon fasting men at muscular rest, and yet as has been repeatedly emphasized in the discussion, there were varying degrees of minor muscular activity during the experiments and it can not be said that the subjects all remained in the same degree of rest. Knowing the marked influence of excessive muscular activity on the heat production as observed in experiments on muscular work, it is important to determine, if possible, to what extent the slight variations in activity observed in these experiments influenced the heat production. Method of estimating energy of external muscular activity. — Aside from the intermittent strength tests with the dynamometer in some of the experi- ments and the 10-minute exercise period on the bicycle ergometer on the first day of experiment No. 71, no measurement of the muscular activity of the subjects was attempted. Our knowledge regarding the energy required to perform the minor movements of the body, even in a so-called " rest experi- ment " is as yet very meager. But since it was important to estimate as nearly as possible the muscular activity on the different days and with the different subjects, a method for estimating these variations in activity has been put in use, although admittedly based on very uncertain evidence. The muscular activity was in general very slight. The visible muscular activity of the subjects was carefully recorded by the observer outside of the calorimeter chamber, the number of times the food aperture was opened and closed was likewise noted on a sheet provided for the purpose, and from the diary kept by tlic subject, records of certain other movements were obtained. These have been combined as has been shown previously in the " body movement " records for each experiment. From these movements the attempt has been made to estimate the muscular activity, expressed in calories. The material used con- Energy. 485 sisted of the records of body movements, the results obtained from a long series of unpublished experiments with different subjects to ascertain the energy required for certain definite motions, and an experiment made to find the energy required to rise from the chair, go to the food aperture, open it, close it, return and sit. Other motions than those which would naturally accompany the act of going to the food aperture were made by the different subjects in the fasting experiments and indeed in considerable number. For the purpose of these estimates, one of the simplest motions, i. e., the act of rising from the chair in which the subject sits, is taken as a unit of movement, and all other movements are expressed in terms of this unit. The actual amount of energy required to go to the food aperture and return has been found from a series of special experiments to be one calorie. In another series, the energy required to undress, be weighed in a hanging chair, dress, and be seated was actually measured. This was found to average 20 calories. The operation at the food aperture was estimated to be 4 times as great as that of rising from the chair. Hence the unit of movement was taken as one-fourth of a calorie. With this as a basis, values were estimated for all the individual movements recorded in the unpublished experiments, the sum of which equaled 20 calories or 80 units. For example, the act of hanging the chair for weighing was estimated to be 5^ units; weighing, 3 units; hanging curtain and taking it down, 2 units; undressing, 16 units; dressing, 25£ units. It is to be noted that a greater total is given for the movements in dressing than in undressing, and estimates were likewise made for different acts in the process of the two operations. Other movements occurring during the experiments are estimated according to their character, such as for example, to rise, weigh, and dress in the morning. To rise from the bed is reckoned as 3 units; hang chair, 5^ units; weigh, 3 units ; remove and reinsert the rubber stopper through which the weighing rod passes to the top of the calorimeter, 1 unit ; weigh clothing, 6 units ; weigh the heat absorbers, 15 units; unhang chair, 5-| units; arrange and fold bed, 5 units; raise table, 3 units; dress (underclothing being already on), 15 units; sit down, 1 unit; total, 63 units. Similarly, to prepare for bed at night, defecate, take the dynamometer test, and numerous other movements are each accorded their different estimated value in terms of these units. In addition to the activity estimated for the first day of experiment No. 71, account should be taken of the bicycle ride during the second (9 to 11 a. m.) period of the day. Previous calibration has shown that each revolution of the wheel of the bicycle ergometer results in the transformation of 0.0233 calorie of heat, and as the wheel made 643 revolutions during the ride, the energy given off equals 643 X 0.0233. The result of this calculation X 5, since the efficiency of the body as a machine is about 20 per cent, = 74.91 calories, the total result for the ride. ■1st; Influence of Inanition on Metabolism. In experiment No. 76, at 8h 2m p. m. March 11, 1905, gymnastics are esti- mated arbitrarily as 30 units. A walk in the chamber at llh 16m a. m. January 8, 1905, in experiment No. 71, is estimated at 10 units. Table 234. — Estimate of visible muscular activity in metabolism experiments with and without food. (a) (ft) Ener- (a) (ft) Ener- Experi- ment Subject and date of Esti- mated mus- gy of esti- mated Experi- ment Subject and date of Esti- mated mus- gy of esti- num- experiment. cular mus- num- ber. experiment. cular mus- ber. activ- cular activ- cular ity. activ- ity. ity. activ- ity. 59.... B.F.D., 1903: Units. Cals. 76 S.A.B., 1905: Units. Cals. Dec. 18-20 182 46 Mar. 11-13 244 61 160 40 166 42 150 38 203 51 68... A.L.L., 1904: 77 S.A.B., 1905: Apr. 27-28 229^ 57 Apr. 8-11 167% 42 265 66 239 60 69.... A.L.L., 1905: 158 40 Dec. 16-19 109 27 215 54 122 31 79 H.E.S., 1905: 128 32 Oct. 13-14 199 50 133 33 174 44 70.... A.L.L., 1905: 80 C.R.T., 1905: Dec. 20-22 129 32 Oct. 27-28 130 33 115 29 131 33 141 35 81 A.H.M., 1905: 71 S.A.B., 1905: Nov. 21-22 160 40 Jan. 7-10 1210 53 131 33 199 50 82 H.C.K., 1905: 175 44 Nov. 24-25 173 43 144 36 179 45 72.... S.A.B., 1905: 83 H.R.D., 1905: 73.... Jan. 11 196 49 Dec. 5-6 199 183 50 46 S.A.B., 1905: Jan. 28-Feb. 1 . . . 174^ 44 85 N.M.P., 1905: 182 46 Dec. 9-10 189 47 203 51 198 50 188 47 89 D.W., 1906: 166 42 Jan. 10 11 156 39 74 S.A.B., 1905: Feb. 2-4 190 201 192 48 50 48 218 55 75.... S.A.B., 1905: Mar. 4-10 176 175 232 224 185 191 160 44 44 58 56 46 48 40 1 No allowance has been made for 75 calories total output due to 10 minutes bicycle ride, 9 to 11 a. m. period. Energy. 48 1 From the above data the estimates given in table 234 were obtained. They are shown both in units and in calories. It must be understood, however, that in obtaining the figures in this table no attempt has been made to estimate the minor involuntary movements, hence the figures obtained give nearer the minimum than the maximum. The figures recorded in column b of table 234 are admittedly but approxi- mate estimations. Perhaps the most striking feature of the table is the fact that the differences between the experimental days when computed on the basis of calories are, relatively speaking, so small. The maximum difference observed in any fasting experiment is that of experiment No. 77, where there were 60 calories for the second day and 40 calories on the third, and yet the difference here recorded, 20 calories, is barely over 1 per cent of the average total heat production per day of this experiment. A comparison of the varia- tions in muscular activity as expressed in calories given in this table with the total heat production on the succeeding days of each experiment shows that, while the fluctuations in the estimated energy of the external muscular activity are very small, they generally follow those appearing in the total heat produc- tion. That the differences in the heat production noted from day to day in the different fasts, however, can be directly caused by variations in visible exter- nal muscular activity is very completely disproven. A careful revision of the estimates given in this table has been made and it is certain that although there may be errors in the apportionment and the estimate of value of the activity, the error can certainly not be over 100 per cent and it is probably very much less. Assuming that an error of 100 per cent exists and that the error is always in the same direction, so as to increase the apparent differences on different days, the amounts even then are far too small to account for the absolute differences in the total heat production. It is clear, therefore, that while the variations in muscular activity are proportional to the variations in the total heat produced, some factor other than external muscular activity must account for the wide variations in the total heat production. Relations of internal muscular activity to total heat production. — Aside from the external muscular activity, there is a considerable amount of muscular work which can be conveniently termed " internal " muscular work, typified perhaps by the work of respiration and circulation. Estimates, which for the most part are based on the respiratory exchange, have been made of the energy required for circulation and respiration. Eecently Loewy & v. Schrotter 163 have made an extensive study of these factors of internal work in man. From the results of their experiments they conclude that the total energy required for circulation is 3.6 per cent of the total energy transformation of a man at rest. The work of respiration is 2.4 times that of circulation, and the work of respiration and circulation combined amounts to about 13 per cent of the total energy output for the day. 163Zeit. f. exper. Path. u. Therapie (1905), 1, p. 197. 488 Influence of Inanition on Metabolism. The arterial tension as well as pulse rate per minute are important factors in estimating the work of circulation, but unfortunately all attempts to measure accurately the blood pressure in these experiments failed.164 It is much to be regretted that in the longer experiments, which are of the greatest value for comparing the variations in the total heat production with the external and internal muscular activity, the records of pulse rate are defective. Furthermore, no records of the rate of respiration were obtained. A comparison, however, of the pulse rate and the total heat production is of in I crest. The subject of experiment No. 75, S. A. B., recorded the pulse with considerable regularity during the day. The first record was made in the morning, not far from 7h 30m a. m., and was undoubtedly influenced by the muscular activity attendant upon rising, weighing himself and his chair, bed- ding, etc. As an inspection of the data on page 170 shows, his pulse exhibits considerable irregularities from day to day during the period. By omitting this first observation and averaging the remainder during the fasting period, a rough approximation of the average pulse rate per day can be obtained. A comparison (see p. 509) of this pulse rate with the total heat production shows a striking uniformity in fluctuations, and similar comparisons with other experiments show in nearly every instance a parallelism. The pulse rates in the 2-day fasting experiments have been tabulated, together with the respiration rate and total heat production. Although the pneumograph method of obtaining pulse rate and respiration was, on the whole, extremely satisfactory, there were a number of days when the observations were more or less irregular and hence difficulties were experienced in obtaining a fair average pulse and respiration rate for each period. An examination of the detailed statistics for the experiments will show the data from which the averages recorded in table 235 were obtained. Examination of the table shows that in general there is a very marked regularity in the results of pulse rate, respiration rate, and total heat produc- tion when they are compared. But there are a few striking anomalies. Per- haps the most noticeable is in experiment No. 81, where the pulse rate and respiration rate for the second period both increased on the second day, and yet the total heat production decreased nearly 80 calories. In the decade in which experiments have been in progress in this laboratory, evidence has been accumulating to correlate the minor muscular activity with heat production, and the experience thus obtained has been utilized in esti- mating the visible muscular activity in terms of calories recorded in table 234. But it is obvious that careful measurements of pulse rate are of the greatest importance in obtaining an estimate of variations in the so-called internal muscular work. During " rest " experiments this factor evidently varies more than does the visible external muscular activity. MCathcart (Biochemische Zeitschrift (1907), 6, p. 109), finds that blood-pressure as well as pulse-rate decreases during inanition. Energy. 489 Table 235. — Relation of pulse rate, respiration rate, and heat production in 2-day fasting metabolism experiments. Exper- iment num- ber. Subject and date. Period. Average pulse rate per minute. Average respira- tion rate per minute. Total heat produc- tion, calories. 79. 79. 80.. 80.. 81. 81. 82. 82.. 83. 83.. 85.. H. E. S., Oct. 13-14, 1905 H. E. S., Oct. 14-15. C.R.Y., Oct. 27-28. C.R.Y., Oct. 28-29. A.H.M.,Nov.21-22. A.H.M-,Nov.22-23... H.C.K., Nov. 24-25. H.C.K., Nov. 25-26.. H.R.D.,Dec.5-6. H. R. D.,Dec. 6-7, [7 a. m to 1 p. m. I 1 p.m. to 7 p.m. ] 7 p. m. to 1 a. m. 1 la. m to 7 a. m. 7 a. m. to 1 p.m. 1 p. m. to 7 p. m. 7 p.m. to 1 a. m. 1 a. m. to 7 a. m. 7 a. m. to 1 p. m. 1 p. m. to 7 p. m. 7 p. m to 1 a. m. 1 a. m. to 7 a. m. 7 a. m. to 1 p. m. 1 p. m. to 7 p. m. 7 p.m. to 1 a. m. 1 a. m. to 7 a. m. 7 a. m. to 1 p. m. 1 p. m. to 7 p. m. 7 p.m. to 1 a. m. 1 a. m. to 7 a. m. 7 a. m. to 1 p. m. 1 p.m. to 7 p.m. 7 p. m. to 1 a. m. 1 a. m. to 7 a. m. ' 7 a. m. to 1 p. m. 1 p. m. to 7 p. m. 7 p. m. to 1 a. m. 1 a. m. to 7 a. m. 7 a. m. to 1 p. m. 1 p.m. to 7 p. m. 7 p.m. to 1 a. m. 1 a. m. to 7 a. m. 7 a. m. to 1 p.m. 1 p. m. to 7 p. m. 7 p. m. to 1 a. m. 1 a. m. to 7 a. m. 7 a. m. to 1 p.m. 1 p.m. to 7 p.m. 7 p.m. to 1 a. m. 1 a. m. to 7 a. m. N.M.P.,Dec.9-10. { 7 a. m. to 1 p. m. . I 1 p. m. to 7 p. m . 7 p. m. to 1a.m.. la m. to 7 a. m. . 55 61 61 61 73 61 56 57 65 78 79 77 77 46 45 43 42 57 46 49 50 66 62 59 56 68 70 72 68 76 76 74 68 85 78 71 71 62 55 58 62 13 18 18 18 19 19 18 18 16 16 15 16 16 21 17 17 15 13 14 14 18 17 17 16 23 21 20 19 24 23 23 19 20 18 18 17 18 17 18 18 12 12 12 12 526 472 492 460 531 525 504 486 527 449 493 483 592 531 486 489 485 491 395 355 540 412 427 400 642 527 584 467 709 607 634 526 565 386 536 424 585 410 478 431 594 505 534 474 190 Influence of Inanition on Metabolism. Table 235. — Relation of pulse rate, respiration rate, and heat production in 2-day fasting metabolism experiments — Continued. Exper- iment num- ber. Subject and date. Period. Average pulse rate per minute. Average respira- tion rate per minute. Total beat produc- tion, calories. 70 14 638 85.. N.M.P.,Dec. 10-11, 1905 - 1 p. m. to 7 p. m 63 73 14 15 559 592 1 a. m. to 7 a. m 76 15 513 f 7 a. m. to 1 p. m 59 17 480 89.. D.W., Jan. 10-11,1906.. - 1 p.m. to 7 p. m 63 56 19 19 606 571 . 1 a. m. to 7 a. m 50 16 492 f 7 a. m. to 1 p. m 59 16 530 89.. D.W., Jan. 11-12 - 1 p. m. to 7 p. m 7 p. m. to 1 a. m 60 63 18 20 630 626 . 1 a. m. to 7 a. m 50 16 467 The differences in total heat production can in only very small part be accounted for by the actual differences in the work of circulation, assuming the accuracy of Loewy & v. Schrotter's 185 estimates of the relative proportion of the total energy output required for circulation, and consequently variations in pulse rate can be taken only as a general index of the fluctuations in the degree of the internal muscular activity including in all probability muscular tonus. ENERGY OF THE URINE. Fat and carbohydrate katabolized in the ordinary diet are, as a rule, wholly broken down to carbon dioxide and water, and the potential energy they originally contained is completely transformed into heat. A portion of the protein molecule, on the contrary, is not completely oxidized, and is excreted as urea and similar compounds in the urine. The energy of these compounds in the urine must be determined in any series of experiments in which the balance of energy is desired. In order to determine the potential energy of urine, it is necessary to burn the dry matter in a calorimetric bomb and measure the heat actually given off. The technique of the determination of the heat of combustion of urine has received special consideration in an earlier paragraph (see p. 16). The errors involved, in all probability render the determinations somewhat too low rather than too high, although the method used in these investigations has given the highest heat of combustion of any method with which we are familiar. In fasting urine we have reason to suppose not only that portions of the protein katabolized are excreted unoxi- 100 Loc. cit. Energy of Urine. 