Nitrogen Partition in Repeated 
 
 Fasting 
 
 , 
 
 By PAUL LDWARD HOWL 
 
 B.5. University of Illinois, 1906 
 M.A. University of Illinois, 1907 
 
 THL5I5 
 
 c''C' r 
 
 SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENT FOR THE DEGREE 
 OF DOCTOR OF PHILOSOPHY IN PHYSIOLOGICAL CHEMISTRY IN THE 
 GRADUATE SCHOOL OF THE UNIVERSITY OF ILLINOIS 
 
 
 Easton, Pa. : 
 
 Eschenbach Printing Company 
 
 . 1911 
 
Nitrogen Partition in Repeated 
 
 Fasting 
 
 By PAUL LDWARD HOWL 
 
 B.5. University of Illinois, 1906 
 M.A. University of Illinois, 1907 
 
 THL5IS 
 
 SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENT FOR THE DEGREE 
 OF DOCTOR OF PHILOSOPHY IN PHYSIOLOGICAL CHEMISTRY IN THE 
 GRADUATE SCHOOL OF THE UNIVERSITY OF ILLINOIS 
 
 19IO 
 
 Easton, Pa. : 
 
 Eschenbach Printing Company 
 1911 
 
Digitized by the Internet Archive 
 in 2017 with funding from 
 
 University of Illinois Urbana-Champaign Alternates 
 
 https://archive.org/details/nitrogenpartitio00howe_0 
 

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 £W„ 
 
 NITROGEN PARTITION IN REPEATED FASTING. 1 
 
 By Paul E. Howe. 
 
 Historical. 
 
 The literature upon fasting is large and may be divided into two classes, 
 that presenting data upon (a) simple fasts of (i) short duration, and 
 dealing with changes in body weight, nitrogen partition, the blood, tem- 
 perature, secretions, and the effect of drugs, or only as a preparation for 
 the study of normal or pathological metabolism; (2) more or less pro- 
 found fasts, which include in addition to those changes mentioned above, 
 losses in the weight and differences in composition of the organs as com- 
 pared to the normal organs; and (6) repeated fasts in which (1) the ob- 
 ject was to study the effect of repeated fasting; or (2) fasts in which 
 the main object was not the study of repeated fasting, but in which the 
 subject was used repeatedly in various investigations, no regard being 
 given to the original conditions, such as body weight, nitrogen equilibrium, 
 etc. 
 
 Experiments on Repeated Fasting . — Investigations upon fasting have 
 been limited mainly to the lower animals, while those upon repeated 
 fasting are confined entirely to them unless the fasts of Sued 1 are to be 
 considered as repeated fasts, which is hardly proper, since each fast was 
 conducted under different circumstances as regards original condition, 
 body weight, time of year, climate, etc. 
 
 Repeated fasts have been conducted upon dogs by Albitsky. 3 His 
 investigations were made upon four dogs, two fasting absolutely, i. e. t 
 without water, to a loss in body weight of 40.4 per cent, and 37.6 per cent., 
 in twenty-nine days, and fifteen days respectively, and the second time 
 with water ad libitum , with a loss of 53.2 per cent, and 45.8 per cent, 
 in fifty-one and thirty days, respectively. The other two dogs fasted first 
 1 Reprinted from the Journal of the American Chemical Society , 33, 215, 1911. 
 
4 
 
 FASTING STUDIES. 
 
 with water, to losses of 41.5 per cent, and 38.7 per cent, in thirty-four days 
 and twenty-five days, respectively, and the second fast to a loss of 43.7 
 per cent, and 49.0 per cent, body weight, in thirty-two days, and thirty- 
 eight days respectively. Positive conclusions cannot be drawn from 
 Albitsky’s results. No study was made of the effect of a properly con- 
 trolled repeated fast, either complete or with water. Hence he had no 
 basis upon which to compare the effect of water upon the length of a 
 fast. A study of the results show in general that water seems to lessen 
 the breakdown of the body tissues, for in three out of the four cases the 
 dogs fasted a greater length of time for the same loss in body weight 
 when they received water than when no water was given. 
 
 Tuvim 4 subjected two dogs to three repeated fasts of three days each, 
 the first fast being absolute, and the two following with a large water 
 ingestion. A third dog was fasted but twice, the first fast being abso- 
 lute and the second with water. His experiments were to study the effect 
 of water upon fasting, and only the respiration data were observed. His 
 results did not show that water had any pronounced effect upon the 
 gas interchange, and no data are presented which would show the effect 
 of repeated fasting upon the nitrogen partition. 
 
 Schulz working with Mangold, 9 Stiibel 10 and Hempel 11 reports results 
 upon repeated fastings in dogs. One dog weighing 16.1 kilograms was 
 fasted for thirty-two days, when the- premortal rise seemed to have set 
 in, and then the animal was fed an insufficient nitrogenous diet for nine 
 days, during which time it stored 23 grams of nitrogen. This nitrogenous 
 feeding was followed by a seven-day carbohydrate feeding period with 
 a loss of 12.93 grams of nitrogen, leaving a storage of 10.07 grams of 
 nitrogen for the sixteen days, during which the animal was fed. This 
 feeding period was followed by another fast of fourteen days with a 
 further loss of 27.61 grams of nitrogen over that gained and lost during 
 the feeding. 
 
 This same dog was subjected to a similar experiment one month later. 
 At the beginning of this experiment the animal weighed 19.6 kilograms 
 and probably had greater fat stores than in the first experiment. The 
 dog was fasted for twenty-seven days, at the end of which time he was 
 so reduced in strength that they feared he would die. Schulz described 
 the condition as a collapse. Food was given, 400 cc. of milk on the first 
 day and 300 grams of meat per day for four days, restoring the dog to a 
 satisfactory condition, after which the dog was fasted further for sixty- 
 one days, at the end of which time the animal was very weak, although 
 no premortal rise had occurred. During this second series of fasts 
 the dog had lost 208.4 grams of nitrogen without regard to the nitro- 
 gen balance of the feeding period, while in the first series of fasts the 
 dog lost 183.2 grams of nitrogen. The daily average nitrogen excretion 
 
FASTING STUDIES. 
 
 5 
 
 during the second part of the first series and the first part of the second 
 series was less than that in the first fast, while the average nitrogen ex- 
 cretion in the final fast of sixty-one days was even lower than the nitro- 
 gen excretion of the feeding period when sugar was ingested. 
 
 Stiibel, 10 working on another dog, obtained a similar lowered nitrogen 
 excretion after a short feeding period in which the dog received an in- 
 sufficient food supply. 
 
 Manassein 5 reports experiments upon the repeated fasts of three rab- 
 bits in absolute fasting, and Albitsky 3 upon nine rabbits fasting either 
 absolutely or alternately with water. In Albitsky ’s experiments the 
 object was the study of the effect of water upon fasting and the following 
 conditions were in force in his work. The first three rabbits were fasted 
 absolutely while the remaining six rabbits were fasted alternately with 
 and without water. The fasts were continued to a loss of 40 per cent, 
 of the body weight and it was found that with each succeeding fast there 
 was a tendency toward a decrease in the daily loss in body weight. Al- 
 bitsky also observed that the recovery was slower with each succeeding 
 fast and that as a result of fasting the animals’ “oxidative” properties 
 were increased so that from one to one and one-half times as much food 
 and twice as much water were needed to keep the animal in a state of 
 constant body weight. Pashutin 2 concludes from the work of these 
 men that with the ingestion of water there is a slower loss in body weight 
 than would occur in absolute fasting. The conditions of experimentation 
 were not controlled accurately in these experiments and only general 
 deductions may be drawn from the results. 
 
 Kagan 6 conducted experiments in repeated absolute fasting upon pigeons 
 and showed in the birds results similar to those obtained upon dogs and 
 rabbits, i. e., a slower loss in body weight upon repeated fasting. This 
 decrease in the loss of body weight does not continue with each succeeding 
 fast but remains fairly constant. Albitsky 3 reports results upon pigeons 
 which confirm his findings upon rabbits. He also gives data showing 
 the influence of water upon fasting. 
 
 Ugrumoff 7 in two experiments upon pups showed that water exerted 
 a sparing influence upon the organs over the loss found by fasting when 
 a salt solution was given. Sadoven 8 working along the same lines sub- 
 jected rabbits to repeated fasts under varying conditions such as abso- 
 lute, with water, or with salt solution. He showed that salt solution 
 exerted a sparing influence upon the loss of body weight as compared 
 with complete fasting, while water exerted a still greater influence. 
 
 Bischoff and Voit 13 made a number of fasts, twenty-eight, upon one 
 dog, under widely varying conditions, no single fast being more than 
 eight days long and some were for only one to two days. 
 
 Seeland 14 conducted experiments on short repeated fasts with pigeons 
 
6 
 
 FASTING STUDIES. 
 
 and roosters and showed that those birds which were subjected to re- 
 peated fasts of one to three days’ duration gained weight more rapidly, 
 their tissues were more solid, and the birds hardier and stronger than 
 the control birds which were fed every day. 
 
 Richet 15 fed and fasted rabbits alternately in periods of six days; 
 under such treatment they died in about a month and a half. Dogs 
 subjected to a similar treatment lived about six months. 
 
 The work upon repeated fasting which has been cited cannot be in- 
 terpreted in any way to show the effect of repeated fasting with water. 
 The only cases in which the animals were fasted repeatedly and absolutely 
 were those of Albitsky’s rabbits which were fasted to a loss of 40 per cent, 
 of the body weight. The most of the data presented deal largely with 
 the changes in body weight. In some cases the respiration data, urine 
 volume, P 2 0 5 and urea excretion were determined but no accurate data 
 are presented upon nitrogen partition. 
 
 Short, repeated fasts upon men are reported by Bucker, 18 Smith, 17 
 Ranke, 18 Pettenkofer and Voit, 19 Oppenheim, 20 Sadoven, 21 and Richet. 22 
 
 Accurate studies of the nitrogen partition in fasting dogs have been 
 made but recently. Previous to the advent of Folin’s 32 method for crea- 
 tine and creatinine no accurate investigations were made showing the 
 nitrogen partition further than urea and ammonia, and in one or two 
 cases the excretion of some of the pathological constituents of the urine. 
 
 Osterberg and Wolf 33 report the nitrogen partition in an experiment 
 in which a dog was fed a low protein diet for a number of days and was 
 then given a large amount of casein, after which the animal was fasted. 
 Their findings cannot be considered normal nitrogen partition data for 
 a fasting dog, inasmuch as the ingestion of a large amount of protein 
 following a low protein diet would influence the nitrogen partition of the 
 succeeding fasting days. Underhill and Kleiner 34 report findings upon 
 the nitrogen partition in the urine of a fasting dog. Their experiment 
 extended over a period of fourteen days, water being taken in varying 
 amounts. Their results upon the nitrogen distribution during fasting 
 as compared with a well fed animal show (1) “A slightly increased per- 
 centage of ammonia nitrogen and a correspondingly diminished urea 
 percentage.” (2) “The total creatinine nitrogen does not bear a strict 
 relation to the total nitrogen” “nor is the quantity excreted con- 
 
 stant.” “ The relatively large output of creatin is also of special interest 
 although its significance at this time is obscure.” They report the pres- 
 ence of allantoin in the urine and also allantoin crystals in the urine 
 after standing, and hold that the presence of this substance in “ the urine 
 of starving dogs shows without question that allantoin may have an 
 endogenous origin.” With regard to the purine and allantoin excretion 
 
FASTING STUDIES. 
 
 7 
 
 they say that: “In inanition no fixed relationship appears to exist be- 
 tween the excretion of purine and allantoin.” 
 
 Schondorf, 35 working with a fasting dog, shows that the percentage 
 of urea nitrogen may fall as low as 75 per cent, of the total nitrogen. 
 
 Leffmann 36 reports results which show that the creatine and creatinine 
 excretions are fairly constant for animals of equal body weight, and that 
 creatine or creatinine ingested with food, by a well-nourished dog, are 
 seemingly excreted as such. Weber 37 shows that in fasting dogs as a 
 result of muscular activity creatinine is excreted, while with uniformly 
 nourished animals creatinine diminishes. Dorner 38 showed that fasting 
 had no influence upon the excretion of creatinine in rabbits but that 
 creatine appeared in increasing amounts. Bohtlingk 39 gives results 
 upon the nitrogen distribution in fasting cats and rabbits. 
 
 The more recent work upon fasting which deals with the occurrence 
 of creatine and creatinine has been presented by van Hoogenhuyze and 
 Verploegh, 40 Benedict, 41 Benedict and Diefendorf, 42 and Cathcart, 43 
 who report results which are in general quite similar, the creatinine remain- 
 ing fairly constant, with a slight decrease as the fast progresses. The 
 appearance of creatine in increasing amounts is reported by all except 
 van Hoogenhuyze and Verploegh, who did not determin creatine. Bene- 
 dict and Diefendorf report a decrease in creatine after the rise. 
 
 Results upon single prolonged fasts in the case of dogs have been re- 
 ported by the following investigators: Howe, Mattill and Hawk, 23 Falck, 24 
 Kumagawa and Miura, 25 Pfliiger, 26 Daddi, 27 Grandis, 28 Krawkaw, 29 
 L’abbe and Vitry 30 and Luciani and Bufalini. 31 The longest normal 
 single fast has been reported by Howe, Mattill, and Hawk. 23 In this 
 prolonged fast the animal fasted 1 1 7 days, during which time he received 
 a constant water ingestion. The next longest normal fast upon which 
 data are presented was reported by Falck. 24 This fast was continued 
 for 60 days. Kumagawa and Miura 25 report a fast of 98 days during 
 which time the dog was subjected to repeated doses of phlorhizin. This 
 fast cannot be compared with true fasts because the phlorhizin intro- 
 duces unknown factors. 
 
 Description. 
 
 Purpose and Plan . — The purpose of this investigation was to study 
 the effect of repeated fasting upon the nitrogen partition in the urine 
 of a dog. Changes in body weight were also observed. As a basis of 
 comparison the nitrogen partition of the urine was determined during 
 a normal feeding period for a number of days preceding the fast. 
 
 The plan of the experiment was as follows : The dog was brought into 
 nitrogen equilibrium and then permitted to fast, the fast continuing 
 until the inception of the premortal rise. At this time the animal was 
 fed, the feeding continuing until the dog had regained its original body 
 
8 
 
 FASTING STUDIES. 
 
 weight and was again in nitrogen equilibrium. Following this came 
 the second fast. 
 