491 dized but that there may be a more or less marked acidosis with an accompany- ing excretion of acids such as /? oxybutyric acid resulting from the partial oxidation of fat. The total energy of the urine on each day of the fasting experiments is given in table 236. Table 236. — Total energy of urine excreted in metabolism, experiments without food. Exper- iment num- ber. Subject and duration of experiment. First day. Second Third day. day. Fourth day. Fifth day. Sixth day. Seventh day. 59.. 68.. 69.. 71.. 73. . 75.. 77.. 79.. 80.. 81.. 82.. 83.. 85.. 89.. B.F.D., Dec. 18-20,1903 A.L.L., Apr. 27-28, 1904 A.L.L., Dec. 16-19, 1904 S.A.B., Jan. 7-10, 1905 S.A.B., Jan. 28-Feb. 1,1905.. S.A.B., Mar. 4-10, 1905 S.A.B., Apr. 8-11, 1505 A.E.S., Oct. 13-14, 1905 C.R.Y., Oct. 27-28, 1905 A.H.M., Nov. 21-22, 1905 H.C.K., Nov. 24-25, 1905 H.R.D., Dec. 5- 6, 1905 N.M.P., Dec. 9-10,1905 D.W., Jan. 10-11, 1906 Average Cals. 91 92 92 58 86 97 90 72 75 80 88 116 96 93 Cals. 106 99 112 91 98 136 157 116 97 109 122 151 99 114 88 115 Cals. Ill 120 99 96 138 157 120 Cals. 110 86 92 146 170 Cals. 92 147 121 120 Cals. 146 146 Cals 139 139 The smallest amount was found on the first day of experiment No. 71, 58 calories, and the largest amount on the fourth day of experiment No. 77, 170 calories. The total energy is invariably somewhat greater on the second day than on the first. For all the experiments the average is 88 calories on the first day and 115 calories on the second. In the 5 experiments which lasted for 3 days or longer, the energy on the third day is a little greater than that on the second day, while in experiments Nos. 75 and 77, there is a marked increase of energy excreted in the urine on the fourth day. The excretion of energy in experiment No. 75 remains fairly constant after the first day. When it is considered that the energy liberated from the body in the form of heat in some of these experiments was as low as 1500 to 1600 calories, it can readily be seen that the energy of the urine may amount to nearly 10 per cent of the total energy and hence especial care should be taken in experiments dur- ing inanition to secure accurate measurements of this factor. The assumption pointed out on page 395 that non-nitrogenous material of high carbon content is excreted in the urine during fasting receives substan- tiation from the figures for the heat of combustion, for it is seen that the large carbon excretion in the urine of experiments Nos. 75 and 77 is paralleled by a large amount of energy. The difficulties attending the direct determi- nation of the energy of urine have led to the attempt to secure a factor for 492 Influence of Inanition on Metabolism. computation from the amounts of nitrogen and carbon known to be contained in the urine. In these experiments not only was the amount of energy determined but also the nitrogen and carbon and organic matter, so that we have the means for determining the ratio of the heat of combustion to the nitro- gen, carbon, and organic matter. These ratios have been calculated and are given in the following table. The results are expressed in terms of the number of calories of energy which accompany 1 gram of nitrogen, carbon, and organic matter, respectively. Thus, on the first day of experiment No. 77, for every gram of nitrogen in the urine, there was found to be 10.216 calories of energy. For every gram of carbon there were on this day 11.292 calories, and for every gram of organic matter, 3.184 calories. An inspection of these ratios shows that the ratio of energy to nitrogen may vary considerably. The lowest ratio observed is on the third day of experiment No. 59, 7.490, and the highest is that observed on the fourth day of experiment No. 77, 14.847. The average ratio is not far from 9. In the nitrogen metabolism experiments, the heat of combustion of the urine was determined on 6 days. The respective ratios of the heat of com- bustion to nitrogen were 8.8, 11.0, 9.6, 9.3, 8.3, and 9.8, respectively. The ratio of energy to carbon is much more nearly constant, the lowest ratio (9.436) observed being on the first day of experiment No. 68 and the highest on the first day of experiment No. 85 (12.468). Here the differences are very much less than in the case of the energy-nitrogen ratio and it is fair to assume that in experiments of this character for every gram of carbon in the urine there are not far from 11.5 calories of energy. The determination of carbon in the urine is, however, not much less trouble- some than the determination of energy and this ratio is, therefore, not of especial value, although it is unusually constant. Since the carbon is in large measure proportional to the total organic matter, the ratio of heat of combus- tion to organic matter is of interest since the organic matter can be determined with little difficulty. These ratios have been computed and are recorded in table 237. They show that for every gram of organic matter there may be energy corresponding to from 2.264 to 3.857 calories, but in a large majority of instances, the ratio is not far from 3.2. While by no means as constant as the energy-carbon ratio, the energy-organic matter ratio is much more satis- factory than that based upon the determination of nitrogen. In practically all of the experiments here reported, the use of the factor 3.2 as the ratio of energy to organic mattter would have given results which would not have seriously affected the final computations. It is of especial importance, how- ever, to note that in these determinations the total transformations of energy were extremely small and consequently in ordinary metabolism experiments where the energy transformations are larger the total energy in the urine may be computed from the organic matter and the results be well within the limits of error. Energy of Urine. 493 Table 237. — Ratio of heat of combustion to nitrogen, carbon, and organic matter of urine in metabolism experiments without food. Experi- ment num- ber. Subject and duration of experiment. First day. Second day. Third day. Fourth day. Fifth day. Sixth day. Seventh day. 59. 68. 69... 71.. 73. 75. 77. 79. 80. 81.. 82. 83. 85. 89. B.F.D., Dec. 18-20, 1903: Nitrogen Carbon Organic matter... A.L.L., Apr. 27-28, 1904: Nitrogen Carbon Organic matter. . . A.L.L., Dec. 16-19, 1904: Nitrogen Carbon Organic matter. . . S.A.B., Jan. 7-10, 1905: Nitrogen Carbon Organic matter. . . S.A.B., Jan. 28-Feb. 1, 1905 : Nitrogen Carbon Organic matter. . . S.A.B., Mar. 4-10, 1905: Nitrogen Carbon Organic matter. . , 8.A.B., Apr. 8-11, 1905: Nitrogen Carbon Organic matter. . H.E.S., Oct. 13-14, 1905: Nitrogen Carbon Organic matter. . C.R.Y., Oct. 27-28, 1905: Nitrogen Carbon Organic matter. . A.H.M., Nov. 21-22, 1905: Nitrogen Carbon Organic matter. . H.C.K., Nov. 24-25, 1905 : Nitrogen Carbon Organic matter. . H.R.D., Dec. 5-6, 1905: Nitrogen Carbon Organic matter. . N.M.P., Dec. 9-10, 1905: Nitrogen Carbon Organic matter. . D.W., Jan. 10-11, 1906: Nitrogen Carbon Organic matter. . 7 K) 2 7 9. 2. 9. 11. 3. 9. 11. 3. 8. 11. 3. 7 . 11. 2. 10. 11. 3. 8. 11. 3. 9. 10. 3. 8, 10, 3 9 11 3 8, 11 3 12 9. 11, 2, 696 448 312 504 436 675 118 071 128 932 111 247 358 513 014 925 916 977 216 292 184 878 483 216 640 549 301 782 638 ,062 ,382 .414 .205 ,755 .885 .127 .443 .468 .066 309 098 951 7. 10. 2. 7. 9. 2. 7. 11. 9, 11 3 10. 11. 3. 14 11 3 8 11 11, 2, 8, 11, 3, 11 11, 3 8 12 3 7 11 2 512 192 275 598 547 712 854 234 902 243 977 942 187 793 ,030 ,924 .826 .553 .564 .263 .679 084 350 055 749 754 225 352 ,498 960 ,496 328 161 ,160 ,815 ,441 .722 ,044 .154 .884 . 550 .855 .490 .165 .265 7. 11. 2, 7, 11 2 12. 3. 10. 12, 3. 14 11 3 979 572 932 557 458 823 319 8 106 12 078 3 599 421 550 299 14 46011 613 3 481 677 038 007 125 810 855 9 01011 102 3 554 967 744 847 669 740 218 660 078 523 621 830 594 146 857 722 257 644 494 Influence of Inanition on Metabolism. Data were at hand for the computation of the ratio of the heat of combustion of urine to organic matter in the food experiments Nos. 70, 72, 74, and 76 and the results are presented in table 238. Table 238 — Relation of heat of combustion to organic matter ism experiments with food. of urine in metabol- Experiment number. Subject and date. (a) Total organic matter of urine. (b) Heat of combustion of urine. (O Ratio of heat of combustion to organic matter (b-s-a). 70 A.L.L.: Dec. 20-21, 1904 Grams. 35.59 29.30 29.61 29.48 29.47 24.06 20.70 38.88 21.41 23.40 Calories. 103 82 91 84 88 68 61 125 66 69 2.894 2.799 3.073 2.849 2.986 2.826 2.947 3.215 3.083 2.949 72 Dec. 21-22, 1904 Dec. 22-23, 1904 S.A.B.: Jan. 11-12, 1905 74 S.A.B.: Feb. 2- 3,1905 Feb. 3- 4, 1905 76 Feb. 4- 5, 1905 S.A.B.: Mar 11 12 1905. . , Mar. 12-13,1905 ., Mar. 13-14, 1905 So far as experiments Nos. 70, 72, and 74 are concerned the ratios are not unlike those obtained in the fasting experiments preceding each, but there is a marked decrease in the ratio between the seventh day of experiment No. 75 and the 3 days of experiment No. 76. The ratio gradually decreases on the 3 successive days of this experiment, ultimately reaching a value not far from that observed in the 2 earlier food experiments with this subject. No data regarding the ratio of heat of combustion to organic matter were secured in the nitrogen metabolism experiments made with this subject. Although the energy was determined in a few instances, no determinations of the organic matter were made. The factor proposed, then, for comparing the total organic matter with the energy, namely, 3.2, represents with reasonable accuracy the ratio of energy to organic matter, found not only during fasting but also in the food experiments. Since the determination of organic matter is relatively simple, this factor should be of value in computing the energy eliminated in the urine where the actual determination of the heat of combustion can not be made. The only other ratios of the heat of combustion to carbon and organic matter in experiments on fasting man with which we are familiar are those obtained in connection with experiments made in this laboratory and reported else- where.1 The ratios of the heat of combustion to nitrogen, carbon, and organic JM U. S. Dept. Agr., Office of Expt. Sta. Bui. 136. Energy of Body Material. 495 matter of urine determined in a 1-day fasting experiment (No. 36) were not published in that report. They were found to be 8.39, 11.12, and 2.59, respectively. The ratio of heat of combustion to nitrogen has been computed for the other fasting experiments published simultaneously with experiment JSTo. 36, although the data were lacking for computing the ratios of heat of com- bustion to carbon and organic matter. The ratio Cal. : 1ST in experiments Nos. 39, 42, and the 2 days of experiment No. 51 were 8.19, 7.66, 8.19, and 7.08 respectively. Eatios of heat of combustion to nitrogen as wide as those obtained in some of these fasting experiments have rarely been observed. A disturbance of the proteid metabolism accompanied by unusually large Cal. : N ratios during residence in high altitudes was observed and reported by Loewy,167 who was able to isolate a considerable amount of amino bodies from the urine. Indeed, there seemed to be a distinct relation between the quanti- ties of amino bodies and the height at which the subject of the experiments dwelt. While we have but little reason to expect an excessive elimination of amino bodies in the urine of fasting men, the high ratio of heat of combustion to nitrogen may well be explained by the formation of acids such as has been pointed out in the discussion on the carbon and energy content of the urine. ENERGY OF KATABOLIZED BODY MATERIAL. In the course of katabolism during inanition, varying quantities of protein, fat, and carbohydrate are disintegrated. The amounts thus katabolized have been discussed in the several sections. Since these compounds are capable of supplying widely varying quantities of energy to the body, it is of interest to note the proportions of the total energy liberated in the body as a result of the katabolism of the three compounds. Proportion of energy derived from different sources. — The proportions of the energy supplied by the protein, fat, and glycogen have been computed and recorded in table 239. In this discussion the net energy of the protein, i. e., the energy of the protein less that of the unoxidized material of the urine, alone is considered. One possible source of error is present in this method inasmuch as it is here assumed that the unoxidized material of the urine is derived wholly from protein. Under normal conditions this is probably the case, but with the possibilities of an acidosis as a result of partial oxidation and cleavage of fat, it is obviously erroneous to deduct from the energy of the protein katabolized, the energy of the urine in case a portion of the energy of the urine has been derived from fat. The effect of this assumption is to decrease the proportion of the total energy derived from the protein and to increase the proportion actually derived from fat. Unfortunately no data are at hand to show the exact magnitude of the error thus involved. 16IDeutsch. med. Wochenschrift (1905), 48, p. 1918. 496 Influence of Inanition on Metabolism. Table 239. -Proportion of energy derived from different sources in metabolism experiments with and without food. Ex- From Ex- From peri- ment Subject and date. net energy From fat. From glyco- peri- ment Subject and date. net energy From fat. From glyco- num- of pro- gen. num- of pro- gen. ber. tein. ber. tein. Experiments Experiments without food. without food. 59.. B.F.D., 1903: P. ct. P.ct. P.ct. 80.. C.R.T., 1905: P.ct. P.ct. P.ct. Dec. 18-19 14.6 67.8 17.6 Oct. 27-28 9.6 68.5 21.9 Dec. 19-20 17.6 70.6 11.8 11.3 85.3 3.4 Dec. 20-21 18.1 81.1 0.8 81.. A.H.M., 1905: 68.. A.L.L., 1904: Nov. 21-22 13.1 80.1 6.8 Apr. 27-28 14.9 63.5 21.6 Nov. 22-23 12.8 87.2 .... Apr. 28-29 15.8 70.3 13.9 82.. H.C.K., 1905: 69.. A.L.L., 1904: Nov. 24-25 10.2 59.1 30.7 Dec. 16-17 12.7 65.3 22.0 Nov. 25-26 14.6 77.9 7.5 Dec. 17-18 17.1 76.8 6.1 Dec. 18-19 18.9 74.5 6.6 83.. H.R.D., 1905: Dec. 19-20 16.4 80.4 3.2 Dec. 5-6 Dec. 6-7 17.0 16.6 76.0 74.0 7.0 9.4 71.. S.A.B., 1905: Jan. 7- 8 7.0 55.2 37.8 85.. N.M.P., 1905: Jan. 8- 9. . . . 15.2 78.1 6.7 Dec. 9-10 13.7 57.4 28.9 Jan. 9-10 19.2 75.7 5.1 Dec. 10-11 12.6 70.5 16.9 Jan. 10-11 16.8 76.8 6.4 89.. D.W., 1906: 73.. S.A.B., 1905: Jan. 10-11 11.2 57.2 31.6 Jan. 28-29 14.2 55.1 30.7 Jan. 11-12 16.4 76.3 7.3 Jan. 29-30 16.8 79.0 4.2 Jan. 30-31 16.6 81.7 1.7 Experiments Jan. 31-Feb. 1. 15.5 79.1 5.4 with food. Feb. 1- 2 12.5 87.5 70.. A.L.L., 1904: Dec. 20-21 16.2 72.4 11.4 75. . S.A.B., 1905: Dec. 21-22 11.2 82.1 6.7 Mar. 4- 5 . . . . 17.7 67.1 15.2 Dec. 22-23 10.2 72.8 17.0 Mar. 5- 6 16.0 78.6 5.4 Mar. 6- 7 17.0 81.7 1.3 72.. S.A.B., 1905: Mar. 7- 8 14.3 79.6 6.1 Jan. 11-12 16.0 81.5 2.5 Mar. 8- 9 13.5 84.4 2.1 Mar. 9-10 14.1 80.0 5.9 74.. S.A.B., 1905: Mar. 10-11 13.2 81.8 5.0 Feb. 2-3 Feb. 3-4 16.1 12.8 77.1 64.9 6.8 22.3 77.. S.A.B., 1905: Apr. 8- 9 11.1 68.3 20.6 Feb. 4-5 10.1 71.0 18.9 Apr. 9-10 10.9 85.9 3.2 76.. S.A.B., 1905: Apr. 10-11 12.1 74.0 13.9 Mar. 11-12 12.4 74.3 13.3 Apr. 11-12 12.3 80.8 6.9 Mar. 12-13 Mar. 13-14 10.4 11.2 57.2 36.5 32.4 52.3 79.. H.E.S., 1905: Oct. 13-14 Oct. 14-15 10.4 18.1 64.5 73.8 25.1 8.1 For purposes of comparison, the proportions of energy from the protein, fat, and glycogen in the experiments with food are included in the table. From 10 to 15 per cent of the total energy produced is derived from protein. The smallest percentage is on the first day of experiment No. 71, where but 7 Katabolism and Heat Production. 497 per cent of the total energy was derived from protein. The largest proportion is on the third day of experiment No. 71, namely, 19.2 per cent. In the food experiments the proportions derived from protein are not materially different on the whole from those observed during fasting. An examination of the proportions of energy derived from fat shows con- siderable variations, which are especially noticeable on the first days of the longer fasting experiments. The apportionment of the carbon other than that of protein between the fat and glycogen would lead to the supposition that as the quantity of katabolized glycogen decreased, the amount of fat would be increased, and this is actually what is found in comparing the amounts of these two materials katabolized. Similar fluctuations in the energy derived from the two compounds are also observed. No regularity appears in the proportion of the energy resulting from fat. It is generally greater than 75 per cent, but in fasting experiments may be as small as 55 per cent. In one instance, namely, on the last day of food experiment No. 76, it fell to 36.5 per cent. The highest proportion was found on the last day of experiment No. 73. Since a much smaller proportion of the total energy results from glycogen rather than from fat katabolism, the fluctuations in the proportions of the energy supplied by the glycogen are very considerable, and it is difficult to ob- tain any general average for the amount of energy furnished by this compound. In two instances there was apparently an absorption of energy due to the formation and storage of glycogen,108 namely, on the last day of experiment No. 73 and the second day of experiment No. 81. This apparent storage or absorption of energy in the form of glycogen could only be derived from body fat or protein, and hence the absorption of energy is apparent and not real. The largest proportion of energy supplied by glycogen on any one day in the fasting experiments was 37.8 per cent. On the last day of food experiment No. 76, a maximum of 52.3 per cent was reached. COMPARISON OF PRODUCTS OF KATABOLISM AND HEAT PRODUCTION. It has commonly been assumed that heat production and katabolism are interdependent and the older methods of computing the heat production were based on this assumption. Therefore, experiments in which all the grosser factors of katabolism have been studied present unusual opportunity for an examination of the regularity of the course of the heat production compared to that of the general katabolism. Such a comparison may be made in several ways. The relations between the grosser factors, oxygen consumption, carbon dioxide and water elimination, and heat production are first studied, and later the numerous other products of katabolism, especially those appearing in the urine, are compared. W8 For errors affecting the storage of glycogen see discussion of ratios of carbon dioxide and oxygen to heat, p. 514. 