 Subject. — The animal used in this experiment was a small fox-terrier 
 bitch, from one to two years old and weighing 3.41 kilograms (see Table 
 
 V, p. 40). She was in good physical condition, well muscled, and had 
 a medium layer of fat under the skin. Life in the experimental cage 
 did not seem to annoy her in any way. 
 
 Articles of Diet: Meat. — The meat fed was beef, which had been prepared 
 according to the method proposed by Gies 44 and modified by Hawk. 45 
 This method of preparation and preservation consisted in freeing the 
 meat from as much visible fat and connective tissue as possible. The 
 trimmed meat was ground three times in a sausage mill, using a smaller 
 plate each time. The meat was passed through the mill and mixed thor- 
 oughly between each grinding. This insured a uniform sample. After 
 the juices had been pressed out of the meat, it was sampled and then 
 wrapped in paraffin paper in cakes of about 400 to 600 g., placed in a 
 museum jar, frozen and kept in cold storage. This method of preser- 
 vation was very satisfactory. The meat remained sweet, and after a 
 period of six months the composition as regards the nitrogen content 
 was practically the same. The analysis of the meat at the time of prepara- 
 tion showed 3.62 per cent, nitrogen and after a period of six months 
 showed 3.69 per cent, nitrogen, or on a moisture-free basis when prepared 
 4.88 per cent.; and after six months 4.86 per cent, of nitrogen (see Table 
 
 VI, p. 41 ). 
 
 Cracker Dust. — Cracker dust was purchased already ground from 
 the manufacturer. This was thoroughly mixed, sampled, analyzed, 
 and kept in air-tight jars. 
 
 Lard. — Pure leaf lard was used. 
 
 Bone Ash. — A good quality of finely ground bone ash was used. 
 
 Water. — Ordinary distilled water was used, such as is supplied to the 
 chemical laboratory. 
 
 Method of Collection and Preservation of Excreta , etc. — The animal was 
 kept in an experimental cage such as described by Gies 46 and modified 
 by Hawk. 45 This cage was provided with a wire netting bottom of about 
 3 / 8 inch mesh. Below this wire bottom, upon which the animal stands, 
 there is arranged a sliding pan, having sides which slope at an angle of 
 about thirty degrees. An opening was provided at the lowest point 
 through which the urine might pass into a graduated receptacle hung 
 from the bottom. With such a cage the urine and feces could be collected 
 separately, the urine flowing into the receiver as soon as passed, where 
 it was preserved by means of thymol. The feces which were, in the feeding 
 periods, hardened by means of the ingested bone ash remained on the 
 wire netting and were picked out with a pair of forceps. The hair and 
 
FASTING STUDIES. 
 
 9 
 
 scurf fell through the netting to the pan and were brushed off each 
 day. At the end of each period the whole cage was carefully washed. 
 The cage washings consisted of scurf and small quantities of soft feces 
 which had fallen through the netting. 
 
 The urine was collected in 24-hour periods and kept in tincture-mouth 
 glass bottles, with ground glass stoppers. The dog was not catheterized 
 for it was found that when the animal was allowed to urinate at will 
 the bladder was satisfactorily emptied each day, or if a portion of the 
 dog’s urine was retained this was a fairly constant quantity. An ex- 
 amination of Table I, page 30, shows that the urine volumes were as a 
 rule very close to 250 cc. per day, corresponding to a daily water ingestion 
 of 250 cc. The other factors, such as the formation of water in metabolism 
 and loss through respiration, would seem to compensate each other. 
 Thymol and refrigeration 47 were used as a means of preservation. 
 
 The feces were collected when passed, dried, pulverized and the hair 
 separated, after which the feces were analyzed. Analyses were made 
 in periods. The hair, scurf, and cage washings were also analyzed in 
 periods and the nitrogen distributed evenly according to the number of 
 days in the period (see Table VI, p. 41). 
 
 The dog was weighed on a platform balance which was accurate to 0.0 1 
 kilogram. 
 
 Analyses and Methods . — Total nitrogen, urea, creatine, creatinine, 
 allantoin, and purine determinations were made upon the urine, and total 
 nitrogen determinations upon the feces, hair and cage washings. All 
 urine analyses were made in duplicate and upon the 24-hour sample, ex- 
 cept for a part of the intermediate feeding period, when analyses were 
 made on two-day composit samples and, in the case of allantoin and 
 purine, which were determined in composit samples of from two to ten 
 days. 
 
 The total nitrogen was determined by the Kjeldahl method, copper 
 sulfate 48 being used in the preliminary oxidation. 
 
 The urea was determined by the Benedict-Gephart 49 modification 
 of the Folin 50 method. While this method has recently 51 been shown 
 to be slightly inaccurate, due to the partial decomposition of uric acid, 
 creatine, creatinine, allantoin, and hippuric acid, still these substances 
 occur in such small amounts in the urine of the dog that the urea results 
 may be accepted as a fairly accurate indication of the amount of urea 
 present in the urine. Ammonia was determined by the method described 
 by Folin. 52 Creatinine was also determined by the Folin 32 procedure. 
 The creatine was hydrolyzed in the autoclave as suggested by Benedict 
 and Meyers. 53 Allantoin and purine were determined upon the same 
 composit sample by the method suggested by Poduschka, 54 and modi- 
 fied by Underhill and Kleiner. 34 
 
IO 
 
 FASTING STUDIES. 
 
 Blank determinations were made in all cases where possible impurities 
 in the reagents might cause an error. 
 
 EXPERIMENTAL. 
 
 Preliminary Feeding Period. 
 
 In an investigation of this character in which the nitrogen distribution 
 is the subject under consideration it is important that there shall be a 
 preliminary period. During this time the subject will live under the 
 same conditions as those which will exist during the experiment proper, 
 and will receive a constant diet. In our case, in addition to obtaining 
 uniform conditions of living and diet, this period served to put the animal 
 in a condition of nitrogen equilibrium, and also to furnish data for the 
 nitrogen partition of the urine of the animal under these normal conditions, 
 and thus serve as a basis of comparison for the fasting and for the in- 
 termediate feeding periods. 
 
 The preliminary feeding period in this experiment extended over fif- 
 teen days. At the end of this time, during which the animal had received 
 a uniform diet and constant water ingestion, a uniform body weight 
 had been secured and the dog was in nitrogen equilibrium. At this 
 point, the nitrogen balance showed an average of — 0.271 gram of nitrogen 
 per day for the eight days immediately preceding the fast. The nitro- 
 gen partition in the urine was determined for this period. The diet con- 
 sisted of 70 grams of meat, 15 grams of cracker dust, 8 grams of lard, 
 3 grams of bone ash, and 250 cc. of water. See Table VI, p. 41. 
 These ingredients were thoroughly mixed together and fed in the form 
 of a broth. This diet contained 2.732 grams of nitrogen, or about 0.8 
 gram per kilogram body weight. 
 
 The data of the preliminary period are found in Tables I and IV, pp. 30 
 and 36. From a consideration of the average data for this period it will 
 be seen that with an ingestion of 2.732 grams of nitrogen per day the dog ex- 
 creted 2.762 grams of nitrogen in the urine, and suffered a further loss of 
 0.235 gram of nitrogen in the form of feces, hair and cage washings. 
 This gives a minus balance of 0.271 gram of nitrogen per day. This 
 is an approximate nitrogen equilibrium (see Table IV, p. 36). 
 
 First Fast. 
 
 At the end of the preliminary period the dog was in a fair nitrogen 
 equilibrium, and had maintained a constant body weight for three days. 
 The last food was given at 2 p . m ., Feb. 4th, and the fast was considered 
 as beginning at 2 p . m ., Feb. 5th, the urine collected at 2 p . m . on Feb. 
 6th being considered as representing the urine of the first day of fast. 
 On the first day the water (250 cc.) was placed in the food pan, but only 
 50 cc. were taken. As a result the urine volume for the first day was 
 only 35 cc. and this was combined with the urine of the following day. When 
 
FASTING STUDIES. 
 
 1 1 
 
 the dog refused water on the second day of fast, the fluid was given by 
 means of a tube and this was continued throughout the fasts. On the 
 seventh day the first feces were passed, which were very small in amount 
 and of a pasty consistency. On the eleventh day the second fasting 
 feces were passed; these feces were removed from the cage immediately 
 after defecation and were greenish-brown in color, becoming black upon 
 exposure to air. It seems probable that the fresh fasting feces of the 
 dog are of a greenish-brown cast and only assume the black color ordi- 
 narily ascribed to them 55 upon coming in contact with the air. More 
 fasting feces were passed on the thirteenth day of fast. The dog con- 
 tinued weaker and could hardly stand. Feces were again passed on 
 the fourteenth day, but in small amount. 
 
 The lower nitrogen percentages for the feces of the feeding periods 
 above those for the fasting periods (see Table VI, p. 41) is probably 
 due primarily to the fact that bone ash was used in the diet of the feeding 
 period thus increasing the bulk of the fecal output although having no 
 influence upon its nitrogen content. At the same time it is no doubt 
 true that under some conditions at least, the percentage of nitrogen in 
 fasting feces is higher than that of the feeding feces from the same animal. 1 
 
 The following entry was made in the diary at the close of the fifteenth 
 day of fast. “February 20th. Soft feces in a.m. Dog could not stand 
 and was in an extremely weak condition. Pulse rose from 54, yesterday, 
 to 140 to-day, and the temperature sank from 37 0 to 36°. Weight 
 fell from 2.05 kg. to 1.85 kg. She has lost over 40 per cent, of her body 
 weight in two weeks. A very striking loss. Dog was so weak we feared 
 to go further, therefore, we fed her (4 p . m .) 15 grams cracker dust, 2 
 grams bone ash, and 3 grams lard. She took one-half of it. Gave her re- 
 mainder of it in the evening and covered her up with a blanket to keep her 
 warm. Pier nose was cold. She wagged her tail when addressed, could not 
 stand, however (6.10 p.m.). After dinner the dog rose as we entered and 
 walked to the front of the cage, wagging her tail. She was a new dog, and yet 
 she had partaken of only about 8 grams of cracker dust, 3 grams of lard, 
 and 2 grams of bone ash. We fed her the remainder of the amount of 
 food, previously mentioned, and then fixed her blanket for her in the 
 corner of the cage and she lay down contentedly. Later in the evening 
 the dog seemed much stronger.” 
 
 Discussion of the Nitrogen Distribution for the First Fast. Total Nitro- 
 gen . — From an examination of Table I, p. 30, and of figure on p. 12, it is 
 seen that in the first two days the nitrogen excretion fell from an average 
 of 2.768 grams during the preliminary period, to an average of 1.3 17 
 grams. This low nitrogen excretion is due, as has been proven by Bene- 
 dict 41 in fasts upon men, to the sparing action of glycogen present in the 
 1 Unpublished results from this laboratory. 
 
REPEATTD FAST • i DOGN s I 
 
FASTING STUDIES. 
 
 13 
 
 body. After the second day the nitrogen excretion in the urine rose rapidly 
 from day to day, taking a slight drop during the twelfth to fourteenth 
 days, and followed by a pronounced premortal rise, the nitrogen output 
 increasing from 2.472 grams on the fourteenth day to 3.944 grams on the 
 fifteenth day. 
 
 The rapidly increasing output of nitrogen mentioned above as occurring 
 during the first twelve days of the fast is somewhat unusual. It might 
 be inferred from such data, according to the commonly accepted theories 
 with regard to the nitrogen excretion in the urine of a fasting dog, that 
 the animal was thin and muscular, without any fat stores of any amount. 
 Or, on the other hand, it might be argued that the dog was organically 
 unsound. We believe both premises to be unfounded and have dis- 
 cussed the matter more at length on p. 22. 
 
 Urea . — The excretion of nitrogen as urea followed the total nitrogen 
 excretion, increasing absolutely, but not relatively. The percentage 
 urea nitrogen as compared with the total nitrogen was practically con- 
 stant throughout the fast, the average being 89.13 per cent, (see Table 
 III, p. 36) and the range being 86.36 per cent, on the sixth day to 91.12 
 per cent, on the seventh day (see Table II, p. 34). This average ex- 
 cretion is slightly higher than that found in the normal feeding period 
 (86.79 P er cent.). 
 
 This constant relation between urea and total nitrogen is contrary to 
 the statements in the literature, 33 » 34 > 35 to the effect that the percentage 
 of nitrogen as urea nitrogen decreases as the fast progresses. This fact 
 holds for men 23 ' 41 > 43 but it does not seem to hold for the dog, as is shown 
 in the experiments reported from our laboratory by Howe, Mattill and 
 Hawk. In their results the variations did not tend in any marked di- 
 rection away from a constant relative value. 
 
 The fact that the method of determining the urea-nitrogen is not 
 absolutely accurate might appear to invalidate our deductions. How- 
 ever, if the method determined all of the nitrogen appearing as creatine, 
 creatinine, etc., the percentage of nitrogen as urea would not be reduced 
 more than 3 to 4 per cent, and this would be a fairly constant quantity 
 throughout the experiment. Therefore, this error does not detract from 
 the accuracy of our conclusions. 
 
 Ammonia . — The ammonia-nitrogen fell from the average excretion 
 per day of 0.143 gram of nitrogen during the feeding period to an average 
 of 0.103 gram per day during the fast. After the first four days the am- 
 monia excretion remained almost constant, rising slightly at the end, 
 with the premortal rise. The relative excretion increased, then fell, 
 and finally increased again to the end. The range in the percentage of 
 ammonia nitrogen as compared to total nitrogen was from 3.64 per cent, 
 on the ninth day, to 6.24 per cent, on the third day, the average per cent. 
 
H 
 
 FASTING STUDIES. 
 
 of total nitrogen appearing as ammonia nitrogen being 4.59 per cent, 
 (see Table II, p. 34, and Table III, p. 36). 
 
 Creatinine . — The creatinine decreased in absolute amount very gradually 
 with but slight variations from day to day. The average daily excretion 
 for the fast (0.025 gram of nitrogen) was below that of the feeding period 
 (0.041 gram of nitrogen). This pronounced drop took place during the 
 first part of the fast and from then on the absolute amount of creatinine 
 excreted per day fell gradually, following in general the decreasing body 
 weight. While the absolute amount of creatinine nitrogen fell very grad- 
 ually, the percentage output decreased much more rapidly. The per- 
 centage of total nitrogen appearing as creatinine was 2.28 per cent, for 
 the first two days, and then fell very regularly, as the fast progressed, 
 to 0.46 per cent, on the last day. The grand average was 1.14 per cent, 
 as against an average of 1.49 per cent, for the normal feeding period 
 (see Table II, p. 34, and Table III, p. 36). 
 
 Creatine . — The creatine nitrogen fell from the feeding level (daily average 
 of 0.056 gram of nitrogen) to an average of 0.018 gram for the first two 
 days, and then remained approximately equal to the amount of crea- 
 tinine nitrogen excreted, being in general somewhat below this value. 
 On the ninth day the amount of creatine nitrogen excreted rose very 
 suddenly from 0.037 gram to 0.054 g ram > and continued on this higher 
 plane, with a tendency toward an increasing excretion, to the end of the 
 fast. The average excretion for these two portions of the fast were 
 0.025 gram of nitrogen for the first eight days and 0.068 gram of nitro- 
 gen for the last seven days, with a grand average of 0.046 gram of nitro- 
 gen. 
 
 The percentage of total nitrogen as creatine nitrogen increased from 
 1.40 per cent, on the first day to 3.10 per cent, on the twelfth day, due 
 to the greatly increased nitrogen excretion. The average percentage 
 of total nitrogen was 1.94 per cent, against 2.01 per cent, for the feeding 
 period (see Table II, p. 34, and Table III, p. 36). 
 
 Allantoin and Purine . 1 — The allantoin and purine nitrogen varied 
 from day to day. The average purine nitrogen excretion was below that 
 of the feeding period, being 0.005 gram per day, for eleven days of the 
 fast, against 0.012 gram of nitrogen during the feeding period. The 
 same average amount of allantoin nitrogen as of purine nitrogen was ex- 
 creted, i. e.y 0.005 gram while there was 0.008 gram of allantion nitrogen 
 excreted per day, during the feeding period. 
 
 The relative amounts of nitrogen excreted as purine and allantoin 
 nitrogen were 0.22 per cent, of purine, and 0.26 per cent, of allantoin 
 nitrogen, while the feeding period gave 0.45 per cent, purine, and 0.31 
 per cent, allantoin nitrogen (see Table II, p. 34, and Table III, p. 36). 
 
 1 These determinations were made by Mr. S. R. Wreath. 
 
FASTING STUDIES. 
 
 5 
 
 Undetermined Nitrogen . — The undetermined nitrogen fell as a result 
 of the fast, being 0.067 gram for an average during the fast, against 
 0.109 gram for the feeding period. 
 
 Intermediate Feeding Period. 
 
 General Considerations . — The object of this feeding period was to re- 
 store the animal to its original condition as regards body weight and 
 nitrogen equilibrium. The data upon the income and outgo of nitrogen 
 are found in Tables I and IV, pp. 30 and 36. During the portion of the 
 feeding period immediately following the end of the first fast the diet 
 was varied. The same constituents were fed, but the animal did not 
 receive the full diet given in the preliminary period. The diet for the 
 first three days did not contain any meat, and there was a minus balance 
 of 3 grams for this period. A positive balance of 1.218 grams occurred 
 on the fourth day, when meat was fed for the first time. During this 
 preliminary period of four days on an extremely low diet, the animal 
 gained 21.08 per cent, in body weight. The original diet was then fed, 
 and continued for fifteen days, during which period the body weight 
 increased but 7.57 per cent. In fact, during the last three days of this 
 period there was practically no increase in weight. 
 
 It appeared that to restore the original body weight, on the normal 
 diet, would require a very long time, 1 hence the diet was increased 50 
 per cent, thus causing the dog to ingest 4.104 grams of nitrogen per day. 
 Upon this diet the dog gained 20 per cent, of its weight in eight days. 
 The diet was again increased 50 per cent, at the end of the eight-day 
 period of increased feeding. The animal was now receiving twice as 
 much nitrogen per day (5.464 grams of nitrogen) as was required to main- 
 tain her constant body weight at the beginning of the experiment. 
 Throughout these periods of increased diet (50 per cent, and 100 per 
 cent.) the animal showed a plus nitrogen balance. After seventeen 
 days of the 100 per cent, increase the dog had more than regained the 
 weight lost during fasting, the weight being 3.59 kilograms, against a 
 weight of 3.41 kilograms at the beginning of the fast. The diet was now 
 reduced to the normal (2.732 grams of nitrogen per day). The weight 
 dropped slightly on the first day, and remained constant for the following 
 two days at 3.54 kilograms and at the same time the dog was in nitrogen 
 equilibrium, there being a plus balance of 0.02 gram of nitrogen per 
 day. 
 
 1 That it was possible to restore the normal body weight of fasting rabbits which 
 were fed the original diet following the breaking of the fast, has been shown by Kagan.® 
 However, in these tests the animals were not brought into a condition of nitrogen equi- 
 librium before being fasted, and the original diet may therefore have been an excessive 
 one, thus enabling Kagan to restore the normal weight of his rabbits with greater 
 facility than otherwise. 
 
i6 
 
 FASTING STUDIES. 
 
 In spite of the fact that the dog was giving a minus balance during the 
 first three days of the feeding period she gained 280 grams in weight. 
 This may have been due to the retention of water, for although the dog 
 had been receiving 250 cc. of water daily throughout the fast, upon feed- 
 ing the urine volumes for the first two days decreased to 215 and 50 cc. 
 respectively. While the method of collecting the urine (without catheteri- 
 zation) does not give absolute values for the urine volumes, the values 
 indicate that there must have been a marked water retention upon the 
 ingestion of food. This has been previously noted in feeding after ab- 
 solute fasts in dogs, 9 and also with men, 23 who had been receiving water 
 during the fast. 
 
 Nitrogen Distribution . — The average relative amount of urea nitrogen 
 excreted increased, as would be expected, with the increase in the diet, 
 and the amount of nitrogen excreted per day. The percentage nitrogen 
 output as urea dropped to the level of the preliminary period upon the 
 resumption of the normal diet. For further discussion see p. 23. 
 
 The absolute amount of ammonia nitrogen increased with the in- 
 crease in the diet. The relative amount of ammonia decreased, how- 
 ever, and was equal to the normal upon return to the original diet. 
 
 The creatinine nitrogen increased from day to day corresponding to 
 the increase in muscular tissue. The average amount of creatinine ex- 
 creted upon return to the original condition and body weight at the end 
 of the intermediate feeding period was practically equal to that excreted 
 under the normal conditions in force at the end of the preliminary feeding 
 period, being 0.040 gram of nitrogen per day, against 0.041 gram. Rela- 
 tively, however, the creatinine excretion decreased with the increase in 
 diet and the amount of total nitrogen excreted. On the same diet (nor- 
 mal) the creatinine nitrogen excretion was 1.49 per cent, of the total 
 nitrogen, during the preliminary feeding period, whereas, at the end of 
 the intermediate feeding period, the value was 1.63 per cent. In each 
 case the dog was in nitrogen equilibrium and of practically the same 
 body weight. 
 
 The amount of creatine nitrogen excreted increased with the increase 
 in the quantity of meat fed. 71 In other words the creatine ingested with 
 the food was in part excreted in the urine. 
 
 Allantoin and purine nitrogen were both excreted in larger amounts 
 during the fasting period. The average purine nitrogen excretion was 
 less (0.008 gram) than that in the preliminary feeding period (0.012 
 gram), whereas the allantoin nitrogen was the same, i. e., 0.008 gram. 
 
 Second Fast. 
 
 At the end of the feeding period, the weight of the animal was constant, 
 within 20 grams, for a period of six days. At the beginning of the second 
 fast the body weight was constant, although a little higher (3.54 kilograms) 
 
FASTING STUDIES. 
 
 17 
 
 than at the beginning of the first fast (3.41 kilograms). The nitrogen 
 balance was +0.02 gram on the day preceding the fast. The animal 
 appeared to be in a much better physical condition than when brought 
 to nitrogen equilibrium at the beginning of the first fast. The coat was 
 much smoother and the muscles were very firm. The activity and at- 
 tentiveness were fully as pronounced as at the beginning of the first 
 fast. 
 
 The last food was given at 2 p.m. on April 7th and the urine collected 
 on April 9th was taken as the first fasting urine. In this fast the water 
 (250 cc. per day) was given entirely by means of a tube. Signs of marked 
 weakness did not appear until about the fourteenth fasting day, and 
 even then the dog was quite active. From then on she became grad- 
 ually weaker, becoming more quiet and less disposed to move about. 
 She became perceptibly weaker on the twenty-ninth day and at 6.15 
 a.m. on the thirtieth day of the fast (16 hours after the beginning of 
 that day) the dog was found lying upon her side in a state of coma. Par- 
 ticulars with regard to the post-mortem examination and the histological 
 characteristics of the tissues will be published in a later article. 
 
 Total Nitrogen . — The analytical data for the second fast are contained 
 in Table I, p. 32, and the data are represented graphically in figure on 
 p. 12. From an excretion of 2.435 grams on the last day of the feeding 
 period, the total nitrogen excretion dropped on the first day of fast to 
 1. 08 1 grams. This was the smallest twenty-four-hour output of nitrogen 
 excreted at any time during the progress of the fast. The nitrogen excre- 
 tion remained fairly constant for the first twenty-six days. The maxi- 
 mum for this time was 1.471 grams on the sixth day (when no water was 
 given) with an average of 1.23 1 grams. From the twenty-sixth day on 
 there was an increased nitrogen excretion from day to day, until on the 
 thirtieth day an output of 2.298 grams was registered for a sixteen-hour 
 period. 
 
 Urea . — The urea nitrogen excretion followed the total nitrogen. Rela- 
 tive to the total nitrogen the average for the whole fast was 86.51 per 
 cent., the extreme values being 76.56 per cent, and 90.88 per cent, (see 
 Table II, p. 35, and Table III, p. 36). The lower value is probably 
 the result of alkaline fermentation, for the urine was found standing on 
 the pan of the cage, the wire gauze which served as a filter having become 
 clogged with hair. Aside from three days, the seventeenth, nineteenth, 
 and twenty-third days of the fast on which alkaline fermentation prob- 
 ably occurred, the variation was between 81.95 per cent, and 90.88 per 
 cent, with an average of 87.46 per cent., there being no marked tendency 
 toward either a drop or a rise in the percentage of total nitrogen as urea. 
 
 Ammonia . — The ammonia nitrogen showed a slight tendency to in- 
 crease, as the fast progressed, both absolutely and relatively. However, 
 
i8 
 
 FASTING STUDIES. 
 
 after the first six days, the excretion remained fairly constant, there 
 being but slight variations in either direction from the mean (0.083 gram 
 of nitrogen). This mean is a little high for it includes the three days 
 on which alkaline fermentation probably took place, and also the last 
 two days when the ammonia excretion was high. Excluding these, 
 the average was 0.068 gram of nitrogen. The variations noted range 
 from 0.039 gram to 0.085 gram of ammonia nitrogen. 
 
 Creatinine . — The creatinine nitrogen fell gradually from 0.032 gram on 
 the first day to 0.012 gram on the twenty-sixth day, with slight varia- 
 tions which may be attributed to the method of urine collection (without 
 catheterization). Unfortunately through accident we were prevented 
 from obtaining creatine and creatinine data on the urine for the last three 
 days of fast. The excretion of nitrogen as creatinine decreased, from day 
 to day, both absolutely and relatively. The maximum output was 
 2.96 per cent, on the first day, and the minimum 0.94 per cent, on the 
 twenty-seventh day, of fast, the average being 1.85 per cent, (see Table 
 II, p. 35, and Table III, p. 36). 
 
 Creatine . — The creatine nitrogen fell below that of the creatinine and 
 remained fairly constant until the fifteenth day, when it became greater 
 than the creatinine excretion, and remained above this value to the end of 
 the fast. On the last two days on which creatine was determined, there was 
 a tendency toward a rise in the creatine nitrogen excretion corresponding 
 to the premortal rise. The average creatine excretion was 0.026 gram 
 of nitrogen per day, the extremes being 0.008 gram on the fourteenth 
 day, and 0.053 gram on the twenty-seventh day of fast. Relative to 
 the total nitrogen the creatine nitrogen excreted was an average of 2.05 
 per cent, with extremes of 0.68 per cent, on the fourteenth day and 3.64 
 per cent, on the twenty-sixth day (see Table II, p. 35, and Table III, 
 P- 36). 
 
 Allantoin and Purine . — The allantoin and purine nitrogen did not show 
 any marked tendency toward a definit variation. The average excre- 
 tion was 0.005 gram of purine nitrogen per day, and 0.004 gram of allan- 
 toin nitrogen per day, which represents 0.38 per cent, of the total nitro- 
 gen as purine nitrogen, and 0.3 1 per cent, as allantoin nitrogen, respectively 
 (see Table III, p. 36). 
 
 Undetermined Nitrogen . — The undetermined nitrogen was low, and 
 practically constant in amount from day to day. It is interesting to 
 note that the average daily amount of undetermined nitrogen in the 
 second fast was only half that excreted during the first fast. 
 
 Discussion of the Two Fasts. 
 
 Data have been presented from two fasts upon a dog, which had pre- 
 viously never been subjected to an experiment of this character. The 
 first fast continued for a period of fifteen days, or until the dog was in 
 
FASTING STUDIES. 
 
 19 
 
 a state of collapse and the premortal rise was in evidence. This fast 
 was followed by an intermediate feeding period of forty-seven days, during 
 which time the animal was restored to its original condition. Following 
 this came the second fast, which continued for thirty days, or twice as 
 long as the first fast. 
 
 Some important factors which affect the length or intensity of a fast 
 are the race or breed, the age, the physical condition and activity, the 
 surrounding temperature, the character of the preliminary feeding period, 
 the general type of the nutritional and metabolic mechanism, and whether 
 the fast is absolute or with water. Since the two fasts were to be com- 
 pared, it was essential that the animal should be as nearly as possible 
 in the same condition as regarded body weight, nitrogen equilibrium, 
 etc., at the commencement of each fast, and, that the experimental 
 procedure throughout the fasts should be similarly regulated. The body 
 weight and nitrogen equilibrium were chosen as constants to be attained 
 before the beginning of the fast, whereas the water ingested was a constant 
 for the whole experiment. 2 57 
 
 In this experiment breed and age do not enter directly. While the 
 age of the animal affects the length of fast, in this case the dog was matured 
 and the time between the two fasts was so short (47 days) that the age 
 could not enter in any material way; Before each fast the dog was 
 brought to a constant body weight, and was practically in nitrogen equi- 
 librium. The body weight was a little greater at the beginning of the 
 second fast, being 3.54 kilograms, while at the beginning of the first 
 fast the weight was 3.41 kilograms. Whether this increase in weight 
 was fat or muscular tissue will be discussed later. A lower nitrogen 
 balance was attained than before the first fast, +0.02 gram of nitrogen 
 against — 0.271 gram before the first fast (see Table IV, p. 37). The 
 physical condition of the animal immediately before the fasts was prac- 
 tically the same in the two instances. However, her coat was smoother 
 and her muscles rather firmer at the commencement of the second fast. 
 