32 498 Influence of Inanition on Metabolism. Table 240. — Proportion of oxygen consumed and of carbon dioxide and water elimi- nated and heat produced during different periods of the day in metabolism experiments without food. Period. 7 a.m. to 10 a.m. 10 a.m. 1p.m. Total, 6 hours . . 1 p.m. to 4 p.m. 4 pm. 7 p.m. Total, 6 hours . . Total, 12 hours. 7 p.m. to 10 p.m. 10 p.m. 1 a.m. Total, 6 hours. . 1 a.m. to 4 a.m. 4 a.m. 7 a.m. Total, 6 hours. . Total, 12 hours . 7 a.m. to 9 a.m. 9 a.m. 11 a.m. 11 a.m. 1 p.m. Total, 6 hours, to 1 p.m 3 p.m 5 p.m 3 p.m 5 p.m 7 p.m Total, 6 hours. , Total, 12 hours, 7 p.m. to 9 p.m. 9 p.m. 11 p.m. 11 p.m. 1 a.m, Total, 6 hours. , to 1 a.m. 3 a.m. 5 a.m. 3 a.m. 5 a.m . 7 a.m. Total, 6 hours. . Total, 12 hours, Exp. No. 59, subject B.F.D., 3 days. Oxygen. Carbon dioxide. Water. Heat. Per cent. 15.54 12.33 27.87 Per cent. 14.78 13.05 27.83 Per cent. 13.22 12.95 26.17 Per cent. 15.04 13.30 28.34 13.04 13.47 26.51 54.38 13.00 13.70 26.70 54.53 12.30 12.82 25.12 51.29 12.72 13.74 26.46 54.80 13.63 10.97 24.60 13.07 11.13 24.20 12.35 11.93 24.28 13.15 10.51 23.66 10.76 10.26 21.02 45.62 10.33 10.94 21.27 45.47 12.13 12.30 24.43 48.71 10.03 11.51 21.54 45.20 Exp. No. 69, subject A.L.L., 4 days. 10.82 7.87 8.24 26.93 11.04 8.08 8.22 27.34 9.85 7.98 8.11 25.92 10.16 7.73 7.82 25.71 9.30 8.63 8.63 26.56 53.49 9.06 8.79 8.98 26.83 54.17 7.94 8.08 8.07 24.09 50.01 9.85 8.98 9.45 28.28 53 . 99 9.17 8.07 7.54 24.78 8.74 8.33 7.65 24.72 7.99 7.59 8.43 24.01 8.51 8.25 8.12 24.88 7.32 7.25 7.16 21.73 46.51 6.86 7.13 7.12 21.11 45.83 8.78 9.21 7.99 25.98 49.99 6.98 7.01 7.14 21.13 46.01 Exp. No. 68, subject A.L.L., 2 days. Oxygen. Carbon dioxide. Per cent. 15.63 12.50 28.13 Per cent. 15.21 13.07 28.28 Water. Heat. Per cent. 13.68 12.46 26.14 Per cent. 13.77 13.36 27.13 12.94 13.72 26.66 54.79 13.03 13.55 26.58 54.86 12.50 12.74 25.24 51.38 13.14 13.58 26.72 53.85 12.98 11.44 24.42 12.95 11.78 24.73 12.08 12.35 24.43 12.97 12.15 25.12 10.50 10.29 10.03 10.38 12.22 11.97 20.79 45.21 20.41 45.14 24.19 48.62 10.43 10.60 21.03 46.15 Exp. No. 71, subject S.A.B., 4 days. 12.56 10.37 8.38 31.31 12.18 10.85 9.07 10.73 9.73 8.49 32.10 28.95 11.21 10.72 9.51 31.44 9.59 8.03 8.63 26.25 57.56 9.01 8.21 8.50 25.72 57.82 8.62 8.50 8.93 26.05 55.00 8.81 7.66 9.77 26.24 57.68 9.25 7.44 6.34 23.03 8.85 7.41 6.64 8.54 7.54 7.62 22.90 23.70 8.08 5.60 8.28 21.96 6.71 6.65 6.05 19.41 42.44 6.40 6.41 6.47 19.28 42.18 7.52 6.49 7.29 21.30 45.00 6.36 7.09 6.91 20.36 42.32 Katabolism and Heat Production. 499 Table 240. — Proportion of oxygen consumed and of carbon dioxide and water elimi- nated and heat produced during different periods of the day in metabolism experiments without food — Continued. Period. 7 a.m. to 9 a.m 9 a.m. 11 a.m 11 a.m. 1 p.m Total, 6 hours . 1 p.m. 3 p.m. 5 p.m. to 3 p.m 5 p.m 7 p.m Total, 6 hours. Total, 12 hours 7 p.m. 9 p.m. 11 p.m. to 9 p.m 11 p.m 1 a.m Total, 6 hours . to 1 a.m 3 a.m 5 a.m 3 a.m, 5 a.m, 7 a.m. Total, 6 hours. . Total, 12 hours. 7 a.m. to 9 a.m. 9 a.m. 11 a.m. 11 a.m. 1 p.m. Total, 6 hours . . 1 p.m. 3 p.m. 5 p.m. to 3 p.m. 5 p.m. 7 p.m. Total, 6 hours. . Total, 12 hours. 7 p.m. to 9 p.m. 9 p.m. 11p.m. 11 p.m. 1 a.m. Total, 6 hours . . 1 a.m. to 3 a.m. 3 a.m. 5 a.m. 5 a.m. 7 a.m. Total, 6 hours . . Total, 12 hours. Exp. No. 73, subject S.A.B., 5 days. Oxygen. Per cent. 11.51 9.70 9.06 Carbon dioxide. Per cent. 11.24 9.88 9.21 Water. Per cent. 9.70 9.44 8.65 30.27 30.33 27.79 29.48 Heat. Per cent. 10.14 10.05 9.29 8.57 8.77 9.04 26.38 56.65 8.91 6.84 9.21 8.38 8.09 8.51 26.76 57.09 24.98 52.77 8.97 8.48 9.62 27.07 56.55 8.37 8.18 5.99 8.40 8.01 6.74 8.49 8.09 7.82 22.54 23.15 24.40 22.38 8.60 7.85 5.93 7.03 6.99 6.79 20.81 43.35 6.32 6.89 6.55 7.70 7.63 7.50 19.76 42.91 22.83 47.23 6.11 7.66 7.30 21.07 43.45 Exp. No. 77, subject S.A.B., 4 days. 11.17 7.92 7.58 26.67 10.86 8.29 7.79 26.94 10.60 8.14 8.06 9.78 8.66 7.96 26.80 26.40 8.09 8.91 8.58 25.58 52.25 8.70 8.56 8.12 25.38 52.32 8.36 7.76 8.31 24.43 51.23 8.56 9.47 8.53 26.56 52.96 8.18 9.85 6.14 24.17 8.54 9.37 7.00 24.91 8.25 8.35 7.93 7.78 8.05 6.88 24.53 22.71 7.68 7.81 8.09 23.58 47.75 7.22 7.78 7.77 22.77 47.86 8.07 8.02 ' 8.15 24.24 48.77 7.16 8.68 8.49 24.33 47.04 Exp. No. 75, subject 8.A.B., 7 days. Oxygen. Per cent. 10.90 8.83 8.74 28.47 Carbon dioxide. Water. Per cent. 10.42 9.44 8.98 Per cent. 10.29 8.69 8.26 28.84 I 27.24 Heat. Per cent. 10.60 10.01 9.00 29.61 8.72 9.08 9.00 26.80 55.27 8.90 8.45 9.07 26.42 55.26 8.40 7.82 7.93 24.15 51.39 8.94 8.50 9.32 26.76 56.37 8.08 8.90 6.10 8.42 8.52 6.79 7.91 8.39 7.99 23.08 23.73 24.29 8.10 7.83 6.01 21.94 6.50 7.71 7.44 6.51 7.06 7.44 21.65 44.73 21.01 44.74 8.07 8.20 8.05 24.32 48.61 6.45 7.79 7.44 21.69 43.63 Experiments Nos. 79-83, 86, and 90. 11.32 7.91 8.69 27.92 10.93 8.81 8.82 28.56 9.29 8.33 8.43 26.05 10.69 8.26 8.63 27.58 8.24 7.95 8.51 24.70 52.62 8.36 7.95 8.12 24.43 52.99 8.13 8.00 8.47 24.60 50.65 8.46 7.58 8.55 24.59 52.17 9.28 8.44 7.47 8.97 8.29 7.80 8.56 8.16 8.18 25.19 25.06 24.90 25.41 9.06 8.34 8.10 7.21 7.35 7.63 22.19 47.38 7.03 7.28 7.64 21.95 47.01 8.13 7.94 8.38 7.21 7.27 7.94 24.45 49.35 22.42 47.83 -.on Influence of Inanition on Metabolism. Proportion of oxygen consumed, carbon dioxide and water eliminated, and heat produced during different periods. — The variations in the amounts of the four factors — oxygen, carbon dioxide, water, and heat — which occur during the different periods of the day are recorded in terms of per cents in table 240. A certain parallelism appears in the proportions of these four factors; thus, the largest katabolic activity occurs, as a rule, during the first period of the day when more oxygen is consumed and carbon dioxide and water given off and heat produced than during any other period of the day. A close examina- tion of the figures shows that in the 2-hour periods during the night from 1 to 7 a. m., the results from period to period are on the whole more nearly constant than during those of any other 6 hours. As has been shown in the foregoing discussion, the influence of external muscular activity is compara- tively unimportant in its effect on the total heat production of a fasting man at rest. The results further show that although fluctuations in the pulse rate (assumed to be due to changes in the work of circulation) do not account for the marked differences in the accompanying heat production, nevertheless the pulse rate has been found to be a remarkably good index of the rise or fall in the amount of heat produced. The external muscular activity as well as the pulse rate is usually lowest during the period from 1 to 7 a. m. During periods of sleep major muscular movements were not impossible, but it is probable that in general the subjects were in about the same degree of muscular activity in all the experiments. It is quite probable that extraneous muscular exertion can be neglected, although undoubtedly on certain nights, when the subjects reported a poor night's sleep or the physical assistant recorded unusual muscular activity, the external muscular activity did have an influence on the total katabolism. It is of especial interest to study the four important factors during this night period. For purposes of comparison, the amount of oxygen consumed, carbon dioxide and water given off, and heat produced per minute, are computed from 1 to 7 a. m. in all the experiments with and without food. These results are given in table 241. The striking feature of these comparisons is the fact that in the longer experiments, except No. 77, which was abnormal in this and other respects, during the night period when the muscular activity is at the minimum, there is a progressive diminution in the total katabolism as indicated by the con- sumption of oxygen, the elimination of carbon dioxide and water-vapor, and the production of heat. This observation is also true when the computations are made on the basis of per kilo of body-weight or per square meter of body surface. Katabolism and Heat Production. 501 Table 241. — Volume of oxygen consumed and amounts of carbon dioxide and water eliminated and neat produced per minute from 1 a. m. to 7 a. m. in metabolism experiments with and without food. Oxygen Carbon dioxide Water Heat Experi- consumed. eliminated. eliminated. production. ment number. Subject and date. Per kilo- gram of body- weight. Per square meter of body surface. Per kilo-! Per ~™\Cr„f square W°f meter bo-dT; of body ""*"• j surface5. Per kilo- gram of body- weight. Per square meter of body surface. Per kilo- P!' gbXofi = body- ! of body weight., surfac* 59... B. F.D., 1903: c.c. c.c. c.c. c.c. Gram. Gram. Calorie. Calorie. Dec. 18-19 .... 3.721 122.8 2.894 95.50 .0096 .3161 .0176 .5814 Dec. 19-20 4.071 133.7 3.067 100.71 .0098 .3211 .0190 .6242 Dec. 20-21 3.966 129.8 3.031 99.23 .0101 .3315 .0202 .6616 68... A.L.L., 1904: Apr. 27-28. . . . 3.523 119.1 2.733 92.40 .0066 .2223 .0167 .5640 Apr. 28-29.... 3.738 125.6 2.819 94.69 .0076 .2553 .0192 .6453 69... A.L.L., 1904: Dec. 16-17. . . . 3.616 123.0 2.737 93.12 .0073 .2486 .0173 .5897 Dec. 17-18. . . . 3.691 125.1 2.630 89.15 .0098 .3308 .0174 .5905 Dec. 18-19.... 3.499 118.2 2.598 87.75 .0076 .2561 .0147 .4957 Dec. 19-20. . . . 3.507 118.1 2.555 86.06 .0069 .2307 .0163 .5489 70... A. L.L., 1904: Dec. 20-21.... 3.328 111.9 2.629 88.38 .0088 .2974 .0165 .5545 Dec. 21-22. . . . 3.242 108.9 2.654 89.15 .0099 .3342 .0177 .5957 Dec. 22-23.... 4.264 143.2 3.412 114.58 .0111 .3719 .0221 .7423 71... S.A.B., 1905: Jan. 7- 8 3.952 124.0 2.946 92.42 .0065 .2027 .0172 .5395 Jan. 8- 9 3.658 114.3 2.956 92.34 .0071 .2211 .0190 .5949 Jan. 9-10 3.434 106.9 2.608 81.20 .0065 .2014 .0178 .5556 Jan. 10-11.... 3.433 106.5 2.561 79.40 .0061 .1880 .0175 .5428 72... S.A.B., 1905: Jan. 11-12. ... 3.324 102.8 2.593 80.17 .0064 .1967 .0171 .5288 73... S.A.B., 1905: Jan. 28-29 3.837 121.0 2.944 92.87 .0074 .2344 .0189 .5961 Jan. 29-30 3.720 116.7 2.734 85.75 .0068 .2144 .0181 .5662 Jan. 30-31 3.657 114.2 2.520 78.68 .0066 .2047 .0171 .5351 Jan. 31-Feb. 1. 3.642 113.2 2.551 79.30 .0065 .2022 .0175 .5454 Feb. 1- 2. . . . 3.727 115.3 2.540 78.59 .0065 .2008 .0169 .5215 74... S.A.B., 1905: Feb. 2- 3. . . . 3.330 102.9 2.562 79.19 .0065 .2007 .0162 .5018 Feb. 3- 4. . . . 3.462 107.2 2.670 82.68 .0063 .1943 .0156 .4826 Feb. 4- 5. . . . 3.703 114.9 2.691 83.47 .0061 .1881 .0160 .4952 75... S.A.B., 1905: Mar. 4- 5 3.779 119.9 2.931 92.94 .0074 .2352 .0187 .5919 Mar. 5- 6.... 4.267 135.0 2.866 90.71 .0073 .2295 .0179 .5660 Mar. 6- 7 3.822 120.4 2.726 85.88 .0077 .2417 .0183 .5766 Mar. 7- 8. . . . 3.460 108.5 2.606 81.73 .0068 .2134 .0184 .5758 Mar. 8- 9.... 3.688 115.3 2.532 79.14 .0069 .2157 .0174 .5435 Mar. 9-10.... 3.342 104.2 2.552 79.57 .0065 .2041 .0164 .5109 Mar. 10-11.... 3.461 107.6 2.548 79.22 .0066 .2062 .0167 .5199 502 Influence of Inanition on Metabolism. Table 241. — Volume of oxygen consumed and amounts of carbon dioxide and water eliminated and heat produced per minute from 1 a. m. to 7 a. m. in metabolism experiments with and without food — Continued. Experi- ment number. Subject and date. Oxygen consumed. Carbon dioxide eliminated. Water eliminated. Heat production. Per kilo- gram of body- weight. Per square meter of body surface. Per kilo- gram of body- weight. Per square meter of body surface. Per kilo- gram of body- weight. Per square meter of body surface. Per kilo- gram of body- weight. Per square meter of body surface. 76... S.A.B., 1905: Mar. 11-12.... Mar. 12-13.... Mar. 13-14. . . . c.c. 3.449 3.243 3.236 c.c. C.C. ' 107.0 2.685 100.6 2.691 100.7 2.943 c.c. 83.30 83.50 91.56 Gram. .0070 .0067 .0069 Gram. .2186 .2085 .2144 Calorie. .0173 .0163 .0168 Calorie. .5372 .5065 .5211 77... S.A.B., 1905: Apr. 8- 9. . . . Apr. 9-10.... Apr. 10-11.... Apr. 11-12.... 3.975 4.091 4.302 4.511 127.3 130.3 135.9 141.6 3.125 2.994 3.064 3.131 100.06 95.35 96.81 98.28 .0073 .0069 .0071 .0076 .2330 .2202 .2241 .2379 .0194 .0206 .0222 .0219 .6216 .6566 .7008 .6862 79... H.E.S., 1905: Oct. 13-14. . . . Oct. 14-15. . . . 4.617 4.736 144.0 146.6 3.562 3.695 111.07 114.40 .0086 .0084 .2683 .2611 .0226 .0244 .7045 .7547 80... C.R.Y., 1905: Oct. 27-28. ... Oct. 28-29 4.056 4.404 134.8 145.0 3.051 3.328 101.37 109.60 .0117 .0110 .3874 .3633 .0196 .2004 .6517 .6722 81... A.H.M., 1905: Nov. 21-22. . . . Nov. 22-23. . . . 3.575 3.753 114.6 119.6 2.656 2.695 S5.17 85.88 .0063 .0077 .2026 .2446 .0160 .0184 .5141 .5868 82... H.C.K., 1905: Nov. 24-25. . . . Nov. 25-26. . . . 3.552 4.299 119.5 144.0 2.923 3.273 98.40 109.61 .0071 .0081 .2377 .2715 .0182 .0208 .6133 .6977 83... H. R. D., 1905: Dec. 5- 6. . . . Dec. 6- 7.... 4.345 4.628 134.5 142.7 3.327 3.295 102.97 101.60 .0086 .0078 .2653 .2394 .0213 .0219 .6592 .6764 85... N.M.P., 1905: Dec. 9-10. . . . Dec. 10-11.... 4.001 4.410 131.9 144.5 3.083 3.420 101.63 112.08 .0076 .0090 .2489 .2964 .0197 .0217 .6498 .7118 89... D.W., 1906: Jan. 10-11.... Jan. 11-12. . . . 3.690 3.430 128.2 118.5 2.871 2.701 99.76 93.36 .0071 .0008 .2460 .2345 .0175 .0168 .6064 .5821 The minimum oxygen consumption per kilo of body-weight per minute occurs on the third day of the food experiment ISTo. 76. The elimination of carbon dioxide is lowest on the third day of the fasting experiment No. 73, while the heat production is lowest on the third day of fasting experiment No. 69. The high heat production on the last night of experiment No. 70 is un- questionably associated with a febrile temperature observed on this day In Katabolism and Heat Production. 503 what manner this high heat production is associated with fever,100 the experi- ments do not show, since the pulse rate and respiration rate are not known. For the food experiments with S. A. B. there was no increase in the heat pro- duction in experiment No. 72 over the last day of experiment No. 71. Simi- larly in experiment No. 74, the heat production during the night was even less than during the fasting period of experiment No. 73. Experiment No. 76 showed, on the average, a slight increase over the heat production of the last days of experiment No. 75. It is reasonable to suppose, therefore, that any variation in the heat production during the ingestion of food is not prolonged into the night period from 1 to 7 a. m. Unpublished experiments made in this laboratory indicate that the extra heat production following the ingestion of food is in large part dissipated shortly after the food is taken, the normal resting metabolism without food being reached after a few hours. With the subject S. A. B. it is apparent that the minimum heat production during inanition is not far from 0.0167 calorie per kilogram of body-weight, or 0.52 calorie per square meter of body surface per minute. Relation between relative humidity, water of oxidation of organic hydrogen, and total heat production. — The water of oxidation of organic hydrogen includes all the organic hydrogen of the protein, fat, and glycogen katabolized. It is, therefore, roughly speaking, a measure of the total katabolism during the day, and since the total katabolism determines the heat production, there should be more or less constant ratios between these two factors. In table 242 are recorded the relative humidity, the water of oxidation of organic hydrogen, and the total heat production. The relative humidity and its influence on the water of vaporization has already been discussed (see p. 428). The conditions determining the relative humidity inside this type of respiration chamber present certain features of unusual interest. The ventilating air-current enters the respiration chamber absolutely dry. The body of the subject gives off water continuously to this air, and, depending upon the rate of ventilation and muscular activity and consequent water output of the subject, the relative humidity may vary. In all the experiments the rate of ventilation was practically constant (see p. 429), but the influence of internal and external muscular activity (measured by the total heat production) on the variations in relative humidity is worthy of note. In general, the relative humidity decreases with a diminution in the total heat production and increases with a rise in the heat production, although it may be noted that in experiments with the same subject a marked varia- tion in the relative humidity of the air inside the chamber may occur on different days of the different experiments when the total heat production 109 The special question of the influence of heat production during fever will be discussed in a later report. 504 Influence of Inanition on Metabolism. does not vary. Thus, on the second day of experiment No. 71 with S. A. B.. lli.' relative humidity was 36.6 per cent and the total heat production was 1844 calories while on the third day of experiment No. 77, the total heat pro- duction was 1840 calories and the relative humidity only 30.2 per cent. Such a lack of agreement is even more noticeable in the experiments with different sub- jects. Thus, on the second day of experiment No. 80, the relative humidity was 54 per cent and the total heat production 2099 calories, while on the first day of experiment No. 85, the total heat production was 2109 calories and the relative humidity 35.6 per cent. In spite of these gross fluctuations there is a tendency, however, for the relative humidity to vary with the total heat production. The fluctuations in the amounts of water resulting from the oxidation of organic hydrogen are remarkably comparable with those of the total heat production, especially after the first day of the fast. The relation between these two products bears out the assertion made previously, that the metabolism on the first day of fasting can not be considered as true fasting metabolism. From the average of these experiments with and without food, it is seen that each gram of water of oxidation of organic hydrogen is accompanied by the production of 9.1 calories and every gram of organic hydrogen oxidized is accompanied by a heat production amounting to 82.4 calories. OXYGEN AND CARBON DIOXIDE THERMAL QUOTIENTS. The number of grams of carbon dioxide produced and oxygen absorbed per 100 calories of heat, the so-called carbon dioxide and oxygen thermal quotients, are recorded for experiments with and without food in table 243. The oxygen thermal quotients show that for every 100 calories of heat there were about 30 grams of oxygen absorbed. The accuracy of the determi- nations of oxygen with this apparatus have been found by check tests, to be within 1 per cent. It is evident that this error allows a fluctuation of from 29.8 to 30.2 in the thermal quotient. In the majority of cases it is believed that the oxygen was determined to within 1 per cent and therefore the values given are of significance. It is, nevertheless, a fact that the determination of the carbon dioxide is much less difficult than that of oxygen, and consequently for any comparison between the heat and oxygen or carbon dioxide the latter factor must be considered the more accurate. The lowest oxygen thermal quotient recorded is 28.84 on the third day of experiment No. 77, and the highest is 31.37 on the last day of experiment No. 73. Katabolism and Heat Production. 505 Table 242. — Daily relative humidity, and daily heat production in metabolism experiments with and without food. Water Water Ex- peri- ment Subject and date. Rel- ative hu- of oxida- tion of Total heat pro- Ex- peri- ment Subject and date. Rel- ative hu- of oxida- tion of or- Total heat pro- num- mid- or- duc- num- mid- duc- ber. ity. ganic hydro- gen. tion. ber. ity. ganic hydro- gen. tion. Experiments Experiments without food. without food. 59.. B.F.D., 1903 : P. ct. Oms. Gals. 80.. C.R.Y., 1905: P. ct. Oms. Cats. Dec. 18-19 Dec. 19-20 46.9 44.4 229 223 2080 2107 Oct. 27-28 51.8 54.0 217 224 1954 2099 Oct. 28-29 Dec. 20-21 43.5 222 2102 81.. A.H.M., 1905: 68.. A.L.L., 1904 : Nov. 2 1-22 30.6 185 1729 Apr. 27-28 49.1 241 2167 178 1781 Apr. 28-29 48.8 237 2217 82.. H.C.K., 1905: 69.. A.L.L., 1904: 38.8 256 2222 Dec. 16-17 45.0 221 1951 Nov. 25-26 41.2 261 2477 Dec. 17-18 51.5 231 2163 Dec. 18-19 48.0 221 2035 83. . H.R.D., 1905: Dec. 19-20 43.7 213 1958 Dec. 5-6 33.5 209 1914 Dec. 6-7 32.2 197 1907 71.. S.A.B., 1905: Jan. 7- 8 45.0 235 1970 85.. N.M.P., 1905: Jan. 8- 9 36.6 200 1844 Dec. 9-10 35.6 240 2109 Jan. 9-10 33.5 191 1746 Dec. 10-11 40.1 252 2305 Jan. 10-11 30.8 176 1606 89.. D.W., 1906: 73.. S.A.B., 1905 : Jan. 10-11 38.8 248 2150 Jan. 28-29... 36.3 210 1866 Jan. 11-12 38.8 244 2254 Jan. 29-30 31.4 196 1791 Jan. 30-31 30.9 188 1739 Experiments Jan. 31-Feb. 1. 29.0 180 1663 with food. Feb. 1- 2 27.9 170 1548 70.. A.L.L., 1904: Dec. 20-21 45.0 226 2104 75.. S.A.B., 1905: Dec. 21-22 48.0 244 2223 Mar. 4- 5. . . . 32.3 195 1765 Dec. 22-23 50.5 275 2457 Mar. 5- 6 29.0 189 ]768 Mar. 6- 7 29.4 187 1797 72.. S.A.B., 1905: Mar. 7- 8 .. 27.2 185 1775 Jan. 11-12 32.0 184 1676 Mar. 8- 9 26.4 173 1649 Mar. 9-10 24.4 166 1553 74.. S.A.B., 1905 : Mar. 10-11 25.1 164 1568 Feb. 2-3 Feb. 3-4 30.0 28.7 184 184 1691 1585 77.. S.A.B., 1905: Apr. 8- 9 30.5 208 1874 Feb. 4-5 29.6 187 1607 Apr. 9-10 30.7 202 1880 76.. S.A.B., 1905: Apr. 10-11 30.2 192 1840 Mar. 11-12 27.2 191 1767 Apr. 11-12 29.7 188 1807 Mar. 12-13 26.5 194 1728 79.. H.E.S., 1905: Oct. 13-14 57.0 220 1951 Mar. 13-14 Average of all experiments. 26.6 206 1753 209 1911 Oct. 14-15 61.2 215 2047 506 Influence of Inanition on Metabolism. Table 243. — Oxygen and carbon dioxide thermal quotients in metabolism experiments with and without food. Exper- iment num- ber. Subject and duration of experiment. First day. Second day. Third Fourth Fifth day. j day. day. Sixth day. Seventh day. 59.. 68.. 69.. 70.. 71.., 72... 73.. 74.. 75.. 76.., 77.., 79.., 80... 81.., 82.. 83.. 85.. 89.. B.F.D., Dec. 18 to 20, 1903: Oxygen Carbon dioxide. . A.L.L., Apr. 27 to 28, 1904: Oxygen Carbon dioxide. . A.L.L., Dec. 16 to 19, 1904: Oxygen Carbon dioxide. . , A.L.L., Dec. 20 to 22, 1904: Oxygen Carbon dioxide. . S.A.B., Jan. 7 to 10, 1905: Oxygen Carbon dioxide. . , S.A.B., Jan. 11, 1905: Oxygen , Carbon dioxide . . S.A.B., Jan. 28 to Feb.1,1905 Oxygen Carbon dioxide. . , S.A.B., Feb. 2 to 4, 1905: Oxygen Carbon dioxide. . , S.A.B., Mar. 4 to 10, 1905 Oxygen Carbon dioxide S.A.B., Mar. 11 to 13, 1905: Oxygen Carbon dioxide. . , S.A.B., Apr. 8 to 11, 1905: Oxygen Carbon dioxide. . , H.E.S., Oct. 13 to 14, 1905: Oxygen Carbon dioxide. . , C.R.T., Oct. 27 to 28, 1905: Oxygen Carbon dioxide . . A.H.M., Nov. 21 to 22, 1905 : Oxygen Carbon dioxide. . H.C.K., Nov. 24 to 25, 1905: Oxygen , Carbon dioxide. . , H.R.D., Dec. 5 to 6, 1905: Oxygen Carbon dioxide. . . N.M.P., Dec. 9 to 10, 1905: Oxygen Carbon dioxide. . , D.W., Jan. 10 to 11, 1906: Oxygen Carbon dioxide. . . 30.25 32. '23 29.53 32.04 29.87 30.74 31.28 30.94 28.98 30.63 29.94 29.86 30.43 32.3S 30.81 31.48 30.19 30.83 29.83 31.64 30.05 |30.81 30.92 31.72 29.59 31.03 29.90 33.96 30.83 31.29 29.17 30.59 30.65 32.64 31.27 31.15 30.28 31.25 30.86 30.81 33.42 32.82 30.24 30.22 29,81 32.29 31.14 30.33 29.84 31.20 29.66 31.98 29.51 32.39 29.49 32.11 29 . 90 30.93 29.45 28.94 28.92 32.43 34.70 30.41 28.84 30.69 30.25 29.57 31.03 29.94 30.50 29.60 29.85 33.35 30.58 31.70 29.76 33.03 30.01 33.60 29.62 30.97 29.07 30.37 29.31 31.21 30.22 31.30 30 31 30 31 30 30 29 30 29 30 70 29 69 65 23 98 26 09 41 14 31 31 29 30 37 15 78 10 30 30 01 7 4 29 30 75 33 Katabolism and Heat Production. 50? The absorption of oxygen results in the combustion of carbon in the case of glycogen assuming that the oxygen of the carbohydrate molecule combines with the hydrogen/ and the combustion of carbon and organic hydrogen in the case of fat and protein, and hence it would be expected that the oxygen thermal quotient would vary with the proportions of protein, fat, and glycogen con- sumed. Since the widest fluctuations in the relative proportions of these com- pounds occur on the first day of the experiments, the oxygen thermal quotients would be expected to be less regular on the first day than on other days of the fasts. It will be noticed that the highest and lowest quotients were observed on the third and fifth days of fasting. This fact is of importance in the subsequent comparison of the heat production with the estimated energy of the material oxidized in the body (see p. 514). When the relative proportions of protein, fat, and carbohydrate become more nearly established as on the second and subsequent days, the quotient should remain practically constant, although the fact must not be lost sight of that there are relatively wide variations in the proportions of glycogen and fat katabolized even on some of the later fasting days. Making due allowance, however, for such deviations, it is clear that marked variations in the oxygen thermal quotients in the later days of fasting are due to one of two things, defective oxygen determinations or defective heat measurements. A further discussion of the relation of oxygen and heat production is given on page 515. The carbon dioxide thermal quotient, like that of the oxygen is determined by the nature of the material burned. The larger the amount of glycogen, the greater the amount of carbon dioxide produced per 100 calories of energy. Consequently we would expect to find on the first day of fasting, where the greater amount of glycogen is consumed, the larger carbon dioxide thermal quotient. It is important to recognize that in comparing these quotients the variations in the total 24 hours' heat production from experiment to experi- ment are without significance. In general, the carbon dioxide thermal quotients are greatest on the first day of the fast. After the relative proportions of protein, glycogen, and fat katabolized in the body have attained a constancy, the quotient has a tendency to remain constant, in general not far from 30.7. The lowest carbon dioxide- thermal quotient observed in any of the experiments is 29.45 on the second day of experiment No. 81 and the highest 34.70 on the third day of food experiment No. 76. 1 The assumption that the absorbed oxygen combines only with carbon is obviously untenable. 508 Influence of Inanition on Metabolism. COMPARISON OF ALL FACTORS OF KATABOLISM. The different relations between the carbon dioxide and heat, and the methods of expressing the output per square meter of body surface and per kilogram of body-weight computed and tabulated in this report are those which have been Fig. 3. found by comparison to be most commonly used in the reports of experiments of a similar nature made in other laboratories. Obviously, various other rela- tions can be computed and it is believed that the data given in the preceding Comparison of Factors of Katabolism. 509 section of this report furnish all the statistical details for such computations as may be desired. The method of showing the relative katabolism by curves has been elaborated Fig. 4. for one typical experiment (No. 75 — a seven days fast), and the results are given in the two curves herewith. The curves are in large part self-explanatory. The uniformity between the curves for water vaporized, heat production, carbon dioxide, oxygen, water of 510 I N'FLUENCE OF INANITION ON METABOLISM. organic hvdrogen, and pulse is especially well marked. It would appear, then, that these factors of katabolism are in a general way directly proportional to the pulse rate even with the inadequate records made in this experiment, and this curve emphasizes more perhaps than could any series of figures, the im- portance of making records of pulse rate in metabolism experiments. Antici- pating this extreme regularity between the pulse rate and the factors of meta- bolism, in the experiments Nos. 79 to 89 inclusive especial care was taken to secure the pulse and respiration rate (see p. 228). The protein, fat, and glycogen katabolized and the amounts of the urinary constituents are shown in the second set of curves. The curve for the protein obviously follows directly the nitrogen of urine. The relation between the katabolism of fat and glyco- gen is shown, since as the katabolism of glycogen decreases, the fat katabolized increases in nearly every instance. The energy of the urine follows in a general way the organic matter and the carbon, but there is little, if any, relation be- tween the nitrogen and the other factors in the urine. ENERGY BALANCE. The increased accuracy in the apportionment of the katabolism between the three compounds — protein, fat, and carbohydrate — in the experiments during inanition, makes the estimate of the energy of material katabolized from the body much more accurate than in the older experiments in which there was no exact knowledge of the amounts of glycogen katabolized, and in all proba- bility the estimated energy of material oxidized in the body computed from the amounts of protein, fat, and glycogen katabolized results in much closer approx- imations to the actual energy transformations than have as yet been available to physiologists. In the experiments during inanition previously reported from this labora- tory 17° the estimated energy of material katabolized was compared with the heat production. The store of glycogen in this comparison was assumed to have remained unaltered. Furthermore, while the heat elimination was in all probability determined with reasonable accuracy the heat production was not measured as accurately as in the present series of fasting experiments. The results show that the differences in three experiments between the esti- mated energy from material actually oxidized and the net heat outgo were some- what over 100 calories. In these three experiments (Nos. 36, 39, and 42), the energy measured as heat was less than the estimated energy of material actually oxidized in the body. The percentage error was — 5.1, — 5.4, and — 5.2 per cent for the three experiments, respectively. On the other hand, on the 2 days of experiment No. 51, the agreement between the energy of material actually oxidized and that measured as heat was extremely close. No wholly satisfactory explanation appears at present for this agreement. 170 U. S. Dept. Agr., Office of Expt. Sta. Bui. 136. Energy Balance. 511 Experiment No. 51 immediately followed a 1-day experiment, No. 50, in which the subject was distinctly ill. He was compelled to stop the work experiment in which he was engaged and lie down during the day, consuming but a small quantity of food. Therefore, the effect of the preliminary day before the 2 days of fasting may have been such as to cause errors either in the results of analyses or the heat measurements. There is a strong probability, too, that after the severe muscular work on the 3 days preceding experiment No. 50, the store of glycogen in the body had been heavily drawn upon and, therefore, did not undergo material alteration during fasting. Against this latter suppo- sition is the fact that for the 3 days of experiment No. 49, although the subject worked very severely, he was supplied with a very large amount of carbo- hydrates in the diet, and analyses showed that during the last 2 days of the experiment, there was a small gain of carbon to the body. On the contrary, there was a marked loss of carbon with probably heavy drafts upon previously- stored glycogen in the 1-day experiment No. 50, which immediately preceded the 2 days of fast. The results, then, in all of the earlier experiments would imply that a very material error is present when the assumption is made that the glycogen content of the body remains constant, save in experiments where the store of glycogen has previously been considerably depleted. Since the apparatus used in connection with these experiments permitted the direct measurement of heat, the estimated energy of material oxidized from the body can be compared directly with the heat production, and since the law of the transformation of energy holds true in the human body, the accuracy of the method of estimating the heat production from the total katabolism can be checked. For purposes of comparison, the estimated energy of katabolized body material, the total heat production, and the differences between the total heat production and the energy from body material, expressed both in terms of calories and per cent, are given in table 244. The widest discrepancy noted on any day during inanition is on the fifth day of experiment No. 73 where the estimated energy of katabolized body material was 66 calories greater than the total heat production, a discrepancy amounting to -|- 4.3 per cent. On the last 2 days of experiment No. 74 (with food), there were discrepancies of -j-4.3 and -(-3.9 per cent, respectively. Usually the discrepancies are extremely small. Thus in the average of 43 days of experiments without food, the estimated energy of katabolized body material, 1937 calories, was but 13 calories greater than the average total heat production, a discrepancy of -+- 0.7 per cent. A somewhat greater discrepancy was observed in the smaller number of food experiments, the estimated energy from body material, 1879 calories, being 20 calories greater than the total heat production, corresponding to a discrepancy of -f- 1.1 per cent. The aver- age for all the experiments here published covering 53 days, with and without food, shows a discrepancy of -f- 0.7 per cent. :> l ■> Influence of Inanition on Metabolism. Table 244. — Comparison of computed and measured heat production, in metabolism experiments with and without food. Experi- ment number. Subject and date. Estimated energy of katabolized body material Total heat pro- duction. Energy from body material greater (+) or less (— ) than output. Amount. Proportion 59. 