 Under the conditions outlined above, the dog experienced two fasts. 
 That these were profound is evidenced by the presence of the premortal 
 rise and the collapse of the animal in each case. The first fast was fifteen 
 days long while the second was twice as long. These are seemingly short 
 fasts, but considering the size and age of the dog they are of average 
 length. The greater length of the second fast is probably due to the 
 resistance of the tissues toward breakdown, acquired as a result of the 
 first fast. During the first fast the tissues seem to have broken down 
 very readily, so that the body lived improvidently upon its reserves. 
 In the second fast, however, the protein and fat needed for the main- 
 tenance of fife were used sparingly, and, as a consequence, the dog fasted 
 longer. This low rate of katabolism may have occurred in one of two 
 
20 
 
 FASTING STUDIES. 
 
 ways: the cells, as a result of the fast and the subsequent rebuilding 
 became more resistant, i. e., they gave up their protein constituents 
 less readily to the blood stream, on the other hand, the amount of pro- 
 tein or energy required for the functioning of the active organs, may 
 have been lessened and consequently the latter did not draw upon the 
 reserve protein as heavily as would have occurred had they not ex- 
 perienced a previous siege of fasting. 
 
 That the difference in the duration of the fasts was not a result of the 
 amount of fat stored up in the body can be shown from consideration of 
 the nitrogen data and body weights. The calculations on this basis 
 follow : 
 
 Weight before the first fast 3.41 kg. 
 
 Weight after the first fast 1.85 kg. 
 
 Percentage loss in weight 45 -75 % 
 
 Weight before second fast 3.54 kg. 
 
 Weight after second fast 1.91 kg. 
 
 Percentage loss in weight 46.04 % 
 
 Weight lost during first fast 
 
 Nitrogen lost during first fast, 35 . 65 gr. 
 
 Equivalent of nitrogen in terms of flesh (3.25 per cent. N) . . . . 
 
 Weight of fat and water lost (by difference) 
 
 Weight gained during feeding 
 
 Nitrogen stored during feeding, 38.25 gr. 
 
 Equivalent of nitrogen in terms of flesh (3.25 per cent. N) . . . . 
 
 Weight of fat and water stored (by difference) 
 
 Weight lost during second fast 
 
 Nitrogen lost during second fast, 41.15 gr. 
 
 Equivalent of nitrogen in terms of flesh (3.25 per cent. N) . . . . 
 Weight of fat and water lost (by difference) 
 
 1560 gr. 
 
 1100 gr. 
 460 gr. 
 1690 gr. 
 
 1180 gr. 
 5io gr. 
 1630 gr. 
 
 1260 gr. 
 370 gr. 
 
 When calculating all of the nitrogen lost as flesh, and the difference 
 between that and the loss in weight as fat, we see that during the feeding 
 period the body increased its muscular tissue by 80 grams, and its fat 
 by 50 grams, over the condition existing at the beginning of the first 
 fast. We can not say positively that that which has been called fat in 
 the above calculation is entirely fat, although the approximation is prob- 
 ably more accurate in the case of muscular tissue. However from a 
 consideration of these data we may conclude that the dog was apparently 
 in the same condition at the commencement of each fast. 
 
 The increased length of the second fast was not acccompanied by a 
 corresponding increase in the amount of muscular tissue and fat katabo- 
 lized. There was an increase of about 5.5 grams in the total weight 
 of nitrogen lost, which amount was lost during the last three days of the 
 fast. The calculations for the second fast show that only 370 grams 
 of fat were lost, while 510 grams were deposited during the intermediate 
 feeding period, leaving 140 grams of this deposited fat in the body at 
 
FASTING STUDIES. 
 
 21 
 
 the end of the fast. The fact that the post-mortem examination showed 
 that the animal was almost free from visible fat would contradict this 
 statement, unless it were in the form of combined fat (phosphatides) 
 as suggested by MacLean and Williams . 58 
 
 Had the increased length of fast and the decreased muscular katabolism 
 been influenced by more extensive fat deposits a much larger amount 
 of fat would have been used than can be accounted for. It seems there- 
 fore that the phenomena were due to the altered rate of the anabolic 
 and katabolic activities of the tissues and organs involved. 
 
 Albitsky 3 has shown that animals subjected to repeated fasting lose 
 body weight less rapidly, and further that the length of time necessary 
 to secure a definit body weight loss is greater than during the initial 
 fast. A long series of repeated fasts conducted at short intervals without 
 bringing the animal back to the normal condition between fasts may be 
 accompanied by serious results, for Richet showed, for example, that 
 dogs fasted repeatedly for five days at a time lived only six months, 
 and rabbits fasted at six-day intervals lived even a shorter time. 
 
 Schulz 12 presents a theory for the explanation of the phenomena of 
 decreased metabolism during interrupted feeding or repeated fasting. 
 He suggests that as a result of the removal of the stimulation 
 brought about by ingested food the activities of what is probably the 
 most important organ involved in metabolism, i. e., the liver, are impaired, 
 and consequently the protein metabolism is led into abnormal paths 
 and autointoxication may result. This phenomenon of autointoxica- 
 tion must be capable of being reduced upon the re-entrance of the stimu- 
 lation of food. With the instigation of a new fast the condition of increased 
 metabolism will not appear and as a result the injury resulting from the 
 loss of the stimulation due to food will be very small, or will be entirely 
 absent. 
 
 It is clear that such a theory takes no account of any metabolic activi- 
 ties other than those centered in the liver. While appreciating to the 
 full the extremely important role played by this organ in the metabolic 
 activities of the animal body, it nevertheless seems highly probable 
 that other organs are involved in connection with the decreased protein 
 katabolism of repeated fasting. 
 
 Schulz’s 12 theory, which would account for the collapse or the pre- 
 mortal rise as due to a defective liver function, is not substantiated in 
 the case of our dog, for it is evident that here the data would tend to 
 show that there was a certain minimum amount of protein which must 
 be present in the body in order that life shall exist. We believe this 
 conclusion to be warranted from the fact that our data show that during 
 each fast there were very similar amounts of nitrogen excreted notwith- 
 standing one fast was twice as long as the other. For example, during 
 
22 
 
 fasting studies. 
 
 the first fast of fifteen days the dog lost 10.45 grams of nitrogen per kilo- 
 gram of body weight, whereas, during the second fast of thirty dyes 
 she lost 11.60 grams of nitrogen per kilogram of body weight. In other 
 words, at the end of each fast there was evidently a similar residual quan- 
 tity of nitrogen-holding substance remaining in the body, the nitrogen of 
 which was not available for the metabolic uses of the organism. 
 
 Total Nitrogen . — The amount of nitrogen excreted in the urine in each 
 fast was approximately the same, i. e., 34.689 grams in the first fast and 
 39.430 grams in the second. The greater absolute amount of nitro- 
 gen excreted during the second fast was due, partly, to the increased 
 amount of nitrogen stored during the intermediate feeding period (80 
 grams of muscular tissue) and partly to the fact that the pre- 
 mortal rise of the first fast was incomplete. The average daily excretion 
 was 2.313 grams of nitrogen for the first and 1.3 13 grams of nitrogen for 
 the second fast. 
 
 During the first fast the total nitrogen excretion increased very rapidly 
 from day to day, denoting a most pronounced acceleration of the dis- 
 integration of muscular tissue. After a drop for three days, a very 
 marked premortal rise occurred on the last day (see Table I, p. 30, and 
 figure on p. 12), resulting in an excretion of 3.944 grams of nitrogen, which 
 was 1.472 grams of nitrogen jnore than on the previous day. This in- 
 crease was greater than the average daily excretion for the whole period. 
 
 The rapid loss of nitrogen during the first fast might be ascribed to one of 
 two things: first, a lack of fat in the body, or second, that the dog was 
 organically unsound. If our calculations hold good (p. 20) there was 
 more fat consumed during the first fast than during the second. The 
 increased fat deposit (due to feeding) amounted to but 50 grams of fat, 
 and was also accompanied by an increase in muscular tissue, and in 
 body weight. It is evident that there must have been a good store 
 of fat in the body before the first fast. This presumption was substan- 
 tiated by the feel of the skin, which denoted an ample subcutaneous 
 fat deposit. Furthermore, the complete recovery of the dog after the 
 first fast, and the fact that this was followed by a subsequent fast of 
 twice this length would denote an organically sound animal. The post- 
 mortem examination substantiated the above conclusion. 
 
 These are general deductions but sufficiently strong to warrant the 
 premise that the first fast was normal and the increased excretion of 
 nitrogen was due to a rapid normal katabolism of the muscular tissues. 
 This is further substantiated by the fact that more fat was consumed 
 during the first fast than in the second, notwithstanding the fact that 
 approximately equal amounts of nitrogen were excreted. 
 
 The rate of nitrogen excretion in the second fast was in marked con- 
 trast to that of the first fast. After the first day the excretion remained 
 
FASTING STUDIES. 
 
 23 
 
 almost constant for twenty-five days (an average of 1.238 grams) when 
 a gradual premortal rise set in, the excretion for the last sixteen hours 
 being 2.298 grams of nitrogen. The low constant excretion during the 
 second fast was that observed in most fasts, and accompanies the increased 
 length of time over which the amount of available nitrogen was metabo- 
 lized, since practically the same total amount of nitrogen was excreted in 
 each fast. 
 
 Urea . — The nitrogen excretion as urea followed the total nitrogen 
 excretion both absolutely and relatively. The average percentage of 
 urea nitrogen excreted was less in the second fast, 86.51 per cent., or, 
 excluding the three days upon which alkaline fermentation seemed to 
 have occurred, 87.46 per cent. In the case of the first fast the percentage 
 excretion was 89.13 per cent. These values are higher than the average 
 values for the normal diet. The fact that the ratio of urea nitrogen 
 to total nitrogen was constant has been discussed, see p. 13. This 
 constant ratio held for both fasts. 
 
 According to Folin’s 56 theory, the relative amount of urea excreted 
 varies with the amount of total nitrogen. This law was enunciated with 
 regard to normal metabolism. It has been shown in our laboratory 23 
 that with fasting men the percentage of urea nitrogen decreases, at 
 least for a time. Schondorf 35 and Underhill and Kleiner 34 hold this 
 to be true for the dog. The latter investigators report a fourteen-day 
 fast, the data from which vary considerably, and the fluctuations in the 
 percentage of urea nitrogen do not agree with those of the total nitrogen. 
 
 In fasting man the decrease in the percentage of urea is probably due 
 to the fatty acids, 60 which by combining with the ammonia prevent 
 the utilization of this ammonia in the formation of urea. The constant 
 relative urea excretion in our experiment would tend to show that fatty 
 acids or acid salts were probably not excreted in an increased amount 
 during the fasting period. 
 
 There is a slight increase in the percentage of urea nitrogen excreted 
 during the first fast, which would substantiate Folin’s theory with regard 
 to the effect of an increased nitrogen excretion. However, this point 
 is more strikingly brought out in the case of the intermediate feeding 
 period. An examination of Table III, p. 36, will show that the percentage 
 of total nitrogen in the form of urea increased from 82.67 per cent, to 
 86.51 per cent., then to 87.40 per cent, and finally to 88.74 P er cent, as 
 the diet w T as progressively increased. 
 
 It appears, therefore, that in a fasting dog the relative urea excretion 
 follows in general the laws of relative urea excretion which obtain for 
 normal feeding, for, with an increased nitrogen excretion, as in the first 
 fast, there was a slight increase in the relative urea-nitrogen excretion 
 and with a low and constant total nitrogen excretion in the second fast, 
 
24 
 
 FASTING STUDIES. 
 
 there was a lower and more constant relative urea excretion. Why this 
 should hold for the dog and not for man is not clear. It may be, perhaps, 
 that the dog, being a carnivorous animal, and living on the same form 
 of food in each case (fresh lean meat, and muscular tissue), excretes in 
 each instance the same percentage of total nitrogen in the form of urea. 
 Man, on the other hand, being unaccustomed to a diet of uncooked lean 
 meat, finds the conditions unusual when forced to exist upon his own 
 tissues during a fasting test, and consequently under these conditions 
 a different quota of the total nitrogen output is eliminated as urea than 
 when he ingests the normal mixed diet. 
 
 Ammonia . — The ammonia-nitrogen excretion was less for the daily 
 average in the second fast, 0.083 gram or, excluding the questionable 
 days already mentioned, 0.068 gram of nitrogen per day, as against 
 0.103 gram of nitrogen in the first fast. Instead of increasing from day 
 to day, as in the fasts of men, the ammonia remained fairly constant, 
 following the total nitrogen excretion, as Folin 56 has pointed out, only in 
 that when the total nitrogen excretion was low, the ammonia was abso- 
 lutely lower, but relatively higher. A rise in the ammonia excretion 
 at the end corresponded in each case with the premortal rise. The average 
 ratio of ammonia nitrogen to total nitrogen was in the first fast 4.59 
 per cent., and in the second fast 6.12 per cent., or, excluding the days 
 of alkaline fermentation, 5.66 per cent., which shows the relatively higher 
 ammonia excretion during the second fast. The facts noted here with 
 regard to the ammonia excretion are those which hold in normal feeding. 
 
 Creatinine . — The marked differences in the total nitrogen excretions 
 were not accompanied by similar changes in the creatinine-nitrogen out- 
 put. In both fasts the creatinine nitrogen fell gradually, the average 
 daily excretion for each period being almost the same, that for the first 
 fast being 0.025 gram of nitrogen and for the second 0.023 gram. The 
 total amount excreted in the second fast was nearly twice as great as 
 that of the first fast, the output of creatinine nitrogen, therefore, varying 
 inversely as the length of the fast. 
 
 Folin 56 first enunciated the law that the amount of creatinine excreted 
 in the urine by a normal individual is quite independent of either the amount 
 of protein in the food, or the total nitrogen in the urine. This has been 
 confirmed by van Hoogenhuyze and Verploegh, 40 Klerker, 61 Closson 62 
 and Shaffer, 63 whereas Cathcart 64 and Paton 65 fail to verify this theory. 
 Folin has further pointed out that the chief factor determining the amount 
 of creatinine eliminated appears to be the weight of the individual, and 
 that it also depends upon the mass of active protoplasmic tissue. Shaf- 
 fer 63 virtually substantiates this theory. The creatinine elimination 
 in fasting man has been observed by van Hoogenhuyze and Verploegh, 
 Benedict, 41 Benedict and Diefendorf 42 and Cathcart, 66 and in the fasting 
 
FASTING STUDIES. 
 
 25 
 
 dog by Underhill and Kleiner, 34 and Howe, Mattill and Hawk, 23 a gradual 
 decrease having been observed in each case. Underhill and Kleiner state 
 that the creatinine excretion is variable. Shaffer lays stress upon the 
 creatinine coefficient (milligrams per kilogram body weight) as a constant 
 for individuals of an equal muscular efficiency, which should then decrease 
 for a decreasing muscular efficiency. 
 
 The data from our experiment confirm, in the main, these theories. 
 With a decreasing body weight (loss of muscular tissue) and muscular 
 efficiency there is a decrease in the amount of creatinin excreted and also 
 in the creatinine coefficient (Table V, p. 38). The quantity of creatinine 
 excreted, per day, bears a direct relation to the mass of active muscular 
 tissue present in the body. The creatinine excreted in our experiment 
 does not bear any direct relation to the nitrogen excretion, for we have 
 a similar excretion of creatinine in the two fasts, notwithstanding the 
 fact that the rate of the nitrogen excretion was markedly different in the 
 two cases and that the average daily excretion of total nitrogen was nearly 
 100 per cent, greater in the first than in the second fast. 
 
 That it is the decrease in the amount of active protoplasm, and not 
 merely in the body weight, which causes a decrease in the creatinine excre- 
 tion may be seen from an examination of the data relative to the creatinine 
 coefficient. If the decrease were coincident with the decrease in weight 
 the creatinine coefficient would remain constant, whereas in reality it de- 
 creases more slowly than the body weight. This dependence upon the 
 mass of muscular tissue is shown in the intermediate feeding period. As 
 the dog gained in weight and stored nitrogen the quantity of creatinine 
 increased, although it did not bear any direct relation to the amount 
 of nitrogen excreted. On the nineteenth day, or at the time 34 per cent, 
 of the body weight lost in the fast, and one-fourth the nitrogen lost had 
 been restored, the creatinine coefficient was nearly equal to that before 
 the fast. Furthermore, on the twenty-fifth day or at the time the animal 
 had regained 50 per cent, of its lost nitrogen and body weight, the creatinine 
 coefficient was the same as that secured before the fast when the dog was 
 of constant body weight and in nitrogen equilibrium. If current ideas 
 regarding this coefficient are correct the findings indicate that the animal 
 
 possessed equal muscular efficiency under the two conditions. 
 
 The above may be substantiated by the following data : 
 
 Total amount of nitrogen lost during first fast 34.689 g. 
 
 Creatinine coefficient before fast =12 mg. 
 
 Nitrogen stored during 19 days of feeding (creatinine coefficient n mg.) 
 
 and 34 per cent, gain in body weight 9 .015 g. 
 
 Nitrogen stored during 25 days of feeding (creatinine coefficient 12 mg.) 
 
 and 50 per cent, in body weight 17.157 g. 
 
 We thus see that by the time the original creatinine coefficient was 
 restored, i. e., on the twenty-fifth day of feeding, the mass of muscle was 
 
26 
 
 FASTING STUDIES. 
 
 only 50 per cent, of that present before the fast. If we are to hold to the 
 idea that the ingested creatine (in the meat) and the amount of nitrogen 
 excreted have no effect upon the creatinine output, then in order that 
 the creatinine coefficient shall be significant, we must suppose a return 
 to the original condition as regards muscular efficiency when the animal 
 had only restored one-half of its nitrogen and body weight. The dog 
 certainly, at this point in the experiment, exhibited all of her normal 
 activity, which fact tends to confirm the above remarks with regard to 
 muscular efficiency. 
 
 Creatine. The excretion of creatine in the two fasts was quite similar, 
 
 being marked by a drop in the creatine-nitrogen excretion at the beginning 
 of the fast, followed by a period during which the creatine nitrogen re- 
 mained fairly constant, and a little less than the creatinine excretion. 
 Toward the end of the fast, and before the premortal rise there was a 
 sudden increase in the amount of creatine nitrogen excreted, being greater 
 than the creatinine excretion. This increased creatine continued to the 
 end. This marked increase of the creatine excretion over the amount 
 of creatinine excreted seems to be an indication of the pronounced metabolic 
 irregularities which finally result in the inception of the premortal rise, 
 and occurs from two to three days before the drop in the total nitrogen 
 excretion, which precedes this rise. 
 
 The amount of creatine excreted in the two fasts is almost equal, corre-^ 
 sponding to the amount of total nitrogen excreted during the periods 
 in which creatine was determined. Benedict, 67 Mellanby, 68 Shaffer, 63 
 Paton, 65 Fowler and Hawk 57 and others have furnished experimental 
 evidence in support of the theory that the creatine in the urine arises as 
 a result of the breakdown of the muscular tissues. Paton suggested 
 that the discrepancy between the amount of flesh which was katabolized 
 as calculated from the creatine output and that calculated from the total 
 nitrogen may be due to the resynthesis of a part of the nitrogen of the 
 broken-down muscular tissue, the creatine being excreted at the time of j 
 the muscle breakdown. 
 
 Calculating upon the basis of the excretions of total nitrogen and crea- 
 tine nitrogen we have derived in each case the weight of the mass of tissue 
 katabolized, assuming the accuracy of the theory that urinary creatine 
 represents disintegrated muscular tissue. The data are appended : 
 
 Flesh Lost as Calculated from the Urinary Total Nitrogen Data and from 
 the Creatine Nitrogen Data. 
 
 First Fast (15 days). 
 
 Nitrogen lost— 34.689 grams. 
 
 Equivalent in terms of flesh — (3.25%) = 1069 grams 
 Creatine nitrogen lost — 0.684 gram 
 Equivalent in terms of flesh (0.123%) = 555 grams 
 
FASTING STUDIES. 
 
 27 
 
 Second Fast (27 days). 
 
 Nitrogen lost — 33.524 grams 
 Equivalent in terms of flesh (3.25%) = 1030 grams 
 Creatine nitrogen lost — 0.689 gram 
 Equivalent in terms of flesh — (0.123%) = 560 grams 
 
 The amount of creatine nitrogen present in the muscles of the dog at 
 the end of the fast was 0.042 per cent., 70 showing a very marked decrease 
 (66 per cent.) in the amount of creatine in the fasting muscle remaining 
 in the body. Dorner has reported a somewhat smaller decrease in the case 
 of rabbits’ muscle. His data show the amount of creatine in this tissue 
 to be reduced from 0.456 per cent, to 0.356 per cent, as a result of fasting. 
 This pronounced decrease of creatine found by us in fasting muscle is 
 a most significant fact and shows clearly that in fasting we cannot with 
 accuracy consider the total amount of excreted creatine as resulting from 
 the complete and permanent disintegration of muscular tissue. How- 
 ever, even if we do consider all of the excreted creatine as having this 
 origin we have shown by calculation that we can account for only about 
 one-half of the total nitrogen on this basis. The discrepancy between 
 the mass of muscular tissue lost, calculated on the two bases, i. e., total 
 nitrogen and creatine nitrogen output becomes all the more striking 
 when we appreciate that a large part of the creatine excreted during the 
 fasts and which is ordinarily considered as representing completely dis- 
 integrated muscular tissue, in reality most certainly does not represent 
 this but rather has been withdrawn from muscular tissue which is still 
 functioning as living tissue within the body of the animal. Unfortu- 
 nately no determination was made of the actual mass of muscular tissue 
 remaining in the body after fasting. However, inasmuch as we have 
 shown in this laboratory 70 that the total nitrogen content of fasting 
 muscle is only slightly lower than that of well-nourished muscle it is evi- 
 dent that but a small fraction of the total nitrogen excreted by the urine 
 arose from the muscular tissue still functioning within the body, whereas 
 a comparatively large part of the creatine had this origin. When all 
 these facts are taken into consideration it is apparent that over 50 per 
 cent, of the total nitrogen of the urine had a source other than the katabo- 
 lism of muscular tissue. The fact that fasting muscle may have its 
 creatine store depleted to one-third the normal while retaining its nitro- 
 gen content but slightly lowered indicates rather conclusively that the 
 creatine is held in such a combination within the muscle that its release 
 is not necessarily accompanied by the disintegration of the other nitroge- 
 nous muscle constituents, which go to form the total nitrogen of muscular 
 tissue. 68 ’ 57 At the same time we must not lose sight of the fact that the 
 muscular tissue whose nitrogen store is seemingly intact at the end of 
 a fast may assume this condition not because the katabolic processes 
 
28 
 
 fasting studies. 
 
 have not been sufficiently profound to alter its inner structure but rather 
 because of certain synthetic reactions which have followed or accompanied 
 the primary disintegration of the tissues. 65 That there is a direct re1 ^ 
 tion between the amount of nitrogenous material katabolized and the 
 creatine excreted is brought out in our results. For example, with prac- 
 tically the same amount of nitrogen excreted under the same conditions 
 i. e. two fasts continuing to the premortal rise, and with two equal 
 but decidedly different rates of nitrogen excretion, practically the same 
 
 amount of creatine was excreted. . 
 
 No direct evidence has been obtained which would show the relation 
 between creatine and creatinine. The creatine appears to be derived either 
 from disintegrating muscular tissue or to be removed in s °“ e “r 
 from such tissues which are still functioning within the body. The 
 creatine excretion was low during the first part of each fast, increasing 
 suddenly a few days before the drop in the total nitrogen excretion wh ch 
 preceded the premortal rise. From this time to the end of the fast the 
 daily output of creatine nitrogen exceeded that of the creatinine nitrogen. 
 Dorner 69 in the case of a rabbit which died after nine days of fasting, 
 found the creatine excretion of the second day to exceed that of creatimne. 
 
 Summary. 
 
 A fox terrier bitch about one year old and weighing 3-4 1 kilograms 
 was brought into nitrogen equilibrium and was subjected to two fasts. 
 On the fifteenth day of the first fast the premortal rise m nitrogen ^out- 
 put was noted and was accompanied by other signs indicating that death 
 would result in a few hours. She was then carefully fed and dunng 
 the feeding period of forty-seven days regained her former weight and 
 was again brought into nitrogen equilibrium, after which she was fasted 
 Tsecond time for a period of thirty days. The animal was in fully as 
 good physical condition at the beginning of the second as she wa 
 
 at the commencement of the first fast. .. 
 
 The water ingestion was uniform (250 «.) throughout the feeding 
 and fasting periods. The loss in body weight was nearly equal in each 
 fast being 45.75 per cent, for the first and 46.04 per cent, for the 
 second fast. However, on the fifteenth day of the second fast ’ t ' e '’ 
 after a period equaling in length the entire first fast, the animal had 
 lost but 25.42 per cent, of her body weight. 
 
 From the data obtained we have shown that: . 
 
 1 With a practically equal total nitrogen excretion, the rate of t 
 excretion was widely different in the two fasts, being high and rapidy 
 increasing during the first fast of fifteen days, and low and fairly con- 
 stant during the second fast of thirty days. A premortal rise occurred 
 
 at the end of each fast. . « 
 
 2. The absolute amount of urea nitrogen excreted was in direct re a- 
 
FASTING STUDIES. 
 
 29 
 
 tion to the total nitrogen excretion. The relative urea-nitrogen ex- 
 cretion remained practically constant throughout the fasts, but increased 
 with an increased total nitrogen excretion during the intermediate 
 feeding period. 
 
 3. The ammonia-nitrogen excretion remained fairly constant, varying 
 with the total nitrogen excretion in that when this was low the ammonia- 
 nitrogen was absolutely lower but relatively higher. 
 
 4. The creatinine-nitrogen excretion decreased very uniformly and 
 gradually as the fasts progressed, and increased uniformly and gradually 
 during the intermediate feeding period. The total output of creatinine 
 nitrogen varied inversely as the length of the fast. On the nineteenth 
 day of the feeding period, or at the time thirty-four per cent, of the body 
 weight lost in the first fast, and one-fourth the nitrogen lost, had been 
 restored, the creatinine coefficient was nearly equal to that before the 
 fast. Furthermore, on the twenty-fifth day or at the time the animal 
 had regained only fifty per cent, of its lost nitrogen and body weight, 
 the creatinine coefficient was the same as that secured before the fast when 
 the dog was of constant body weight and in nitrogen equilibrium. 
 
 5. The total amount of creatine nitrogen excreted was practically the 
 >ame during each fast. The average daily amount of creatine nitrogen, 
 fowever, was nearly twice as great during the first fast as during the 
 second fast. 
 
 6. The allantoin and purine nitrogen excretions decreased as a result 
 )f the fast and were practically equal for each fast. 
 
 7. The undetermined nitrogen decreased as a result of fasting. The 
 iverage daily amount of undetermined nitrogen in the second was 
 me-half that in the first fast. 
 
 8. A consideration of the summation of the nitrogen balances shows 
 hat there is a minimum amount of nitrogen which must be present in 
 he body in order that life shall exist. 
 
 9. The excretion of urinary creatine increased suddenly a few days 
 >efore the drop in the total nitrogen excretion which precedes the pre- 
 tiortal rise. From this time to the end of the fast the daily output of 
 reatine nitrogen exceeded that of the creatinine nitrogen. 
 
 10. Assuming the accuracy of the theory that the urinary creatine 
 epresents disintegrated muscular tissue and calculating accordingly, 
 
 • discrepancy exists between the calculated mass of muscular tissue 
 3st, when considered from the standpoint of total nitrogen and creatine 
 itrogen respectively. When all the facts in this connection are taken 
 ito consideration it is apparent that over fifty per cent, of the total 
 itrogen had a source other than the muscular tissue. 
 
 11. The creatine content of muscle showed a marked decrease (over 
 ixty per cent.) as a result of fasting, while the nitrogen content of similar 
 
Tabus I. — General Data. 
 
 FASTING STUDIES. 
 
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 944 3-544 
 
FASTING STUDIES. 
 
 31 
 
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Table I ( continued ). 
 
 3 2 
 
 FASTING STUDIES. 
 
 a 
 
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 33 
 
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34 
 
 Day. 
 
 I 
 
 2 
 
 3 
 
 4 
 
 5 
 
 6 
 
 7 
 
 8 
 
 ;! 
 
 3 
 
 4 
 
 5 
 
 6 
 
 7 
 
 8 
 
 9 
 
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 ii 
 
 12 
 
 13 
 
 14 
 
 15 
 
 2-3 
 
 4 
 
 5 
 
 6 
 
 7 
 
 8 
 
 9 
 
 io 
 
 ii 
 
 12 
 
 13 
 
 14 
 
 15-16 
 
 17-18 
 
 19 
 
 20 
 
 21-22 
 
 FASTING STUDIES. 
 