68. 69. 71. 73. 75. 77. 79. 80. 81. Experiments without food. B.F.D., 1903: Dec. 18-19 Dec. 19-20 Dec. 20-21 A.L.L., 1904: Apr. 27-28 Apr. 28-29 A.L.L., 1904: Dec. 16-17 Dec. 17-18 Dec. 18-19 Dec. 19-20 S.A.B., 1905: Jan. 7- 8 Jan. 8- 9 Jan. 9-10 Jan. 10-11 S.A.B., 1905: Jan. 28-29 Jan. 29-30 Jan. 30-31 Jan. 31-Feb. 1 Feb. 1- 2 S.A.B., 1905: Mar. 4- 5 Mar. 5- 6 Mar. 6- 7 Mar. 7-8 Mar. 8- 9 Mar. 9-10 Mar. 10-11 S.A.B., 1905: Apr. 8- 9 Apr. 9-10 Apr. 10-11 Apr. 11-12 H.E.S., 1905: Oct. 13-14 ... Oct. 14-15 C.R.Y., 1905: Oct. 27-28 Oct. 28-29 A.H.M., 1905: Nov. 21-22 Nov. 22-23 Calories. 2122 2115 2159 2181 2179 1972 2166 2069 2008 2013 1860 1800 1653 1847 1831 1782 1678 1614 1796 1790 1785 1734 1636 1547 1546 1885 1910 1775 1770 1961 2046 1974 2125 1751 1763 Calories. 2080 2107 2102 2167 2217 1951 2163 2035 1958 1970 1844 1746 1606 1866 1791 1739 1663 1548 1765 1768 1797 1775 1649 1553 1568 1874 1880 1840 1807 1951 2047 1954 2099 1729 1781 Calories. + 42 + 8 + 57 + 14 -38 + 21 + 3 + 34 + 50 + 43 + 16 + 54 + 47 -19 + 40 + 43 + 15 +66 + 31 + 22 —12 — 41 — 13 — 6 -22 + 11 + 30 -65 -37 + 10 — 1 + 20 + 26 + 22 -18 Per cent. + 2.0 + 0.4 + 2.7 + 0.6 — 1.7 + 1 + 0 + 1 + 2, + 2.2 + 0.9 + 3.1 + 2.9 -1.0 + 2.2 + 2.5 + 0.9 + 4.3 + 1.8 + 1.2 — 0.7 -2.3 -0.8 — 0.4 — 1.4 + 0.6 + 1.6 -3.5 -2.0 + 0.5 + 1.0 + 1.2 + 1, -1 Energy Balance. 513 Table 244. — Comparison of computed and measured heat production, in metabolism experiments with and without food — Continued. Experi- ment number. Subject and date. Estimated energy of katabolized body material. Total heat pro- duction. Energy from body material greater (+) or less (— ) than output. Amount. Proportion. 82 H.C.K., 1905: Nov. 24-25 Calories. 2261 2494 Calories. 2222 2477 Calories. + 39 + 17 Per cent. + 1.8 + 0.7 Nov. 25-26 H.R.D., 1905: Dec. 5-6 1960 1914 + 46 + 2.4 85 Dec. 6-7 1855 1907 -52 -2.7 N.M.P., 1905: 89 Dec. 9-10 , 2117 2273 2109 2305 + 8 -32 + 0.4 — 1.4 Dec. 10-11 D.W., 1906: Jan. 10 11 2197 2150 + 47 + 2.2 Jan. 11-12 2284 2254 + 30 + 1.3 Experiments with food. 70 A.L.L., 1904: 72 Dec. 20-21 2089 2253 2474 2104 2223 2457 -15 + 30 + 17 -0.7 + 1.3 + 0.7 Dec. 21-22 Dec. 22-23 S.A.B., 1905: 74 Jan. 11-12 1727 1676 + 51 + 3.0 8.A.B., 1905: Feb. 2-3 1711 1691 + 20 + 1.2 76 Feb. 3-4 1653 1669 1585 1607 + 68 + 62 + 4.3 + 3.9 Feb. 4-5 , S.A.B., 1905: Mar. 11-12 1772 1699 1743 1937 1767 1728 1753 1924 + 5 -29 -10 + 13 + 0.3 -1.7 -0.6 + 0.7 Mar. 12-13 Mar. 13-14 , Average of experiments: Without food (43 days). With food (10 days) 1879 1859 + 20 + 1.1 All experiments (53 days) 1926 1912 + 14 + 0.7 Discrepancies such as appear in certain of the experiments between the estimated energy of katabolized body material and the total heat production may be caused by a number of factors. The assumptions involved in the com- putations as used with the present form of apparatus are (1) that there are no differences in the amounts of partially oxidized material in the body at the end of each day; and (2) that the material in the urine represents the incompletely oxidized nitrogenous fragment of the protein molecule. The influence of any 33 514 Influence of Inanition on Metabolism. errors involved in these assumptions on the total metabolism can hardly be estimated with our present knowledge of physiological chemistry. Since for the average of all the experiments, the discrepancy between the computed and measured heat production is almost insignificant, it would appear that it is reasonable to assume that in the few instances where noticeable discrepancies appear there are specific errors other than those involved in the assumptions above outlined, which have inevitably crept into the chemical and physical measurements. Fortunately, the simultaneous determination of the oxygen absorbed and carbon dioxide and heat production furnishes an unusual check on the accuracy of the method in general and the computations in particular. From the ratios between the oxygen and carbon dioxide and the heat, clues regarding the causes of the few marked discrepancies observed in the com- parisons may be obtained. It is necessary, therefore, to compare the data given in table 244 with those in table 243. The most noticeable discrepancies between the computed energy of body material and the total heat production are to be observed on the last day of experiment No. 73, the third day of experiment No. 77, and the last 2 days of experiment No. 74. Examining the carbon dioxide and oxygen thermal quotients for these days, certain valuable conclusions may be drawn. Thus, on the fifth day of experiment No. 73, the oxygen thermal quotient is abnormally high while the carbon dioxide thermal quotient is approximately normal for this particular experiment. Hence, on this day, at least, the figures point strongly to the conclusion that there was an error in the oxygen determination rather than in the heat determination, and consequently the discrepancy of +4.3 per cen+ implies that the heat production was accurately determined while the estimated energy of katabolized body material was somewhat too high. Again, on the third day of experiment No. 77, the oxygen thermal quotient is abnormally low while the corresponding carbon dioxide thermal quotient is not abnormal. It is evident, then, that in this case the error is again with the determination of oxygen, so that the total heat production is probably not far from correct. Another factor enters into food experiment No. 74. The oxygen-thermal quotient is high on both days but the carbon dioxide thermal quotient is likewise abnormally high, hence it would appear that on these 2 days the error is with the measurement of heat rather than the computed products of katabolism. It is of course not impos- sible that minor compensating errors may enter into the comparison of these three factors, but certainly the simultaneous determination of the carbon diox- ide elimination, oxygen consumption, and heat production is of the greatest value in deducing the correct balance between the total heat production and the estimated energy of katabolized body material. It is, furthermore, signifi- cant that none of the ratios seem to point toward marked discrepancies in the carbon dioxide determination, bearing out the results of check tests which show tli at the carbon dioxide determinations with this apparatus are extremely Energy Balance. 515 accurate. While, then, it is possible with accurate determinations of carbon dioxide and oxygen, in addition to the ultimate analyses of the urine, to com- pute the total heat production with an accuracy in all probability well within 2 or 3 per cent and with a large number of experiments an average accuracy of less than 1 per cent, accurate heat measurements are essential in any study of the energy transformations in the body. The carbon dioxide determination alone is insufficient, likewise the oxygen determination, although it would appear that the relation between the oxygen consumption and heat production is much more constant on the different days of the fast than that between the carbon dioxide output and heat production. From the molecular weights and heats of combustion of fat and carbohydrate the amount of energy liberated when 1 gram of oxygen is used to burn either of these bodies may be computed. The energy resulting from the combustion of 1 gram of carbon in the form of fat or carbohydrate may also be found. From the results of the computations it will be seen that the heat production per gram of oxygen in the combustion of both fat and carbohydrate is nearly the same, while the heat production per gram of carbon dioxide produced is considerably more in the case of the combustion of fat than in the combustion of carbohydrate. Hence, it can be seen that irrespective of whether fat or glycogen is burned in the body, the energy production per gram of oxygen consumed will always be nearly the same, while the energy output per gram of carbon dioxide will vary, depending upon the relative amounts of fat and carbohydrate oxidized. Theoretically, there should be no discrepancy between the estimated energy of the material oxidized in the body and the total heat production, when com- puted from factors as accurately determined as were the nitrogen, carbon dioxide, and oxygen of these experiments. The fact remains, however, that experiments of this nature are being made with an extremely intricate and complex organism and not in a test tube or flask. The probable limit of error we have arbitrarily set as not far from 1.5 per cent. About half of these experiments come inside of this estimated limit of error, and yet during fasts the assumptions regarding the intermediary metabolism are greater perhaps than in any other class of experiments. On the whole it is doubtful if the experi- ments could have been conducted so as to secure much closer agreement. Cer- tain known errors which could not be properly corrected unavoidably crept into certain of the experiments. The carbon dioxide and oxygen thermal quo- tients, however, suggest strongly the nature of the error in all experiments with marked discrepancies. Undoubtedly many other unknown errors likewise found their way into the conduct of the experiments and the computations. On the whole, however, there is an agreement between the computed energy and that directly determined that is as satisfactory as physiological experiments could ordinarilv demand. 5iG Influence of Inanition on Metabolism. Experiments with Food. All the observations made on man during inanition would imply that during a prolonged fast the whole alimentary tract is quiescent."1 Consequently the ingestion of food after a prolonged fast would naturally be expected to result in a more marked stimulation of peristalsis and metabolism in general, and at first sight it would appear that a study of the influence of the ingestion of food could best be made immediately after a prolonged fast. Data regarding the influence of the ingestion of food can be obtained in three different ways: (1) by studying the influence on the digestion of food in general, with special reference to the effect of the long quiescence of the ali- mentary tract on the degree of the absorption of food materials after fasting; (2) by observing the effect on the total metabolism of the ingestion of food after fast; and (3) by studying the course of the recovery of the body to its normal condition including both a study of the length of time required for recovery and the extent to which the losses during inanition were compensated. This problem is closely related to that attending study of recoupment after fever. The first of these methods demands for its success that the amount of food ingested be large, and that a correct separation of the feces be obtained. The large amounts of food are necessary to produce the maximum effect and like- wise to produce a sufficient amount of feces to permit proper separation. If the normal conditions of metabolism are established in a few days, in order to notice the particular effect of the transition the metabolism should be most carefully studied on the first day. It is practically impossible even under the most advantageous conditions of experimenting to separate the feces corresponding to the food of any one day. In experiments in which a special diet is ingested for 2 days, the separation may be made, although only with considerable difficulty. But when the previous period has been one of inanition, with the delayed expulsion of fecal matter belonging to the food period prior to the fast, the difficulties of separating the feces for 2 or even 3 days are considerably increased. The second method of study, namely, observation of the effect on the total metabolism to be observed during the height of digestion, should also be accompanied with large amounts of food; for as has been shown by a number of other researches in which the respiratory quotient has been the criterion, the effect of the ingestion of food is relatively not large, and consequently the quantities of food ingested in experiments of this kind should be as large as possible. mBoldireff (Archives des Sciences Biologiques (1905), 11, p. 1) has, however, noted periodical movements of the intestine even when no food has been ingested. Influence of Ingestion of Food. 517 The third method, namely, study of the course of the recovery after fasting, obviously requires a longer time than the 2 or 3 days' sojourn in the respiration calorimeter after a fasting period. It is necessary, therefore, to resort to studies of metabolism which can be made outside of the respiration chamber, namely, those of the intake and output of nitrogen, sulphur, phosphorus, etc., and such studies as have been made with S. A. B., in connection with these experiments receive special discussion beyond. Unfortunately, the effect of the ingestion of food immediately after the fast could not be studied in these experiments under conditions which even approximated the ideal conditions outlined above. In connection with the fasting experiments here reported four experiments in which food was ingested were made, during which the subjects remained inside the respiration chamber continuously from 1 to 3 days. It was soon seen that a practical difficulty was experienced that had not been anticipated, namely, that it was difficult, if not indeed dangerous, to administer a large amount of food on the first day fol- lowing a fast. Consequently, the quantity of food administered had to be very much restricted and the effect on metabolism and digestibility was correspond- ingly diminished. Usually not until the second day after the fast could any considerable amount of food be taken. Thus the important observations on the first day were made under great disadvantages. Furthermore, the separation of feces was in every case extremely difficult. While no difficulty was experienced in the separation at the end of the food period, the sharp separation of the feces properly apportioned to the food period from the mass of fecal matter that had been retained in the colon during the whole of the fast was extremely difficult. The feces that had been for a long time in the alimentary tract had been deprived of their water, so that, at least in one instance, the water content was as low as 50 per cent. This mass became more or less mixed with the softer, fresher feces as they were forced through the colon and it was separated after expulsion only with diffi- culty. Under these circumstances, then, the results obtained during the 3-day calorimeter experiments in studying the problem of the effect of the ingestion of food after fasting from the standpoint of the influence on digestibility and on the total metabolism have a value considerably less than it was hoped they would have. It was, furthermore, apparent that this method of studying the effect of the ingestion of food after fasting was impracticable, and that an entirely different type of experiment must be used for a proper study of this problem. Accord- ingly, during the past year experiments have been in progress in this laboratory in which large quantities of food were ingested after a short fast (12 to 18 hours) rather than after a period of inanition covering several days. The method of studying the effect of the ingestion of food after a short fast was suggested by the data secured in the fasting experiments here reported. ;,ls Influence of Inanition on Metabolism. These data seemed to indicate that in general the fasting katabolism is reached after 24 hours of inanition. The results of the experiments on food ingestion after short fasts will he published in a subsequent report. Since these embody a type of experiment somewhat different from the food experi- ments here reported, it seems best to include with the fasting experiments the data obtained from the food experiments immediately following them. The results obtained have much general interest other than that regarding the matter of digestibility or the effect on general metabolism, and although the special discussion of the effect of the ingestion of food on metabolism will be taken up in a subsequent publication certain features are here discussed. Furthermore, in the discussion of the fasting experiments the data for the sub- sequent days with food have frequently been inserted in the derived tables for the purpose of comparison. Thus, some discussion of the effect of ingestion of food has already been given. It is the purpose of this discussion to outline some of the more important points to be emphasized in connection with the data for food experiments Nos. 70, 72, 74, and 76. Aside from the data obtained from the experiments with food inside the respiration chamber, the data for the food, feces, and urine were obtained for periods of considerable length following food experiment No. 76 and fasting experiment No. 77, and many interesting data