 Table II. — Percentage Nitrogen Distribution. 
 
 Urea N. Ammonia N. Creatinine N. Creatine N. Purine N. Allantoin N. Undeter N. 
 
 Preliminary Feeding Period. 
 
 86.96 
 
 384 
 
 1.88 
 
 2-95 
 
 
 
 0.70 
 
 88.10 
 
 4.18 
 
 1.68 
 
 3-59 
 
 
 0.09 
 
 i -45 
 
 86.65 
 
 4.64 
 
 i -52 
 
 1.25 
 
 
 0 . 1 1 
 
 5.82 
 
 86.82 
 
 4-57 
 
 1.47 
 
 2 .01 
 
 0.58 
 
 0.16 
 
 4-38 
 
 84.85 
 
 5.67 
 
 1. 41 
 
 0.51 
 
 0.24 
 
 0.90 
 
 6.42 
 
 85.15 
 
 6.92 
 
 i -45 
 
 2-49 
 
 0-54 
 
 0-34 
 
 3-io 
 
 89-93 
 
 4.24 
 
 1 .08 
 
 3.28 
 
 
 0.24 
 
 1.25 
 
 85.84 
 
 7.29 
 
 i -45 
 
 First 
 
 2 .62 
 
 Fast. 
 
 
 
 2.79 
 
 8751 
 
 4.48 
 
 2.28 
 
 1.40 
 
 0.23 
 
 0.49 
 
 3.60 
 
 86.84 
 
 6.24 
 
 i -95 
 
 1. 41 
 
 
 
 3-56 
 
 89-54 
 
 4.44 
 
 1.63 
 
 1.63 
 
 
 
 2-75 
 
 89.78 
 
 5.87 
 
 1 .60 
 
 1 .02 
 
 0.27 
 
 0.18 
 
 1 .29 
 
 86.36 
 
 6.07 
 
 1.23 
 
 1 .61 
 
 0.32 
 
 0.21 
 
 4.08 
 
 91 . 12 
 
 3-99 
 
 113 
 
 1 .02 
 
 
 
 2.74 
 
 88.26 
 
 5 -ii 
 
 M3 
 
 1 .60 
 
 0.04 
 
 0.04 
 
 3.8i 
 
 90.97 
 
 3.64 
 
 0.98 
 
 2.05 
 
 0.23 
 
 
 2 .12 
 
 90.00 
 
 3-67 
 
 0-93 
 
 2.23 
 
 0.22 
 
 0.07 
 
 2.88 
 
 89.84 
 
 3-78 
 
 0.86 
 
 2.70 
 
 0-43 
 
 0.14 
 
 2.23 
 
 88.92 
 
 4-39 
 
 0.76 
 
 3. 10 
 
 0.04 
 
 0.08 
 
 2.70 
 
 89.19 
 
 4.14 
 
 0-59 
 
 2 .80 
 
 0.16 
 
 O.16 
 
 2 .96 
 
 89.56 
 
 4.45 
 
 0.48 
 
 2-79 
 
 0.28 
 
 0.28 
 
 2.14 
 
 89.86 
 
 4.06 
 
 0.46 1.77 
 
 Feeding Periods. 
 Preliminary. 
 
 
 0.30 
 
 3.60 
 
 91 .01 
 
 3-59 
 
 0-34 
 
 1-52 
 
 
 
 4.01 
 
 82.45 
 
 7 .00 
 
 1.50 
 
 2 <~9 
 
 0.42 
 
 0.48 
 
 5-15 
 
 74-53 
 
 1105 
 
 1.63 2.56 
 
 Normal Diet. 
 
 0.46 
 
 0-93 
 
 8.84 
 
 85-55 
 
 5.82 
 
 0.96 
 
 2.65 
 
 0.44 
 
 0.07 
 
 4-50 
 
 81.30 
 
 11.46 
 
 0.92 
 
 2-34 
 
 0-33 
 
 0.05 
 
 3-59 
 
 85-37 
 
 6-93 
 
 0-93 
 
 0.87 
 
 0-35 
 
 0.06 
 
 5 48 
 
 86.58 
 
 6-33 
 
 113 
 
 2-57 
 
 0.38 
 
 0.06 
 
 2-95 
 
 89.27 
 
 4-95 
 
 1. 14 
 
 3 05 
 
 0.38 
 
 0.06 
 
 1 14 
 
 87.14 
 
 4-74 
 
 1. 14 
 
 3-54 
 
 o.33 
 
 
 3” 
 
 86.99 
 
 5.23 
 
 1 .04 
 
 316 
 
 0.31 
 
 
 327 
 
 87.31 
 
 5.26 
 
 0.94 
 
 2.47 
 
 0.32 
 
 
 3-68 
 
 87.96 
 
 512? 
 
 0.78 
 
 1 .84 
 
 0.31 
 
 
 384 
 
 87.06 
 
 5 08) 
 
 2.31 
 
 0.40 
 
 0-59 
 
 3-37 
 
 85.62 
 
 6.61 
 
 1 . 10 
 
 2.91 
 
 0.27 
 
 0-43 
 
 3 04 
 
 88.27 
 
 5-24 
 
 1 . 12 
 
 2 . 11 
 
 0.36 
 
 0.64 
 
 2 .26 
 
 86.21 
 
 6.90 
 
 1.38 2.86 
 
 50% Increase in Diet. 
 
 0-34 
 
 0-59 
 
 1.82 
 
 86.21 
 
 6.90 
 
 1 .00 
 
 3-04 
 
 0.27 
 
 0.46 
 
 2.12 
 
 87.23 
 
 5.19 
 
 0.94 
 
 3-44 
 
 0.23 
 
 0.68 
 
 2-43 
 
FASTING STUDIES. 
 
 35 
 
 Table II ( continued ). 
 
 Day. 
 
 Urea N. 
 
 Ammonia N. 
 
 Creatinine N. Creatine N. 
 50% Increase in Diet. 
 
 Purine N. 
 
 Allantoin N. Undeter. : 
 
 23-24 
 
 87-59 
 
 5 • 11 
 
 1 .01 
 
 3-67 
 
 O.24 
 
 0.69 
 
 1 .61 
 
 25-26 
 
 87-73 
 
 5 - 3 2 
 
 1. 17 
 
 4-23 
 
 O.27 
 
 O.44 
 
 0.77 
 
 27 
 
 88.25 
 
 4.14 
 
 1.32 4.52 
 
 100% Increase in Diet. 
 
 0-34 
 
 O.23 
 
 1 .21 
 
 28 
 
 88.25 
 
 4-13 
 
 0-95 
 
 3.26 
 
 O.24 
 
 O.16 
 
 2.99 
 
 29-30 
 
 89.21 
 
 4-38 
 
 0.86 
 
 3-13 
 
 
 0-37 
 
 2.05 
 
 31-32 
 
 87.56 
 
 4.46 
 
 o -99 
 
 3-69 
 
 
 O.27 
 
 3-02 
 
 33-34 
 
 87.62 
 
 4-73 
 
 0.88 
 
 2 .81 
 
 
 O.17 
 
 3.78 
 
 35-36 
 
 86.51 
 
 4-97 
 
 1 .00 
 
 3-72 
 
 0.28 
 
 0.21 
 
 3.05 
 
 37-38 
 
 88.02 
 
 4.17 
 
 1 .00 
 
 3-39 
 
 0-37 
 
 0.05 
 
 2.97 
 
 39-40 
 
 87.83 
 
 4.67 
 
 o -99 
 
 3-33 
 
 0-39 
 
 0.05 
 
 2 .69 
 
 41-42 
 
 87.82 
 
 4 05 
 
 1 .01 
 
 2.94 
 
 O.36 
 
 0.05 
 
 3 - 7 i 
 
 43-44 
 
 87.87 
 
 4.08 
 
 1.03 3-47 
 
 Normal Diet. 
 
 0-37 
 
 0.05 
 
 3 05 
 
 45-46 
 
 87.19 
 
 4-63 
 
 1.67 
 
 3.46 
 
 0-33 
 
 0.20 
 
 2 .72 
 
 47 
 
 86.44 
 
 4-56 
 
 1.60 1.89 
 
 Second Fast. 
 
 0-33 
 
 0.21 
 
 4-97 
 
 1 
 
 87.05 
 
 4-25 
 
 2.96 
 
 2.68 
 
 
 0.28 
 
 2.78 
 
 2 
 
 86.54 
 
 4.84 
 
 2 .67 
 
 2-59 
 
 
 O.25 
 
 3 09 
 
 ■ 3 
 
 87-51 
 
 4-30 
 
 2 .48 
 
 1 .16 
 
 0-33 
 
 O.4I 
 
 3- 80 
 
 | 4 
 
 88.10 
 
 3-25 
 
 2 .82 
 
 2-33 
 
 0-33 
 
 O.42 
 
 2-74 
 
 5 
 
 86.81 
 
 4-95 
 
 2 .48 
 
 2.56 
 
 O.32 
 
 O.4O 
 
 2 .48 
 
 6 
 
 89-53 
 
 3-46 
 
 2.31 
 
 1.77 
 
 O.27 
 
 o-34 
 
 2.31 
 
 7 
 
 87.32 
 
 5 05 
 
 2-39 
 
 1 . 60 ... 
 
 O.36 
 
 0.44 
 
 2 .84 
 
 8 
 
 88.38 
 
 5-12 
 
 1 .90 
 
 i -75 
 
 O.29 
 
 0.36 
 
 2 . 19 
 
 9 
 
 N 
 
 00 
 
 6.07 
 
 2.05 
 
 1 -3 1 
 
 0-33 
 
 0.41 
 
 2.38 
 
 10 
 
 88 . 44 
 
 6.28 
 
 2.18 
 
 1 .01 
 
 0-34 
 
 0.42 
 
 i -34 
 
 11 
 
 88.60 
 
 6.03 
 
 2 .07 
 
 1 .21 
 
 0-34 
 
 o -43 
 
 1 .29 
 
 12 
 
 88.06 
 
 5-85 
 
 2 .02 
 
 i -43 
 
 0.30 
 
 o -37 
 
 i -95 
 
 13 
 
 87.84 
 
 6.26 
 
 1 .84 
 
 1 .03 
 
 
 0.30 
 
 2-73 
 
 14 
 
 86.97 
 
 7 .02 
 
 2.03 
 
 0.68 
 
 
 0.34 
 
 2 .96 
 
 15 
 
 90.88 
 
 6.96 
 
 1 .90 
 
 1 .32 
 
 
 0-33 
 
 
 16 
 
 88.02 
 
 4.91 
 
 1.49 
 
 1 - 7 1 
 
 
 0.30 
 
 
 17 
 
 78.54 
 
 14.92 
 
 1 .61 
 
 1 .61 
 
 
 0.36 
 
 2-95 
 
 18 
 
 85.92 
 
 7.12 
 
 1.76 
 
 2 . 10 
 
 
 0-34 
 
 2-77 
 
 19 
 
 Si -95 
 
 10.30 
 
 1.50 
 
 2 .62 
 
 0-37 
 
 0.15 
 
 3 07 
 
 20 
 
 88.07 
 
 4-93 
 
 i -43 
 
 2.86 
 
 O.40 
 
 0. 16 
 
 2.15 
 
 21 
 
 86.49 
 
 5-49 
 
 1 - 3 1 
 
 2.87 
 
 O.41 
 
 0.16 
 
 3-2 
 
 22 
 
 76.56 
 
 6.32 
 
 1 . 16 
 
 3-16 
 
 O.42 
 
 1 • l 7 
 
 12.307 
 
 23 
 
 82 .05 
 
 10.57 
 
 0.99 
 
 2-51 
 
 O.46 
 
 0.23 
 
 3-19 
 
 24 
 
 88.08 
 
 6.74 
 
 0.98 
 
 1.97 
 
 O.49 
 
 0.25 
 
 1 .48 
 
 25 
 
 87-59 
 
 6.46 
 
 1.05 
 
 2.36 
 
 O.52 
 
 0.26 
 
 1-75 
 
 26 
 
 86.23 
 
 5-33 
 
 1 .06 
 
 3-64 
 
 0-53 
 
 0.27 
 
 2-93 
 
 27 
 
 88.16 
 
 4-35 
 
 0-94 
 
 3-54 
 
 O.41 
 
 0.20 
 
 2.41 
 
 28 
 
 87.49 
 
 5-46 
 
 
 
 
 
 
 29 
 
 87.41 
 
 7-33 
 
 
 
 
 
 
 30 
 
 88.59 
 
 7.92 
 
 
 
 
 
 
36 
 
 FASTING STUDIES. 
 
 Table III. — Summary of Nitrogen Excretion. 
 
 
 Total 
 
 Urea Ammonia 
 
 Crea- 1 
 
 Creatine 
 
 Purine Allantoin Unde- 
 
 Period (diet). 
 
 N. 
 
 N. N. 
 
 tinine N. 
 
 N. 
 
 N. 
 
 N. ter. N. 
 
 
 
 Period Averages — Grams per Day. 
 
 
 
 Preliminary . 
 
 . .2 .768 
 
 2.405 0.143 
 
 O.041 
 
 O.056 
 
 0.012 
 
 0.008 0.109 
 
 First Fast . . 
 
 . . . 2.313 
 
 2.065 0.103 
 
 O.025 
 
 O.O46 
 
 0.005 
 
 0.005 0.067 
 
 Prelim. Feed. 
 
 .. . 1.356 
 
 1.162 0.105 
 
 0.012 
 
 O.O29 
 
 
 0.070 
 
 Normal 
 
 .. .I.833 
 
 1.587 0.112 
 
 0.020 
 
 O.054 
 
 
 0.076 
 
 50 % 
 
 . . .2.689 
 
 2.351 0.142 
 
 0.028 
 
 O. 102 
 
 - 0.008 
 
 O . 008 O . 045 
 
 100% 
 
 . . .4071 
 
 3.576 0.178 
 
 O.O4O 
 
 0.134 
 
 
 O.124 
 
 Normal 
 
 
 2.132 0.113 
 
 O.O4O 
 
 0.072 _ 
 
 
 0.085. 
 
 Second Fast . 
 
 . • . 1.314 
 
 1.141 0.083 
 
 0.023 
 
 0.026 
 
 0.005 
 
 O.OO4 O.032 
 
 
 
 Period Averages — Per < 
 
 cent. 
 
 
 
 Preliminary . 
 
 . . ... 
 
 86.79 5-17 
 
 I.49 
 
 2 .01 
 
 0-45 
 
 0.31 4-03 
 
 First Fast. . . 
 