on the third point, i. e., recoup- ment after fasting, were secured. In these experiments the total income of food and the outgo in urine and feces were determined. Since the primary object was to study the intake and output of nitrogen, they have been designated Nitrogen Metabolism Experi- ments, Nos. 1 and 2. The results of these experiments on the long-continued ingestion of food are here presented and discussed. The detailed statistical data have been given on pages 274 to 299. Pbecautions in Feeding. It is commonly considered among professional fasters and others whose experience with fasting subjects has been extended, that especial caution should be exercised in administering food after a fast. The ingestion of a large amount of solid food into the stomach after a long period of rest, causes serious disturbance. Hence, in the experiments reported herewith, special care was taken to avoid any complications which might result from over-feeding. In general, the subjects were given small amounts of milk (less than half a glass) from time to time, and no solid food was administered until several hours after the fast ended. The previous experience of the subject S. A. B. had convinced him that his i'nsts were best broken by taking orange juice. Hence, his diet for the first day contained only orange juice and small quantities of milk. In subsequent experiments,, the orange juice was supplemented by a small amount Digestibility of Food. 519 of apple and in one case by a solid food in the shape of gluten crackers and shredded wheat. After fasting experiment No. 77, the subject did not remain in the chamber and his diet for that day taken outside of the chamber was chosen at will. During the day he consumed milk, bread, tomatoes, lemons, and oranges, but the total quantity of food ingested was small, furnishing but 43.47 grams of protein, and 1311 calories of energy (see table 176). On the next day the quantity of protein and energy was more than doubled, while on the third day, the subject consumed 120 grams of protein and 3649 calories of energy. The food experiments inside the respiration calorimeter which fol- lowed certain of the fasting experiments, formed only a transitional period from fast to complete food, inasmuch as the diet consumed was generally lim- ited, especially as regards protein. Thus, in experiment No. 76, the food fur- nished but 37.6 grams of protein and 1841 calories of energy per day, and while the energy was sufficient to meet the requirements of the body during the inac- tive period in the calorimeter, the protein was not sufficient to equalize the draft upon body protein. On the first day after food experiment No. 76, i. e., the fourth day after the end of the fast, the subject consumed a diet furnishing 131.5 grams of protein and 5064 calories of energy, and on the following day, the amounts were even larger, i. e., 167.5 grams of protein and 6612 calories of energy. While, therefore, the subjects were unable to consume large amounts of food on the first day after fast, there seemed to be no difficulty in ingesting large quantities on the second day. This is especially noticeable on the second day after experiment No. 77, as has been pointed out above. THE DIGESTIBILITY OF FOOD AFTER INANITION. The experiments with food which followed the fasting experiments in the respiration calorimeter furnish some data regarding the digestibility of food material after fasting. It is conceivable that after a long period of rest the stimulus due to the ingestion of food would result in an excessive flow of the digestive juices and increased peristalsis, and in an expulsion of feces from the colon. On the other hand, the quantity of fecal material required to distend the colon before defecation might result in a delay in the expulsion of feces. The feces from ordinary digestion experiments are preceded by feces of a character not especially different from those to be analyzed. Thus, separation between the experimental feces and those resulting from the food preceding is not based upon physical condition. Eegularity in the time of meals and the amounts taken is also of importance in securing uniform con- sistency of the feces. The irregularity in the ingestion of food, especially on the first day following fasting, is one of a number of factors which tend to hinder the accurate separation of feces. The small amounts of food ingested on the first day, in some instances containing only 38 grams of protein, 520 Influence of Inanition on Metabolism. obviously increase markedly the percentage error, since any slight error in the separation of feces would affect materially the coefficient of digestibility of protein. Diet. — Milk was the chief article of diet in this series of food experiments in the calorimeter. In the nitrogen metabolism experiments a very much more elaborate selection of foods was made, but they consisted in large part of milk, cream, fruit, and vegetables, although occasionally meat was taken. Feces. — In the discussion of feces after fasting it was pointed out that in none of the calorimeter experiments here reported was there a sufficiently sharp and accurate separation of feces made to enable any special quantity to be designated as fasting feces. In only one instance was there any approximation to such a separation and a subsequent consideration of the digestibility of the food in the period following fast shows that in all probability the so-called fasting feces were in reality a portion of the feces resulting from food preced- ing the experiment. Separation of feces. — It has already been stated that much difficulty was experienced in obtaining sharp separations of the feces. In ordinary diges- tion experiments where reasonably marked alterations in diet between the experimental period and those preceding and succeeding it are concerned, the technique of the separation of feces is one that is considerably more elaborate than is commonly considered. In the case of the separation of feces imme- diately following a fasting period the problem is even more complex, and extreme difficulty has been experienced in nearly every instance in designating the fecal mass that properly belonged to the period of experimenting with food. As has been stated above, no typical fasting feces were isolated in these experi- ments. Consequently, whatever errors were involved in the separation of the feces between the preliminary food days and the first feces of the feeding experiment proper increased considerably the liability of error in the proper apportionment of feces to the experimental period. The marked alteration in the character of the feces resulting from their long sojourn in the colon was an added difficulty.172 Analyses of the feces. — Ordinarily, in the analyses of feces, the determina- tions of the nitrogen, carbon, organic hydrogen, water, ash, and heat of combustion presented no unusual difficulties. The determination of fat, how- ever, demands special comment. On the assumption that only small amounts of soap exist in the feces, the amounts of fat have usually been determined by extraction with ether. In the case of the feces from experiment No. 70 it was found that the amount of fat was small, but when the heat of combustion 1,2 In one instance 20.4 grams of feces were passed which contained but 50 per cent of water. This is the lowest proportion of water ever found in feces in this laboratory, and indeed we are not familiar with any records of fecal material that contained so small a proportion of water as this. Feces. 521 was computed from the proximate analysis, there was a very decided discrep- ancy between the computed heat of combustion and that actually found by determination with the bomb calorimeter, and the per cent of ash was much higher than ordinarily found. It thus appeared that some material of a high energy content insoluble in ether was present in the feces, and by treating the feces with hydrochloric acid and alcohol, large amounts of fatty acids were liberated from the very considerable quantity of soap present. After this pre- liminary treatment with hydrochloric acid and alcohol, the feces were dried and again extracted and the percentage of fat was increased enormously, so much, in fact, that the sum of the percentages of water, protein, fat, and ash was more than 100 per cent. This, of course, eliminated all carbohydrates and further showed that the determination of one of the constituents was slightly too high. A consideration of the quantity of ash in the feces showed that this was unusually large. Since the modified method for fat determination had shown the presence of a large amount of soap, in the absence of definite ash analyses, it was assumed that the soap present was combined with calcium, and when charred the calcium remained in large part in the form of calcium carbonate, which would hold carbon dioxide at the low temperature of incinera- tion of ash and thus yield a result for the percentage of ash higher than should be obtained. On the basis of this assumption it was decided to estimate the ash by difference and hence the sum of the percentages of water, protein, and fat deducted from 100 per cent was taken as the percentage of ash. Unfortunately, the presence of this large proportion of calcium soap in the feces was not discovered until the samples of feces from some of the experi- ments had been entirely used in making different determinations, so that it was necessary in some instances to assume a percentage of fat based upon the marked increase in the ether soluble material of other samples after treatment with hydrochloric acid and alcohol. The feces belonging to metabolism experiment No. 74 gave, on analyses by the method ordinarily used : For sample No. 3838, 9.06 per cent protein, 4.41 per cent fat, 7.93 per cent carbohydrates, 3.52 per cent ash, with a heat of combustion of 1.533 calories per gram; for sample No. 3839, 6.54 per cent protein, 3.28 per cent fat, 15.90 per cent carbohydrates, and 6.77 per cent ash, with a heat of combustion of 2.016 calories per gram. After experience had shown that by the new method of analysis a larger percentage of fat was obtained in the feces for experiments No. 70 and No. 76, it was thought advisable to treat these two sample of feces by the same method, but unfortunately both samples had previously been exhausted. It seemed reasonable to suppose, however, that the determinations of fat in these two samples as originally made were too low, and that this was the case seemed to be borne out, particularly in sample No. 3839, not only by the high ash content but also by the fact that their heats of combustion by calculation varied very widely from those actually obtained. ,v>-j Influence of Inanition on Metabolism. In experiment No. 76, a redetermination of the fat in the feces changed the result from 3.53 per cent to 7.22 per cent, i. e., it practically doubled the amount of fat in the feces, and reduced correspondingly the percentage of carbohydrates. It was, therefore, decided as the only known method of approximation to the true figures for fat for the feces passed in experiment No. 74 to double the percentage of fat in the two samples, giving, therefore, for No. 3838, 8.82 per cent fat, in place of 4.41, and for No. 3839, 6.56 in place of 3.28, which reduced the carbohydrates in the first instance to 3.52 per cent, and in the second to 12.62 per cent. The result of doubling the amounts of fat in these two samples was to bring the heats of combustion as calculated much nearer to the heats of com- bustion found by burning in the bomb calorimeter. The calculated heats of combustion had previously been much lower than those found, namely, 1.256 calculated against 1.533 found, and 1.338 calculated against 2.016 found. The calculated results after the fat was doubled were 1.487 calculated against 1.533 found and 1.510 calculated against 2.016 found. It seems evident, therefore, that the per cent of fat assumed for No. 3839 is still far too low. DIGESTIBILITY OF FOOD IN SHORT EXPERIMENTS. Recognizing that feces consist chiefly of metabolic products rather than of undigested food, and in view of the fact that the methods of separating them are imperfect, it is impossible to state absolutely the digestibility of any given food material. In discussing these experiments, the custom is followed of assuming that the feces are composed of undigested food. Since the metabolic products of feces result primarily from the ingestion of food, it is not seriously wrong to make such an assumption. Furthermore, digestion experiments have value only for comparison with previous experiments on digestibility, and consequently it seems best to adhere to the usual method of computation. The digestibility of the simple diets given in the short food experiments inside the calorimeter is shown in table 245. It was impossible to separate the feces for experiment No. 72, so that no computations regarding the digestibility were made. As is the common experience with diets containing a large amount of milk the digestibility of all the nutrients is high for the three experiments. In experiments Nos. 70 and 74, the average digestibility for protein was about 94.5 per cent, fat, 96, and oarbohyd rates, 99 per cent. Slightly more energy was absorbed in experiment No. 74 than in experiment No. 70, the average coefficient of digestibility being 92 per cent. In discussing the digestibility of the food in experiment No. 74, it is important to bear in mind that the determinations of fat and carbohydrates in the feces are at best approximate. Indeed, it is very much to be questioned whether in diets so simple as these any appreciable amounts of undigested carbohydrates appear in the feces and in all probability the carbohydrates were completely absorbed. Digestibility. 523 In experiment No. 76 there was a much larger proportion of vegetable protein owing to the shredded wheat and gluten crackers and hence the digestibility of protein is very considerably less. The coefficient of digestibility in this instance is only about 78 per cent. The corresponding coefficients for the fat, carbohydrates, and energy are 88.7, 96.0, and 86.6. Table 245. — Data regarding digestibility of food- Nos. 70, 74, and 76. -Metabolism experiments Labora- tory number. Kind of food. (a) (6) Weight Total. of mate- orgamc !?„i matter nal- \(d + e+f) (c) Nitro- gen. (d) Pro- tein (2VX 6.25).1 (e) Fat. (/) Carbo- hy- drates. (9) Ash. (h) Heat of com- bus- tion. Experiment No.70. 3806-8 Milk 3773. . Plasmon 3809. Grams. 4844.50 19.23 Total. Feces Urine Ain't available. Coefficients of di- gestibility 4863.73 182.90 3826. 3827. 3828. 3829. 3838-9 3841. 3842. . 3844. 3845. 3843. 3858. Grams. ' Grams. 1003.23; 23.42 15.68 2.29 1018.91 49.89 25.71 1.08 969.02 Per ct. 95.1 24.63 Per ct. 95.8 Grams. Grams. Grams. 146.29 642. 231214.71 14.33 .03 1.32 160.62 642.26 6.79 43.10 216.03 Grams. 30.66 1.66 32.32 11.58 153.83 599.16 Per ct. , Per ct. 95.8 93.3 216.03 Perct. 100.0 20.74 Perct. 64.2 Cals. 7637 93 7730 448 276 7006 Perct. 90.6 Experiment No. 74- Grams. \ Grams Milk 3600.00 715.32 368.10 50.31 938.70 110.37 105.00 97.92 Apple Orange juice Graham crackers. Total. Feces. . .. Urine. . . . Am't available . . . Coefficients of di- gestibility 5011.80 106.30 973.92 25.68 Experiment No. 76 Milk Apple Shredded wheat Gluten crackers. . . . Orange juice 948.24 Per ct. 97.4 Grams. 17.28 .12 .75 .96 19.11 1.27 17.84 Per ct. 93.3 Grams. 107.28 .54 4.59 5.43 117.84 7.92 109.92 Per ct. 93.3 Grams. Grams. 433.08 174.96 49.77 105.78 81.66 10.83 443.91 7.84 436.07 Perct. 98.2 Grams. 23.76 .81 3.48 1.71 Cals. 5322 198 411 468 412.17 9.92 29.76 5.95 6399 196 217 402.25 Per ct. 97.6 23.81 5986 Per ct. Per ct. 80.0 93.6 Grams. 1952.10 368.10 533.40 30.30 938.70 Total. Feces Urine 3822.60 332.80 Am't available. . . Coefficients of di- gestibility Grams. 365.04 47.97 485.28 28.62 114.45 1041.36 79.34 962.02 Perct. 92.4 Grams. 3.33 1.29 9.48 4.32 .93 Grams. 20.81 8.06 59.25 27.00 5.81 19.35 4.26 15.09 Per ct. 78.0 120.93 26.63 994.30 Per ct. 78.0 Grams Grams. Grams. \ Cals. 202.83 141.40, 4.50 2529 .... 39.91' 1.05 195 9.33 416.70 9.60 2184 .09 1.53! .72 165 .... 1108.64 3.84 450 212.25 708.18 24.03 28.68 19.71 12.51 5523 478 260 188.22 679.50 Per ct. 88.7 Per ct. 96.0 7.20 4785 Per ct. Per ct. 36.5 86.6 1 With some food materials other factors are used. 1 Ash by difference. See p. 521. 52 l Influence of Inanition on Metabolism. If the nature of the foods ingested and the imperfect method of separation of feces as well as the short duration of the experiments are taken into con- sideration, an examination of the results of these digestion experiments shows no abnormality. It is, therefore, impossible to distinguish from the data any effect of inanition on the digestibility of the kinds of food here ingested during these short experiments. digestibility of food in the nitrogen metabolism experiments. In the long feeding experiments which immediately followed food experi- ment No. 76 and fasting experiment No. 77, it was possible to study only the absorption of protein and energy. Pressure of other work prevented the com- plete analysis of both food and feces, and since these digestion experiments are at best but approximate, it was deemed inadvisable to sacrifice other important determinations. The digestibility or absorption, computed in the usual man- ner, is recorded in table 246 for both nitrogen metabolism experiments. The feces were separated at the end of each week. The first nitrogen metabolism experiment continued 25 days, while the second experiment lasted 2 weeks. The coefficients of digestibility for each week during the experiments have been computed. Table 246. — Digestibility (availability) of protein and energy in nitrogen metabol- ism experiments Nos. 1 and 2. Food. Feces.1 Absorbed Energy of urine.2 Energy avail- able. Per cent digested Average for whole experi- ment. Nitrogen Metabolism Experi- ment No. 1. Mar. 14 to 20, 1905 : Total weight kilos.. Protein grams.. Energy calories . . Mar. 21 to 27, 1905 : Total weight kilos.. Protein grams.. Energy calories.. Mar. 28 to Apr. 3, 1905 : Total weight kilos.. Protein grams.. Energy calories . . Apr. 4 to 7, 1906 : Total weight kilos.. Protein grams . Energy calories.. Nitrogen Metabolism Experi- ment No. %. Apr. 12 to 18, 1905: Total weight kilos... Protein grams Energy calories... Apr. 19 to 25 : Total weight kilos... Protein grams... Energy calories... 24.476 923.67 38,700 23.096 953.70 39,193 19.687 760.39 27,118 9.658 391.75 14,996 18.271 858.01 23,764 19.141 849.31 23,445 0.395 102.70 2011 .664 169.72 3417 .443 121.83 2297 .234 63.18 1082 .349 107.77 1667 .358 103.39 1713 820.87 36,689 783.98 35,776 638.56 24,821 3-J8.57 13,913 750.24 22,097 745.92 21,732 668 666 718 427 890 787 36,021 35,ii6 24J03 13,486 21,207 20',946 88.9 93.1 82.2 89.6 84.0 88.9 83.9 89.9 87.4 89.2 87.8 89.3 84.9 90.4 (7.6 S9.3 1 Partially dried. 2 Energy of urine assumes 9 calories for each gram of urinary nitrogen. General Physical Condition. 525 An examination of the figures shows that the digestibility of both protein and energy was much greater the first week than in the three following periods of the first experiment. This might imply a more thorough absorption of food during this first week, but owing to changes in the diet, the food for the different weeks was not strictly comparable, though still remaining in most cases mainly a milk, fruit, and vegetable diet. An examination of the kinds and amounts of food consumed (see pp. 277 to 288) shows that there was a tendency to diminish the amounts of cream consumed. The most marked change in the diet was due to the fact that a large amount of peanut butter was consumed during the first week, but much less in the weeks which followed. In the second experiment the digestibility during both weeks is practically the same. The per cent of protein digested averages somewhat more than in the first nitrogen metabolism experiment but the absorption of energy in both is practically the same, i. e., about 90 per cent. It would appear, then, that there is no definite information furnished regard- ing the influence of inanition on the digestibility of food and it is clear that experiments on a plan markedly different from that here employed are neces- sary to study the problem satisfactorily. INFLUENCE ON GENERAL PHYSICAL CONDITION. None of the subjects of these experiments showed any symptoms of an alarming nature as a result of inanition (see p. 334), but there was in all cases a daily loss of body-weight, and in general a decrease in pulse rate, respiration rate, and muscular strength. The influence of the subsequent ingestion of food on these grosser functions is of importance. Body-weight. — In table 247 are given the body-weights of the subjects for the food experiments which continued inside the respiration chamber. Table 247. — Body-weights in metabolism experiments with food. Experiment number. Subject and duration of experiment. First day. Second day. Third day. 70 A.L.L., Dec. 20 to 22, 1904 Kilos. 70 . 952 70.841 Kilos. 70.841 70.653 Kilos. 70.653 70.680 72 S.A.B., Jan. 11, 1905 — .111 -.188 + .027 55.069 55.189 74 S.A.B., Feb. 2 to 4, 1905 + .120 .... 54 . 997 55.228 55.228 55.570 55.570 55.770 76 S.A.B., Mar. 11 to 13, 1905 . + .231 + .342 + .200 55.836 55.531 55.531 56.057 56.057 56.319 — .305 + . 526 + .262 526 I m ••].! ence of Inanitiox ox Metabolism. The data in table 247 show that on the first and second days of experiment No. 70, and the first day of experiment No. 76, the subjects still continued to lose weight, as would probably be implied by the fact that the amount of food ingested was barely enough for maintenance. The body-weights of S. A. B. in the nitrogen metabolism experiments are given in table 248. Table 248. — Daily body-weight (including clothes) — Nitrogen metabolism experi- ments Nos. 1 and 2. Experiment number and date. weight Experiment number and date. Body- weight. Nitrogen metabolism exp't No. 1. 1905, Mar. 19, after urinating and Mar. 20 Kilos. 60.99 61.70 61.88 63.50 62.69 62.78 62.76 63.23 63.57 63.85 63.59 63.60 63.55 63.79 63.60 63.34 63.60 63.72 63.93 63.84 Nitrogen metabolism exp't No. 2. 1905, Apr. 13 Kilos. 58.30 59.17 60.32 60.75 60.62 61.15 61.02 61.63 61.82 61.22 61.80 61.32 Apr. 15 Mar. 21 Apr. 16 Mar. 22 Apr. 17 Apr. 18 Mar. 23 Mar. 24 Apr. 19 Apr. 20 Mar. 26 Apr. 21 Mar. 37 Apr. 22 Mar. 28 Apr. 23 Mar. 29 Apr. 24 Mar. 30 Mar. 31 Apr. 1 Apr. 2 Apr. 3 Apr. 5 Apr. 6 During these experiments relatively large amounts of food were consumed and the body-weight attained constancy about 8 or 9 days after the close of each fast. Of special interest is the table of body-weights after fasting of some of the subjects of the 2-day experiments. These subjects were required to come to the laboratory and be weighed for several weeks after the conclusion of their fasts. Table 249 shows that there was a marked tendency for all of these subjects to gain weight and indeed ultimately to exceed their initial weights. A rough comparison with the weights of a number of college students taken during the same period of the year shows that while there was a general tendency for the weights to increase during this portion of the year, all the subjects of these experiments increased in weight very considerably more than did their fellows. This seems to suggest that a short period of inanition may so stimulate anabolism as to result subsequently in a permanent increase in body-weight. Influence on General Metabolism. 527 Body temperature, pulse, and strength, as affected by the ingestion of food. — Under the corresponding sections in the discussion regarding fasting experi- ments it has been pointed out that the ingestion of food resulted in an increase in the pulse and a marked increase in the strength. By examination of the detailed data for pulse, and dynamometer tests, the relative increase in these factors may be observed. Table 249.— Body- weights after fasting. C. R. Y. A. H. M. H.C. K. H. R. D. N. M. P. Experiment 80. Experiment 81. Experiment 82. Experiment 83. : Experiment 85. Date. Weight. Date. Weight. Date. Weight. Date. Weight. Date. Weight. 1905. Kilos. 1905. Kilos. 1905. I KUo8m 1905. Kilos. 1905. Kilos. Oct. 27. 1 69.34 Nov. 21. 162.02 Nov. 24. m.49 ! Dec. 5. 155.64 Dec. 9. !67.63 Nov. 28. 61 . 53 Dec. 4. 73 . 84 Dec. 8. 54.29 Dec. 12. 66.69 1906. Dec. 4. 62.23 Dec. 11. 73.44 Dec. 11. 54.08 Dec. 18. 67.92 Jan. 22. 2 71. 37 Dec. 11. Dec. 18. 62 . 87 62.87 Dec. 18. 1906. 73.38 Dec. 18. 1906. 55.41 1906. Jan. 20. 67.06 1906. Jan. 29. 76.68 Jan. 8. 56.31 Feb. 14. 69.09 Jan. 15. J 64.62 Jan. 15. 57.23 May 19. 68.86 Jan. 22. 64.58 Jan. 22. 58.38 Jan. 30. 58.30 1 Beginning of fast. - Made comparable with weights during fast by including 0.54 kilo, assumed weight of underclothes. INFLUENCE ON GENERAL METABOLISM. The general factors of metabolism, the urine, respiratory products, and heat production are all influenced by the ingestion of food and in the discussion beyond attention is called to the degree to which they are affected. Urine, ingested water, and water vaporized. — The ratios of the water ex- ereted in the urine and of the water of respiration and perspiration to water ingested during the food experiments are given in table 250. The amount of water ingested in the food and drink is above 1400 grams in all instances and hence the ratios of water in urine to ingested water do not exhibit such wide variations as occur in the fasting experiments. In general, about 72 per cent of the water of food and drink is excreted in the urine. The ratios of water of respiration and perspiration to water ingested are also more nearly constant. Nitrogen balance. — That the small amounts of nitrogen in the food of experiments ISTos. 70, 72, 74, and 76 were not sufficient in any instance to restore complete nitrogen equilibrium, since the body lost nitrogen persistently, may be seen from an inspection of the nitrogen balances given in table 251. ■V>S Influexce of Ixaxitiox ox Metabolism. Table 250. — Ratio of water excreted in different ways to water ingested in metabolism experiments with food. Experi- ment number. Subject and date. 70. 72. 74. 76. A.L.L., Dec. 20, 1904. Dec. 21, 1904. Dec. 22, 1904. (.a) Amount of water ingested in food and drink. (fi) (c) (d) Amount Water Ratio of water in of in urine urine. urine. to water ingested (C-r-Cl). (e) Water of respira- tion and per- spiration. (/) Katio of water of respi- ration and per- spiration to water ingested (e-=-a). Total, 3 days Average per day. . . S.A.B., Jan. 11, 1905... S.A.B., Feb. 2, 1905 . . Feb. 8, 1905 Feb. 4, 1905 Total, 3 days Average per day. . . S.A.B., Mar. 11, 1905... Mar. 12, 1905... Mar. 13, 1905 . . . Total, 3 days Average per day. . . Average of all ex- periments with food Grama. Grams. 1445.33 |1031.50 1561.89 |1042.70 1599 28 i 624.70 4606.50 2698.90 1535.50 899.63 Grams. 991.37 1007.04 586.16 2584.57 861.52 0.686 .645 .367 0.561 Grams. 840.30 1002.78 1059.07 2902.15 967.38 0.581 .642 .662 0.630 1837.94 1496.40 1463.33 0.796 544.84 0.296 1939.34 2358.54 2066.24 6364.12 2121.37 1518.90 1894.40 1605.00 5018.80 1672.77 1485.94 1866.17 1579.48 4931.59 1643.86 0.766 .791 .764 0.775 584.88 527.61 524.88 1637.37 545 . 79 0.302 .224 .254 0.257 1941.61 1964.01 1991.01 1735.80 1313.70 1603.20 5896.63 1965.54 4652.70 1550.90 1695.18 1290.45 1576.11 4561.74 1520.58 0.873 .657 .792 0.744 616.81 582.30 611.39 1810.50 603.50 0.318 .296 .807 0.307 1870.52 1386.63 1354.12 0.724 689.49 0.369 Table 251. -Balance of income and outgo of nitrogen in metabolism experiments with food.1 Experi- ment number. Subject and date. Total. Per kilogram of body-weight. In food. In feces. In urine. Loss to body. In food. In feces. In urine. Loss to body. 70 A.L.L., 1904: Dec. 20-21 Gms. 8.54 Gms. 0.36 Gms. 13.04 Gms. 4.86 Gms. 0.120 Gms. 0 . 005 Gms. 0.184 Gms. 0.069 Dec. 21-22 8.60 .36 9.84 1.60 .122 .005 .139 .023 Dec. 22-23 8.57 .36 10.15 1.94 .121 .005 .144 .027 72 S.A.B., 1905: Jan. 11-12 6.24 .... 10.66 4.42 .113 .193 .080 74 S.A.B., 1905: Feb. 2-3 6.37 6.37 6.37 .42 .42 .42 10.74 8.25 6.78 4.79 2 . 30 .83 .116 . 115 .114 .008 .008 .008 .195 .149 122 .087 .042 .015 Feb. 3-4 76 S.A.B., 1905: Mar. 11-12 6.45 1.42 10.17 5.14 .116 .026 .183 .092 6.45 1.42 7.15 2.12 .116 .025 .128 .038 Mar. 13-14 6.45 1.42 7.82 2.79 .115 .025 .139 .050 v„, -a T~5l UULa on nurogen metaboli rsos. <6 and 77, respectively, see table 256. Protein Katabolism. 529 The marked decrease in the nitrogen excretion on the second and third days with food shows a tendency of the body to adjust itself rapidly to the new conditions of protein ingestion and it is worthy of note that on the third day of experiment No. 74, after a prolonged draft upon body protein not only during the fasting period but likewise during the first two days of food, the body was nearly in nitrogen equilibrium on but 6.37 grams of nitrogen. It is greatly to be regretted that this experiment was not continued with the inges- tion of small amounts of protein and the nitrogen balance further studied. In subsequent fasting experiments this point will be taken into consideration. During the period, then, with low nitrogen intake there was a marked tendency for the body to adjust itself to new conditions and retard the great drafts upon body protein made during the fasting experiment. Further discussion of the effect of the ingestion of nitrogenous material upon the total katabolism of protein is taken up in connection with the data for the nitrogen metabolism experiments. 'Protein Jcatabolized in metabolism experiments with food. — The ingestion of protein after fasting generally results in a diminution in the katabolism of protein. The course of the protein metabolism has already been indicated in discussing the income and outgo of nitrogen (table 251), and the actual amounts of protein katabolized in the food experiments are recorded in table 252. Taele 252. — Protein katabolized per kilogram of body-weight in metabolism experiments with food. Experi- ment number. Subject and duration of experiment. First day. Second day. Third day. Total. Per kilogram of body- weight. Total. Per kilogram of body- weight. Total. Per kilogram of body- weight. 70 72 74 76 A.L.L., 1904: Dec. 20 to 22 Grams. 78.24 63 .96 64.44 61.02 Grams. 1.104 1.160 1.169 1.096 Grams. 59.04 49.50 42.90 Grams. 0.835 .894 .769 Grams. 60.90 40.68 46.92 Grams. 0.862 .731 .835 S.A.B., 1905: 8.A.B., 1905: Feb. 2 to 4 S.A.B., 1905: Mar. 11 to 13 66 . 92 1.132 50.48 0.833 49 . 50 0.809 A comparison of the results given in this table with table 226 for experi- ments without food shows that in experiment No. 70, the protein katabolized on the first day with food was practically the same as that on the last fasting day. But on the second and third days the katabolism of protein decreased to that of the first day of the fast. In experiment No. 72 with S. A. B. the pro- tein katabolized on the first day with food was essentially the same as that on 34 :,:]() Influence of Inanition on Metabolism. the last day of fast, while in experiment No. 74 the katabolism of protein was 4 grams larger than on the last day of experiment No. 73. The extremely small amounts katabolized on the second and third days of experiment No. 7 I were 10 to 20 grams less than the lowest amount katabolized on any day of experiment No. 73. Similarly, with experiment No. 76, the katabolism on the first day was essentially that of the last day of the fast, but on the second and third days with food the katabolism was greatly diminished. The tendency of the body to attain nitrogen equilibrium with such small quantities of protein in the diet is of especial interest when compared to the length of time required to attain a condition of equilibrium with excessive quantities of nitrogen in the diet.173 Fat katabolized. — The katabolism of fat in experiments with food following fast is shown in table 253. Table 253. — Fat katabolized per kilogram of body-weight in metabolism experiments with food. Experi- ment number. Subject and duration of experiment. First day. Second day. Third day. Total. Per kilogram of body- weight. Total. Per kilogram of body- weight. Total. Per kilogram of body- weight. 70 72 74 76 A.L.L., 1904: Dec. 20 to 22 Grams. 158.59 147.47 138.30 137.95 Grams. 2.237 2.675 2.509 2.477 Grams. 193.87 112.48 101.92 Grams. 2.740 2.030 1.827 Grams. 188.90 124.26 66.67 Grams. 2.673 2.232 1.187 S.A.B., 1905: S.A.B., 1905: Feb. 2 to 4 S. A. B., 1905: Mar. 11 to 13 145.58 2.475 136.09 2.199 126.61 2.031 With regard to experiment No. 70, it is seen that large amounts were kata- bolized on the second and third days, which may in part be accounted for by the high pulse rate and febrile temperature observed at the end of this experi- ment. The amount of fat broken down in experiment No. 72 was 15 grams more than on the last day of experiment No. 71. In experiment No. 74 on the other hand, the fat katabolism on the first day was 10 grams less than on the last day of experiment No. 73, and throughout this experiment it was unusually low. In experiment No. 76, it increased on the first day but 5 grams over the last day of fasting and then continued small for the remaining two days of the food experiment. On the whole, then, the katabolism of fat was diminished on the ingestion of food after fasting. 73 See discussion of gain of nitrogen during nitrogen metabolism experiment No. 1, p. 541. Balance of Matter and Energy. 531 Glycogen katabolized. — Owing to the marked losses of glycogen observed during fasting, the katabolism of glycogen in food experiments following fast is of especial interest. The amounts katabolized are shown in table 254. Table 254. — Glycogen katabolized per kilogram of body-weight in metabolism experiments with food. Experi- ment number. Subject and duration of experiment. First day. Total. Per kilogram of body- weight. Second day. Total. Per kilogram of body- weight. Third day. Total. Per kilogram of body- weight. 70..., 72... 74 ... , 76... A.L.L., 1904 Dec. 20 to S.A.B., 1905 Jan. 11. S.A.B., 1905 Feb. 2 to S.A.B., 1905: Mar. 11 to 13. 22. Grams. 56.46 10.31 27.82 56.37 Average. 37.74 Grants. 0.796 .187 .505 1.012 0.625 Grams. 35.95 87.88 131.62 Grams. 0.508 1.586 2.359 Grams. 99.90 75.23 217.47 85.15 1.484 130.87 Grams. 