 
 89 13 4-59 
 
 I. 14 
 
 1.94 
 
 0.22 
 
 0.26 2.89 
 
 Prelim. Feed. 
 
 
 82.67 7.21 
 
 I . II 
 
 2.36 
 
 0-45 
 
 0.71 6.00 
 
 Normal 
 
 
 86.51 6.13 
 
 1.05 
 
 2.51 
 
 0-35 
 
 0.28 3.25 
 
 50 % 
 
 
 87-40 5-33 
 
 1 .09 
 
 3.78 
 
 O.27 
 
 0.50 1.63 
 
 100% 
 
 
 88.74 4.40 
 
 0.97 
 
 3.30 
 
 O.4O 
 
 0.15 3 03 
 
 Normal 
 
 
 86.81 4.59 
 
 I.63 
 
 2.68 
 
 0-33 
 
 0.20 3.56 
 
 Second Fast . 
 
 
 86.51* 6.12f 
 
 1.85 
 
 2.05 
 
 O.38 
 
 0.31 2.58 
 
 
 Total Weight of Nitrogen Excreted in Urine (Grams). 
 
 
 First Fast. . . 
 
 . .34-689 
 
 30.972 1.550 
 
 0-373 
 
 0.684 
 
 
 
 Second Fast . 
 
 . - 39-430 
 
 34.224 2.483 
 
 0.6ll 
 
 0.689 
 
 
 
 
 
 Table IV. — Nitrogen Balances. 
 
 
 
 
 N in hair. 
 
 Total N 
 
 
 
 
 
 N in urine. 
 
 feces and cage 
 
 output. 
 
 N in food. 
 
 
 Day. 
 
 Grams. 
 
 wash. Gram. 
 
 Grams. 
 
 
 Grams. 
 
 N balance. 
 
 
 
 Preliminary Period. 
 
 
 
 
 1 
 
 2.707 
 
 0.235 
 
 2.942 
 
 
 2.732 
 
 0.210 
 
 2 
 
 2.202 
 
 0.235 
 
 2-437 
 
 
 2.732 
 
 + O . 295 
 
 3 
 
 2.63O 
 
 0.235 
 
 2.865 
 
 
 2.732 
 
 — 0.133 
 
 4 
 
 2.580 
 
 0.235 
 
 2.815 
 
 
 2.732 
 
 O.083 
 
 5 
 
 2-555 
 
 0.235 
 
 2.790 
 
 
 2.732 
 
 — O.O58 
 
 6 
 
 2.964 
 
 0.235 
 
 3-199 
 
 
 2.732 
 
 — O.467 
 
 7 
 
 3-683 
 
 0.235 
 
 3.918 
 
 
 2.732 
 
 I . l86 
 
 8 
 
 2.825 
 
 0.235 
 
 3.060 
 
 
 2.732 
 
 — O.328 
 
 
 
 First Fast. 
 
 
 
 
 
 2.634 
 
 0.064 
 
 2.762 
 
 
 
 2 . 762 
 
 3 
 
 I.489 
 
 0.064 
 
 1-553 
 
 
 
 — 1-553 
 
 4 
 
 I .960 
 
 0.064 
 
 2 .024 
 
 
 
 — 2 .024 
 
 5 
 
 2.250 
 
 0.064 
 
 2.314 
 
 
 
 — 2.314 
 
 6 
 
 1.862 
 
 0.064 
 
 1 .926 
 
 
 
 — 1 .926 
 
 7 
 
 2.558 
 
 0.064 
 
 2.622 
 
 
 
 — 2 .622 
 
 8 
 
 2.308 
 
 0.064 
 
 2.372 
 
 
 
 — 2.372 
 
 9 
 
 2.637 
 
 0.064 
 
 2 .701 
 
 
 
 — 2.701 
 
 io 
 
 2 .780 
 
 0.064 
 
 2.844 
 
 
 
 — 2.844 
 
 * See discussion, p. 17. 
 
 -j- See discussion, pp. 17-18. 
 
FASTING STUDIES. 
 
 37 
 
 Table IV (< continued 
 
 
 
 N in hair, 
 
 Total N 
 
 
 
 
 N in urine. 
 
 feces and cage 
 
 output. 
 
 N in food. 
 
 
 Day. 
 
 Grams. 
 
 wash. Gram. 
 
 Grams. 
 
 Grams. 
 
 N balance 
 
 
 
 
 First Fast. 
 
 
 
 II 
 
 2.777 
 
 0.064 
 
 2 .841 
 
 
 — 2 .841 
 
 12 
 
 2.483 
 
 0.064 
 
 2-547 
 
 
 — 2-547 
 
 13 
 
 2-535 
 
 0.064 
 
 2-599 
 
 
 — 2.599 
 
 14 
 
 2.472 
 
 0.064 
 
 2.536 
 
 
 — 2.536 
 
 15 
 
 3-944 
 
 0.064 
 
 4.008 
 
 
 — 4 . 008 
 
 
 
 Feeding Periods. 
 
 
 
 
 
 Preliminary. 
 
 
 
 I 
 
 2.893 
 
 0. 161 
 
 3-054 
 
 0.198 
 
 — 2.856 
 
 3 
 
 1 .670 
 
 0. 161 
 
 1.992 
 
 I .848 
 
 — O.144 
 
 4 
 
 0.860 
 
 0. 161 
 
 1 .021 
 
 2.239 
 
 + I. 218 
 
 
 
 Normal Diet. 
 
 
 
 5 
 
 1-356 
 
 0. 161 
 
 I- 5 I 7 
 
 2.465 
 
 + O . 948 
 
 6 
 
 1 .840 
 
 0. 161 
 
 2 .001 
 
 2.732 
 
 + O.731 
 
 7 
 
 1.716 
 
 0. 161 
 
 1.877 
 
 2.732 
 
 + 0.855 
 
 8 
 
 1-595 
 
 0.161 
 
 i -756 
 
 to 
 
 G> 
 
 to 
 
 + 0.976 
 
 9 
 
 i -576 
 
 0 . 161 
 
 1-737 
 
 2.732 
 
 + O.995 
 
 10 
 
 1-835 
 
 0. 161 
 
 1.996 
 
 2.732 
 
 +0.736 
 
 11 
 
 1.929 
 
 0. 161 
 
 2 .090 
 
 2.732 
 
 4- 0.642 
 
 12 
 
 1.900 
 
 0. 161 
 
 2 .061 
 
 2.732 
 
 4 - 0.671 
 
 13 
 
 1.952 
 
 0. 161 
 
 2.113 
 
 2.732 
 
 4- 0.619 
 
 14 
 
 I- 5 I 5 
 
 0. 161 
 
 1 .676 
 
 2.732 
 
 4 - 1.056 
 
 15-16 
 
 4.368 
 
 0.322 
 
 4.690 
 
 5-464 
 
 + 0.774 
 
 17-18 
 
 3-889 
 
 0.322 
 
 4.211 
 
 5.464 
 
 + 1-253 
 
 19 
 
 2.030 
 
 0 . 161 
 
 2 . 191 
 
 2.732 
 
 + 0.541 
 
 
 
 50% Increase in Diet. 
 
 
 
 20 
 
 2-594 
 
 0.247 
 
 2.841 
 
 4.104 
 
 + 1.263 
 
 21-22 
 
 5-545 
 
 0.494 
 
 6.039 
 
 8.208 
 
 4 - 2 . 169 
 
 23-24 
 
 5-535 
 
 0.494 
 
 6 .029 
 
 8.208 
 
 + 2.179 
 
 25-26 
 
 5-183 
 
 0.494 
 
 5-677 
 
 8.208 
 
 + 2.531 
 
 27 
 
 2.655 
 
 0.247 
 
 2 .902 
 
 4.IO4 
 
 4- 1 .202 
 
 
 
 100% Increase in Diet. 
 
 
 
 28 
 
 3-682 
 
 0.256 
 
 3-938 
 
 5-464 
 
 + 1.526 
 
 29-30 
 
 7-858 
 
 0.512 
 
 8.370 
 
 IO.928 
 
 + 2.558 
 
 31-32 . 
 
 7.286 
 
 0.512 
 
 7.798 
 
 IO.928 
 
 + 3 -i 3 o 
 
 33-34 
 
 8-577 
 
 0.512 
 
 9.089 
 
 IO.928 
 
 + 1.839 
 
 35-36 
 
 7 .606 
 
 0.512 
 
 8.118 
 
 IO.928 
 
 4- 2 .810 
 
 37-38 
 
 8.56O 
 
 0.512 
 
 9.072 
 
 IO.928 
 
 + 1.856 
 
 39-40 
 
 fO 
 
 00 
 
 cs 
 
 00 
 
 0.512 
 
 8-795 
 
 IO.928 
 
 + 2.133 
 
 41-42 
 
 8.762 
 
 0.512 
 
 9.274 
 
 10.928 
 
 + 1-654 
 
 43-44 
 
 8.592 
 
 0.512 
 
 9.104 
 
 IO.928 
 
 4- 1.824 
 
 
 
 Normal Diet. 
 
 
 
 45-46 
 
 4.920 
 
 0.634 
 
 5-554 
 
 5-464 
 
 — 0.090 
 
 47 
 
 2.435 
 
 0.317 
 
 2.752 
 
 2.732 
 
 4- 0.020 
 
Day. 
 
 I 
 
 2 
 
 3 
 
 4 
 
 5 
 
 6 
 
 7 
 
 8 
 
 9 
 
 io 
 
 ii 
 
 12 
 
 13 
 
 14 
 
 15 
 
 16 
 
 17 
 
 18 
 
 19 
 
 20 
 
 21 
 
 22 
 
 23 
 
 24 
 
 25 
 
 26 
 
 27 
 
 28 
 
 29 
 
 30 
 
 iy. 
 
 6 
 
 7 
 
 8 
 
 i 
 
 2 
 
 3 
 
 ;S 
 
 6 
 
 7 
 
 FASTING STUDIES. 
 
 Table IV ( continued ). 
 
 
 N in hair, 
 
 Total N 
 
 
 - 
 
 N in urine. 
 Grams. 
 
 feces and cage 
 wash. Gram. 
 
 output. 
 
 Grams. 
 
 N in food. 
 Grams. 
 
 N balance. 
 
 
 Second Fast. 
 
 
 
 I .081 
 
 0.054 
 
 i -135 
 
 
 — 1 135 
 
 i .197 
 
 0.054 
 
 1 .251 
 
 
 — 1*251 
 
 1.209 
 
 0.054 
 
 1.263 
 
 
 — 1263 
 
 1 .202 
 
 0.054 
 
 1.256 
 
 
 — I .256 
 
 1 .251 
 
 0.054 
 
 1305 
 
 
 — 1.305 
 
 1 .471 
 
 0.054 
 
 1.525 
 
 
 — 1.525 
 
 1.128 
 
 0.054 
 
 1 . 182 
 
 
 — I . 182 
 
 1.368 
 
 0.054 
 
 1 .422 
 
 
 — I .422 
 
 1 .218 
 
 0.054 
 
 1 .272 
 
 
 — I.272 
 
 1. 194 
 1.158 
 
 0.054 
 
 0.054 
 
 1 .248 
 
 1 .212 
 
 
 — I .248 
 
 I .212 
 
 1-332 
 
 0.054 
 
 1.386 
 
 
 — I .386 
 
 1 -357 
 
 0.054 
 
 1 .411 
 
 
 I .41 * 
 
 1 . 182 
 
 0.054 
 
 1.236 
 
 
 I .236 
 
 1.207 
 
 0.054 
 
 1 .261 
 
 
 I .261 
 
 1-344 
 
 0.054 
 
 1*398 
 
 
 — I .398 
 
 1 .119 
 
 0.054 
 
 1.273 
 
 
 — I.273 
 
 1 . 193 
 
 1 -335 
 
 0.054 
 
 0.054 
 
 1.247 
 
 1.389 
 
 
 —I.247 
 — 1.389 
 
 1.257 
 
 1 .221 
 
 0.045 
 
 0.054 
 
 1 .311 
 
 1*275 
 
 
 — i - 3 11 
 
 — 1.275 
 
 1 .203 
 
 0.054 
 
 1.257 
 
 
 — 1*257 
 
 1 .315 
 
 0.054 
 
 1.369 
 
 
 — 1369 
 
 1 .216 
 
 0.054 
 
 1 .270 
 
 
 — 1 .270 
 
 1 . 145 
 1.126 
 
 0.054 
 
 0.054 
 
 1. 199 
 
 1 .180 
 
 
 — 1. 199 
 
 — 1 . 180 
 
 1 -495 
 
 0.054 
 
 i *549 
 
 
 — 1*549 
 
 i« 5*9 
 
 2 .089 
 
 0.054 
 
 0.054 
 
 i *573 
 
 2*143 
 
 
 1 * 573 
 
 — 2.143 
 
 2.298 
 
 0.054 
 
 2.352 
 
 
 — 2.352 
 
 Table V. — Body Weight and Creatinine Coefficient. 
 
 Body 
 
 weight. 
 
 Kgs. 
 
 Gross gain 
 or loss. 
 Kgs. 
 
 Daily gain 
 or loss. 
 
 Kgs. 
 
 Per cent, 
 gross gain 
 or loss. 
 
 Per cent, 
 daily gain 
 or loss. 
 
 Creatinine 
 
 coefficient. 
 
 3-42 
 
 3.40 
 
 3-4* 
 
 
 Preliminary Period. 
 
 
 N N 
 
 OOO 
 
 O 06 
 
 
 
 First Fast. 
 
 
 
 
 3.26 
 
 3*9 
 
 3**3 
 
 — 0.15 
 
 0.22 
 
 0.28 
 
 — 0.15 
 
 — 0.07 
 
 — 0.06 
 
 — 4.40 
 
 6-45 
 
 8.21 
 
 — 4.40 
 
 2.15 
 
 1.88 
 
 O . 009 
 
 0.009 
 
 O .009 
 
 2-95 
 
 O .46 
 
 — 0.18 
 
 13-49 
 
 5-75 
 
 O.OII 
 
 2.91 
 
 2.86 
 
 — 0.50 
 
 — 0-55 
 
 — 0.04 
 
 — 0.05 
 
 14.66 
 
 16.13 
 
 1-35 
 
 I .72 
 
 0 .008 
 
 0.010 
 
FASTING STUDIES. 39 
 
 Table V (continued). 
 
 
 Body 
 
 weight. 
 
 Gross gain 
 or loss. 
 
 Daily gain 
 or loss. 
 
 Per cent, 
 gross gain 
 
 Per cent, 
 daily gain 
 
 Creatinine 
 
 Day. 
 
 Kgs. 
 
 Kgs. 
 