1 .414 1.351 3.870 2.212 At the end of the longer fasts the amount of glycogen katabolized was as a rule not much over 15 to 20 grams per day. Hence, it is seen that on the inges- tion of food in practically all instances there was a very considerable increase in the katabolism of glycogen,* an increase which persisted throughout the whole experiment. The largest amount katabolized on any day with food was on the last day of experiment No. 76, namely, 217.5 grams. BALANCE OF MATTER AND ENERGY. In experiments with food the main comparison is that which shows in how far the matter and energy of the food sustained the body and to what extent material was gained or lost by the body. From the determinations of the protein, fat, and glycogen katabolized and the quantities of the protein, fat, and carbohydrates absorbed from the food, the gains or losses of these com- pounds in the body may be determined. They are recorded in the first three columns of table 255. The gain or loss of preformed water is obtained from the water of food and drink and total water katabolized. This is recorded in the fourth column, while in the last column the gain or loss of energy is recorded. Although from 38 to 53 grams of protein were ingested in the food, the body lost on the first day of each experiment about 30 grams of protein. This was with a diet which contained practically enough energy for maintenance. On the second day the loss of protein was considerably less, a decrease, in fact, of more than one-half from that on the first day in all the experiments. On the * Carbohydrates as a whole. :,■;•? Influence of Inanition on Metabolism. third day of experiments Nos. 70 and 76 there was an increase over the second, while on the third day of experiment No. 74 there were but 5 grams of protein lost. For all the experiments there was an average loss per day of about 19 grams of protein. During experiment No. 70 the body stored fat in considerable amounts on the first day, but stored much less on the second day, and none on the third. In the 1-day experiment, No. 72, there were 25.8 grams of fat lost. Experi- ment No. 74 showed a slight loss on the first day and substantial gains on the 2 succeeding days with an average gain for the whole experiment of about 9 grams per day. In experiment No. 76 there was a marked loss of fat on the first day, a loss of about 25 grams on the second, and a gain of about 10 grams on the third, the average loss being a little more than 25 grams per day. Table 255. — Protein, fat, carbohydrates, preformed water, and energy gained or lost in metabolism experiments with food. Experi- ment number. Subject and date of experiments. Protein, gain (+) or loss (— ), Fat, gain (+) or loss (— ). Carbohy- drates, gain (+) or loss (— ). Pre- formed water, gain(+) or loss (— ). Energy, gain (+) or loss (— ) 70... 72... 74... 76.. A.L.L., 1904: Dec. 20-21 . Dec. 21-22. Dec. 33-23. Average per day. S.A.B., Jan. 1905 11.. S.A.B., 1905: Feb. 2-3.. Feb. 3-4. Feb. 4-5.. Average per day. .A.B., 1905: Mar. 11-12 Mar. 12-13 Mar. 13-14 Average per day. Orams. — 29.16 — 9.60 — 11.64 Orams. + 34.06 + 4.68 - .40 Grams. + 16.64 + 30.57 — 20 90 — 16.80 +12.78 | + Orams. — 160.3 -203.6 + 149.6 .77 - 71.5 Cals. + 213 + 131 — 121 + 74 -30.66 -25.78 + 61.99 i- 37.7 -108 -28.74 -13.80 -4.98 - 4.50 + 31.45 + 9.57 + 103.70 + 43.53 + 56.37 -15.84 + 8.84 + 67.83 + 95.3 + 52.2 + 106.1 + 127.4 + 289 + 415 + 400 + 36S -30.84 -12.72 -16.74 -20.10 — 61.20 -25.12 + 10.12 - 25 . 40 + 133.68 + 58.36 - 27.47 — 179.0 + 285 . 4 !— 9.9 + 54.86 + 32.1 -310 — 112 — 140 -154 1 Water of feces taken into account. The carbohydrate balance shows that on only 2 days of this series of experi- ments was there a loss of glycogen to the body. On the last day of experi- ments Nos. 70 and 76 there was a loss of 21 and 27 grams, respectively. This gain of glycogen in an organism which had previously been deprived of glyco- gen indicates strongly the tendency of the body to replace its lost glycogen. On the first days of experiments Nos. 74 and 76, respectively, there were over 100 grams of glycogen gained. The relation of the gain or loss of glycogen to the gain or loss of protein is extremely interesting. Thus, aside from the Becovery after Inanition. 533 first day of each experiment, the largest gains of glycogen are accompanied by the smallest losses of protein, while in the 2 instances where glycogen was lost there is a noticeable increase in the loss of protein over the preceding day. It is interesting to note that in some instances where fat was actually lost from the body, there was, nevertheless, a marked gain in glycogen, as, for instance, on the first 2 days of experiment No. 76. On the other hand there was a slight gain of fat on the third day of experiment No. 76 accompanied by a loss of 27 grams of glycogen. The gains and losses of preformed water underwent marked variations. In experiment No. 70 there was a marked loss on the first 2 days of the experi- ment followed by a gain of 150 grams on the third day. In experiment No. 74 there was a gain on all 3 days. In experiment No. 76 there was a loss of 179 grams on the first day and a gain of 285 grams on the second. No striking relation appears between the gain or loss of preformed water and the other gains or losses to the body. The energy shows an average gain of 74 calories per day in experiment No. 70. In experiment No. 72 the loss was 108 calories. The diet in experi- ment No. 74 was plainly in excess of maintenance since there was an average of 368 calories stored per day. On the other hand, with experiment No. 76 the diet was deficient in energy and there was an average loss of 154 calories. Of special interest, perhaps, is the fact that on the first day of experiment No. 76, with a loss of 61 grams of fat and a total loss of 210 calories of energy, there was a positive gain of 134 grams of glycogen. A corresponding observa- tion may be made in connection with experiment No. 72 where there was a marked loss of fat and loss of energy, but nevertheless a gain of glycogen. The most noticeable feature of the effect of the ingestion of food following fast is the tendency of the body to restore its depleted glycogen. RECOVERY AFTER INANITION. Experiments on physiological fasting have been numerous, and yet the problem of the recovery after fast has been but imperfectly studied. The nitrogen metabolism experiments which followed experiments Nos. 76 and 77 gave opportunity for a more or less extended study of the degree and rapidity with which the body recovered its equilibrium so far as the nitrogen, phos- phorus, and sulphur were concerned. Data for computing the storage of fat and glycogen are not obtainable after the subject leaves the respiration calo- rimeter, so that aside from general observations regarding the amount of food eaten and the body-weight, there is no direct evidence at hand regarding the gain of fat or glycogen. Dietetic habits. — The subject of these experiments was of average height and weight (see p. 107), but in recovering from the fasts he consumed liberal amounts of food in proportion to his body-weight and muscular activity. An 534 Influence of Inanition on Metabolism. examination of the statistical tables (pp. 277 to 284) shows that on some days the food consumption was enormous. While no accurate record was kept of the amount of muscular activity engaged in by this subject between the fasts, except for an occasional walk of considerable length, no extraordinary physical muscular exercise was noted. While hardly of a phlegmatic tem- perament, he was not muscularly active or quick. Urine. — From the statistical table 183 the determinations in the urine may be obtained. No special abnormalities were noticed in the urine at any time during the experiments. Nitrogen balance. — Since nitrogen was determined not only in the food and feces but also in the urine, the data were at hand for striking a complete nitrogen balance in both nitrogen metabolism experiments. While the nitro- gen in each day's food and urine was definitely determined, it was not possible to know the exact excretion of fecal nitrogen per day since daily separations were impossible. Accordingly, the feces for each week were separated, dried, and analyzed and the nitrogen per day was calculated according to the methods explained on page 294 and included in table 181. From the quantities of nitrogen in the food, feces, and urine, therefore, the gain or loss could be computed. The results of these computations are given in table 256 herewith. It is believed that the daily gains or losses of nitrogen obtained by this method as indicated in the last column of this table are not far from the true values, although it is to be borne in mind that the quantities excreted in the feces were determined only for the week and not for each individual day. The most striking feature of this table is the enormous gain of nitrogen shown during the first and second weeks of the first experiment. Even during the third week, there is a material gain amounting to more than 22 grams. Nitrogen equilibrium is approximated only on the last few days. During the 4 days of the last period there was a gain of but 5.1 grams of nitrogen or 1.27 grams per day, as compared with a gain of 30 grams for the corresponding 4 days of the first week. During the second nitrogen metabolism experiment there is likewise a very considerable gain of nitrogen during both weeks, but contrary to the first experiment, the larger gain was in the second week. Since the subject was obliged to leave Middletown, the experiment was stopped at the end of 2 weeks. During the first week the body gained an average of over 3 grams and in the second week an average of over 4.5 grams of nitrogen per day. Becovery after Inanition. 535 Table 256. — Intake and output of nitrogen — Nitrogen metabolism experiments Nos. 1 and 2. [Amounts actually determined.] Date. (a) In food. (6) In feces.1 (c) In urine. Gain ( + ) or loss ( — ) a-(b + c). Experiment No. 1. 1905. First week: Mar. 14-15. Mar. 15-16. Mar. 16-17. Mar. 17-18. Mar. 18-19. Mar. 19-20. Mar. 20-21. Total, 1st week. . Average per day. Second week: Mar. 21-22. Mar. 22-23. Mar. 23-24. Mar. 24-25. Mar. 25-26. Mar. 26-27. Mar. 27-28. Total, 2d week. . , Average per day. Third week : Mar. 28-29. Mar. 29-30. Mar. 30-31. Mar.31-Apr.2. Apr. 2-3 Apr. 3-4 Total, 3d week.., Average per day. Fourth week:2 Apr. 4-5. . . Apr. 5-6. . . Apr. 6-7,... Apr. 7-8. . . Total, 4 days Average per day. Grams. 20.55 25.09 21.65 14.90 26.34 17.10 22.14 147.77 21.11 22.14 19.90 15.01 21.55 20.80 25.55 27.64 152.59 21.80 16.62 12.83 11.74 24.69 22.65 19.14 13.99 121.66 17.38 14.03 17.91 18.01 12.73 62.68 15.67 Grams. 2.28 2.79 2.40 1.66 2.92 1.91 2.46 16.42 2.35 3.94 3.53 2.69 3.83 3.70 4.56 4.92 27.17 3.88 2.67 2.05 .89 .96 .63 .06 .24 19.50 2.79 2.27 2.90 2.91 2.06 10.14 2.54 Grams. 12.61 13.19 11.07 6.31 9.55 11.89 9.55 74.17 10.60 10.17 9.35 8.02 8.64 11.95 11.26 14.53 73.92 10.56 12.54 9.27 9.00 26.14 12.77 10.11 79.83 11.40 12.65 11.56 11.87 11.36 47.44 11.86 Grams. + 5.66 + 9.11 + 8.18 + 6.93 + 13.87 + 3.30 + 10.13 + 57.18 + 8.17 + 8.03 + 7.02 + + + + 4.30 9.08 5.15 9.73 + 8.19 + 51.50 + 7.36 + 1.41 + 1.51 + .85 + 13.61 + 3.31 + 1.64 + 22.33 + 3.19 - 0.89 + 3.45 + 3.23 - 0.69 + 5.10 + 1.27 1 Daily amount calculated. See explanation, p. 291. 2 A fasting expei-iment immediately followed this nitrogen metabolism experiment. There was a lag in the feces for the feeding experiment so that the separation was not obtained until April 13. 536 Influence of Inanition on Metabolism. Table 256. — Intake and output of nitrogen — Continued. Date. (a) In food. Kb) In feces.1 (c) In urine. Experiment jVo. 2. 1905. First week: Apr. 12-13. Apr. 13-14. Apr. 14-15. Apr. 15-16. Apr. 16-17. Apr. 17-18. Apr. 18-19. Total, 1st week. . Average per day. Second week: Apr. 19-20. Apr. 20-21. Apr. 21-22. Apr. 22-23. Apr. 23-24. Apr. 24-25. Apr. 25-26. Total, 2d week. . . Average per day. Grams. 7.10 16.23 20.04 23.81 21.67 28.98 19.45 137.28 19.61 Grams. 0.90 2.03 2.51 2.98 2.72 3.63 2.45 17.22 2.46 Grams. 11.14 11.65 15.08 13.89 16.53 17.93 12.72 98.94 14.13 (d) Gain (+) or loss ( — ) a-(b + c). Grams. + + + + + + 4.94 2.55 2.45 6.94 2.42 7.42 4.28 + 21.12 + 3.02 19.05 22.72 20.22 14.93 21.57 20.28 17.12 135.89 19.41 2.31 2.76 2.46 1.82 2.63 2.46 2.08 16.52 2.36 13.80 14.18 12.09 11.13 10.66 13.30 12.19 87.35 12.48 + + + + + + + 2.94 5.78 5.67 1.98 8.28 4.52 2.85 + 32.02 + 4.57 1 Daily amounts calculated. See explanation, p. 294. Intake and output of phosphoric acid and sulphur. — The phosphoric acid (P205) and sulphur (S03) were determined in the food and urine for each day and in the feces for each week. The apportionment of the phosphoric acid and sulphur for the feces for each day was carried out on substantially the same plan as that for nitrogen, namely, the total phosphorus in the food for the week was taken as 100 per cent and the proportion of this amount ingested each day was taken as the proportion for each day of the total phosphorus de- termined in the total feces for the week. The sulphur was apportioned by a like method. From the phosphoric acid and sulphur in the food, feces, and urine, the gains or losses were computed. These are recorded in table 257. Two features of special interest may be noted in these tables: First, the excretion of a small amount of phosphorus in the urine during the first week of the first feeding experiment compared with that during fasting experiment No. 75 ; second, the marked gain of phosphorus after inanition. By examining the data in column c of table 257 and comparing it with the phosphoric acid excretion in experiment No. 75, it will be seen that for this first week the phosphoric acid in the urine was on the whole very much smaller than during the fasting period. In the experiment following No. 77 the same peculiarity may be observed. The most noticeable feature in the data for phosphoric acid is the gain on all the days of the first nitrogen metabolism experiment. The gains were very considerable during the first 2 weeks and not so marked in the last 2 periods. Eecovery after Inanition. 537 Table 257. — Intake and output of phosphoric acid (P2OJ and sulphur (80J in nitrogen metabolism experiments Nos. 1 and 2. Date, 1905. Phosphoric acid (P2O5). (a) In food. (6) In feces.1 (c) In urine (by ti- tration) (d) Gain (+) or loss (— ) a—(b+c). Sulphur trioxide(S03). (e) In food. it) In feces.1 (0) In urine. (ft) Gain (+) or loss (— ) e-(f+g). Experiment No. 1. First week: Mar. 14-15 Mar. 15-16 Mar. 16-17 Mar. 17-18 Mar. 18-19 Mar. 19-20 Mar. 20-21 Total, 1st week. . Average per day. Second week: Mar. 1-22 Mar. 22-23 Mar. 23-24 Mar. 24-25 Mar. 25-26 Mar. 26-27 Mar. 27-28 Total, 2d week. . Average per day, Third week: Mar. 28-29 Mar. 29-30 Mar. 30-31 Mar.31-Apr. 1 Apr. 1- 2 Apr. 2- 3 Apr. 3- 4 Total, 3d week.. Average per day. Fourth week: Apr. 4-5 Apr. 5-6 Apr. 6-7 Apr. 7-8 Total , 4 days . . . Average per day. Experiment No- 2. First week: Apr. 12-13 Apr. 13-14 Apr. 14-15 Apr. 15-16 Apr. 16-17 Apr. 17-18 Apr. 18-19 Total, 1st week. .. . Average per day. . . Grams. 10.257 15.696 11.823 9.685 8.922 7.833 10.770 Orams. Grams. 1.930 2.953 2.224 1.822 1.656 2.196 1.268 1.086 1.679 1.978 1.474 2.322 2.026 2.118 Grams. + 6.671 + 10.547 + 8.331 + 6.777 + 5.265 + 4.037 + 6.626 74. 986' 14. 108 12.624 10.712 2.015 1.804 Grams. 4.547 5.086 2.851 .836 4.025 2.834 7.652 Grams. 0.613 .685 .384 .113 .542 .382 1.032 Grams. 2.440 + 2.176 + 1.622 + 1 . 126 - 1.838 + 1.876 + 1.769 + + 48.254 27.831 3.751 + 6.893 3.976| .536 12.847 Grams. 1.494 2.225 .845 .403 1.645 .576 4.851 + 11.233 1 835 + 1.605 7.926 2.540 9.161 2.935 5.175 1.658 8.510: 2.727 10.3851 3.328 9.176 2.940 10.558 3.384 60.891 19.512 8.699 2.787 2.631 + 2.755 3.839 0.829 2.482 + 3.744 3.938 .850 .... ... 2.274 + 1.243 2.733 .590 . • . . ... 2.456 + 3.327 3.916 .845 .... 2.670 + 4.387 4.403 .950 .... ... 2.966 + 3.270 4.797 1.035 .... ... 3.234 + 3.940 5.143 1.110 .... 18.713 + 22.666 28.769 6.209 2.673 + 3.238 4.110 .887 .... . . . 6.311 4.701 4.401 7.301 7.376 5.922 5.281 2.122 3.571 1.580 2.513 1.480 2.635 2.454: 2.479 1.991: 3.459 6.660 0.618 .608 .286 + 3.084 + 1.775! 2.840 + 41.293 13.881 5.899 1.983 .472 .666 21.678 + 5.734 3.097+ .819 3.606 2.730 2.586 '4.187 .4.015 3.541 2.426 23.091 3.299 0.697 .528 .500 .810 .776' .685 .469 4.4651 .638 5.025 5.651 7.263 4.413 22.352 5.588 1.677 1.886 2.423 1.472 2.934 3.093 2.992 2.162 7.458 11.181 1.8651 2.795 + 0.414 2.671 + .672! 3.560 + 1.848: 3.020 + .779 2.471 3.71311.731 .928 2.933 0.554 .741 .627 .513 1.906 2.435 .609 + 0.487 2.713 4.484 5.897 6.320 6.035 9.854 5.923 1.030 1.702 2.238 2.399 2.291 3.740 2.248 1.493 + 1.142 + 2.225 + 2.189 + 2.994I + 3.280 + 2.472 + 0.190 1.640 1.434 1.732 .750 2.834 1.203 1.522 3.183 3.624 4.312 3.805 5.084 3.155 41.226 15.648 15.795+ 9.783 24.685 5.889 2.235 2.256!+ 1.398, 3.526 0.242 .507 .577 .686 .605 .809 .502 3.928 .561 2.026 - 2.412 + 2.524' + 2.063 + 2 . 536 + 2.555! + 2.033 + 0.746 .264 .523 1.563 .664 1.720 .620 16.149 + 2.307 + 4.608 .658 1 Calculated, see table 180, p. 294. 538 Influence of Inanition on Metabolism. Table 257. — Intake and output of phosphoric acid (P2Ot) and sulphur (80a) — Continued. Date, 1906. Phosphoric acid (P2O5). 8ulphur trioxide (S03). (0) In food. (&) In feces.1 (c) In urine (by ti- tration). (d) Gain (+) or loss (— ) a-(b+c). (e) In food.