 Kg. 
 
 or loss. 
 
 or loss. 
 
 coefficient. 
 
 8 
 
 ft 
 
 2.77 
 
 — O.64 
 
 First Fast. 
 
 — 0.09 
 
 18.77 
 
 3-15 
 
 0.009 
 
 9 
 
 IO\ 
 
 2.60 
 
 — O.81 
 
 — 0.17 
 
 23-75 
 
 6.14 
 
 0.010 
 
 j 
 
 ii 
 
 2.49 
 
 — O.92 
 
 — 0. 11 
 
 26.98 
 
 4-23 
 
 0.009 
 
 12 
 
 2.31 
 
 — I .IO 
 
 — 0.18 
 
 32.26 
 
 7-23 
 
 0.008 
 
 13 
 
 2.22 
 
 — I.I9 
 
 — 0.09 
 
 34 - 6 o 
 
 3-90 
 
 O.OO7 
 
 14 
 
 2.05 
 
 — 1.36 
 
 — 0.17 
 
 39.88 
 
 7.66 
 
 0.005 
 
 15 
 
 I.85 
 
 — 1.56 
 
 — 0.20 
 
 45-75 
 
 9-75 
 
 0.008 
 
 i 
 
 i *95 
 
 + 0.10 
 
 Feeding Periods. 
 
 Preliminary. 
 
 + 0.10 +6.41 
 
 + 5-40 
 
 0.005 
 
 2 
 
 2.25 
 
 + 0.40 
 
 + 0.30 
 
 25-63 
 
 15 - ,38 
 
 0.006 
 
 3 
 
 2-35 
 
 + 0.50 
 
 + 0.10 
 
 32.04 
 
 4.44 
 
 0.006 
 
 4 
 
 2 .24 
 
 + 0.39 
 
 — 0. 11 
 
 24.99 
 
 — 4.68 
 
 0.006 
 
 5 
 
 2 .20 
 
 + 0.35 
 
 Normal Diet. 
 
 — 0.04 
 
 22.43 
 
 — 1.78 
 
 0.006 
 
 6 
 
 2 . 12 
 
 + 0.27 
 
 — 0.08 
 
 17.30 
 
 — 3-64 
 
 O.OOS 
 
 7 
 
 2 . 19 
 
 + 0.34 
 
 + 0.07 
 
 21.80 
 
 + 3-30 
 
 0.008 
 
 8 
 
 2 .24 
 
 + 0.39 
 
 + 0.05 
 
 24.99 
 
 2.28 
 
 0.008 
 
 9 
 
 2.25 
 
 + 0.40 
 
 + 0.01 
 
 25.63 
 
 0-45 
 
 0.008 
 
 IO 
 
 2 .26 
 
 + 0.41 
 
 + 0.01 
 
 26.27 
 
 0-44 
 
 0.009 
 
 ji 
 
 2 .27 
 
 + 0.42 
 
 + 0.01 
 
 26.91 
 
 0-44 
 
 O.OO9 
 
 12 
 
 2 .29 
 
 + 0.44 
 
 + 0.02 
 
 28.20 
 
 0.88 
 
 0.008 
 
 *3 
 
 2.28 
 
 + 0.43 
 
 — 0.01 
 
 27.56 
 
 — 0.44 
 
 0.008 
 
 14 
 
 2.31 
 
 + 0.46 
 
 + 0.03 
 
 29.48 
 
 + 1. 31 
 
 0.008 
 
 15 
 
 2-34 
 
 + 0.49 
 
 + 0.03 
 
 31.40 
 
 1.29 
 
 0.010 
 
 16 
 
 2.36 
 
 + 0.51 
 
 + 0.02 
 
 32.68 
 
 0.85 
 
 0.010 
 
 17 
 
 2.36 
 
 + 0.51 
 
 0.00 
 
 32.68 
 
 0.00 
 
 0.009 
 
 1 8 
 
 2.36 
 
 + 0.51 
 
 0.00 
 
 32.68 
 
 0.00 
 
 0.009 
 
 19 
 
 2.38 
 
 + 0.53 
 
 + 0.02 
 
 33-96 
 
 0.85 
 
 O.OII 
 
 20 
 
 2 .42 
 
 + 0.57 
 
 50% Increase in Diet. 
 
 + 0.04 +36.53 
 
 + 1.68 
 
 O.OII 
 
 21 
 
 2.49 
 
 + 0.64 
 
 + 0.07 
 
 41 .01 
 
 2.89 
 
 O.OII 
 
 22 
 
 2.50 
 
 + 0.65 
 
 + 0.01 
 
 41.65 
 
 0.40 
 
 0.010 
 
 23 
 
 2*57 
 
 + 0.72 
 
 + 0.07 
 
 46.14 
 
 2.80 
 
 O.OII 
 
 24 
 
 2 .61 
 
 + 0.76 
 
 + 0.04 
 
 48.70 
 
 i -55 
 
 O.OII 
 
 25 
 
 2.63 
 
 + 0.78 
 
 + 0.02 
 
 49.98 
 
 0-77 
 
 0.012 
 
 26 
 
 2 .67 
 
 + 0 . 82 
 
 + 0.04 
 
 52.55 
 
 1-52 
 
 O.OII 
 
 27 
 
 2-75 
 
 + 0.90 
 
 + 0.08 
 
 57-67 
 
 3.00 
 
 0.013 
 
 28 
 
 2.84 
 
 + 0.99 
 
 100% Increase in Diet. 
 
 + 0.09 63.44 
 
 3-27 
 
 0.013 
 
 29 
 
 2.85 
 
 + 1 .00 
 
 + 0.01 
 
 64.08 
 
 0-35 
 
 0.012 
 
 30 
 
 2.92 
 
 + 1 .07 
 
 + 0.07 
 
 68.57 
 
 2-45 
 
 0.012 
 
 31 
 
 3-00* 
 
 + J 
 
 + 0.08 
 
 73-70 
 
 2.74 
 
 0.012 
 
 32 
 
 3 00 
 
 -4 J JK 
 
 00 
 
 73-70 
 
 0.00 
 
 0.012 
 
33 
 
 34 
 
 35 
 
 36 
 
 37 
 
 38 
 
 39 
 
 40 
 
 4 i 
 
 42 
 
 43 
 
 44 
 
 45 
 
 4 6 
 
 47 
 
 I 
 
 2 
 
 3 
 
 4 
 
 5 
 
 6 
 
 7 
 
 8 
 
 9 
 
 io 
 
 ii 
 
 12 
 
 13 
 
 14 
 
 15 
 
 16 
 
 17 
 
 18 
 
 19 
 
 20 
 
 21 
 
 22 
 
 23 
 
 24 
 
 25 
 
 26 
 
 27 
 
 28 
 
 29 
 
 FASTING STUDIES. 
 
 Table V ( continued ). 
 
 Body 
 
 weight. 
 
 Kgs. 
 
 Gross gain 
 or loss. 
 Kgs. 
 
 Daily gain 
 or loss. 
 
 Kg. 
 
 Per cent, 
 gross gain 
 or loss. 
 
 Per cent, 
 daily gain 
 or loss. 
 
 
 100% Increase in 
 
 Diet. 
 
 
 3 09 
 
 + 1.24 
 
 + O.09 
 
 79.46 
 
 3-00 
 
 3.10 
 
 + 1.25 
 
 + 0.01 
 
 80.10 
 
 O.32 
 
 3.20 
 
 + i -35 
 
 + 0.10 
 
 86.50 
 
 3.22 
 
 3-24 
 
 + i -39 
 
 + 0.04 
 
 89.08 
 
 I.25 
 
 3-32 
 
 + i -47 
 
 + 0.08 
 
 94.20 
 
 2-47 
 
 3-35 
 
 + 1.50 
 
 + 0.03 
 
 96.12 
 
 O.90 
 
 3 40 
 
 + i -55 
 
 + 0.05 
 
 99-33 
 
 I.49 
 
 3 -46 
 
 + 1 .61 
 
 + 0.06 
 
 103.17 
 
 1.76 
 
 3-50 
 
 + 1.65 
 
 + 0.04 
 
 105.74 
 
 1 . 16 
 
 3*52 
 
 + 1.67 
 
 + 0.02 
 
 107.02 
 
 0-57 
 
 3-56 
 
 + 1. 71 
 
 + O.O4 
 
 109.58 
 
 1. 14 
 
 3-59 
 
 + 1-74 
 
 + 0.03 
 
 Normal Diet. 
 
 hi .50 
 
 0.84 
 
 3-56 
 
 + 1. 71 
 
 — 0.03 
 
 109.58 
 
 — 0.83 
 
 3-54 
 
 + 1.69 
 
 — 0.02 
 
 108.30 
 
 — 0.56 
 
 3-54 
 
 + 1.69 
 
 0.00 
 
 108.30 
 
 0.00 
 
 
 
 Second Fast. 
 
 
 
 3-42 
 
 — 0.12 
 
 — 0.12 
 
 — 3-39 
 
 — 3-39 
 
 3-32 
 
 — 0.22 
 
 — 0. 10 
 
 6.21 
 
 2 .92 
 
 3.26 
 
 — 0.28 
 
 — 0.06 
 
 7.91 
 
 1. 81 
 
 3 -n 
 
 — 0.43 
 
 — 0.15 
 
 12.15 
 
 4.60 
 
 3 07 
 
 — 0.47 
 
 — 0.04 
 
 13-28 
 
 1.28 
 
 3 05 
 
 — 0.49 
 
 — 0.02 
 
 13.84 
 
 0.65 
 
 2-95 
 
 — 0-59 
 
 — 0. 10 
 
 16.66 
 
 3.28 
 
 2.89 
 
 — 0.65 
 
 — 0.06 
 
 18.36 
 
 2.03 
 
 2.86 
 
 — 0.68 
 
 — 0.03 
 
 19.21 
 
 1 .04 
 
 2.80 
 
 — 0.74 
 
 — 0.06 
 
 20.90 
 
 2 . 10 
 
 2.77 
 
 — 0.77 
 
 — 0.03 
 
 21-75 
 
 1.07 
 
 2.72 
 
 — 0.82 
 
 — 0.05 
 
 23.16 
 
 1 .80 
 
 2.68 
 
 — 0.86 
 
 — 0.04 
 
 24.29 
 
 1.47 
 
 2.64 
 
 — 0.90 
 
 — 0.04 
 
 25-42 
 
 1.49 
 
 2.58 
 
 — 0.96 
 
 — 0.06 
 
 27 . 12 
 
 2 .27 
 
 2.51 
 
 — 1.03 
 
 — 0.07 
 
 29.09 
 
 2.71 
 
 2.49 
 
 — 1.05 
 
 — 0.02 
 
 29.66 
 
 0-79 
 
 2-45 
 
 — 1.09 
 
 — 0.04 
 
 30.79 
 
 1.60 
 
 2.40 
 
 — 1. 14 
 
 — 0.05 
 
 32.20 
 
 2 .04 
 
 2-37 
 
 — 1. 17 
 
 — 0.03 
 
 33-05 
 
 1-25 
 
 2.36 
 
 — 1. 18 
 
 — 0.01 
 
 33-33 
 
 0.42 
 
 2.31 
 
 — 1.23 
 
 — 0.05 
 
 34-74 
 
 2 . 12 
 
 2.21 
 
 — 1-33 
 
 — 0. 10 
 
 37-57 
 
 4-33 
 
 2.18 
 
 — 1.36 
 
 — 0.03 
 
 38.42 
 
 i -35 
 
 2.15 
 
 — 1-39 
 
 — 0.03 
 
 39-26 
 
 i -37 
 
 2 . 11 
 
 — i -43 
 
 — 0.04 
 
 40.39 
 
 1.86 
 
 2.05 
 
 — 1.49 
 
 — 0.06 
 
 42.09 
 
 2.84 
 
 1.97 
 
 — i -57 
 
 — 0.08 
 
 44-35 
 
 3-90 
 
 1. 91 
 
 — 1.63 
 
 — 0.06 
 
 46.04 
 
 
 Creatinine 
 
 coefficient. 
 
 0.013 
 0.012 
 0.012 
 0.012 
 0.013 
 0.013 
 0.013 
 0.012 
 0.013 
 0.013 
 0.013 
 O .012 
 
 O.OII 
 
 0.012 
 
 O.OII 
 
 0.009 
 
 0.009 
 
 O.OO9 
 
 0.010 
 
 0.010 
 
 0.009 
 0.009 
 0.009 
 0.009 
 0.008 
 0.010 
 o .009 
 0.009 
 0.009 
 O.OOS 
 0.006 
 O .008 
 0.008 
 0.008 
 0.007 
 0.006 
 0.006 
 0.005 
 0.005 
 0.006 
 0.006 
 
FASTING STUDIES. 4 1 
 
 Table VI. — Nitrogen in Diet, Feces, Hair and Cage Washings. 
 
 Feeding periods. 
 
 Preliminary 
 
 period. 
 
 11 days. 
 
 First 
 
 fast. 
 
 15 days. 
 
 Pre- 5056 
 
 liminary. Normal, increase. 
 
 4 days. 15 days. 8 days. 
 
 100$ 
 
 increase. 
 17 days. 
 
 Normal. 
 
 3 days. 
 
 Second 
 
 fast. 
 
 30 days. 
 
 
 
 Food (per day). 
 
 
 
 
 Meat 2.533 
 
 
 3.618* 2.533 3-799 
 
 5.066 
 
 2-533 
 
 
 Crackers 0.1977 
 
 
 2.930* 0.1977 0.303 
 
 0-395 
 
 O.1977 
 
 
 Lard 0 . 0008 
 
 
 0.003* 0.0008 0.0012 
 
 0.002 
 
 0.0008 
 
 
 Bone ash. . . . 0.0003 
 
 
 0 . 003* 0 . 0003 0 . 0005 
 
 0.0005 
 
 0.0003 
 
 
 Total (grams) 2 . 732 
 
 
 6.554* 2.732 4.104 
 
 5.464 
 
 2.732 
 
 
 H a O 250 cc. 
 
 250 CC. 
 
 250 cc. 250 cc. 250 cc. 
 
 250 CC. 
 
 250 CC. 
 
 250 CC. 
 
 
 
 Feces. 
 
 
 
 
 Grams 41 .89 
 
 3-5 
 
 76.6 61.0 
 
 92 .0 
 
 20.0 
 
 9.6 
 
 Wt. of N 1.691 
 
 O.309 
 
 2.143 1.596 
 
 3-540 
 
 0-739 
 
 0.776 
 
 % N 4.037 
 
 8.874 
 
 2.798 2.616 
 
 3-839 
 
 3-695 
 
 8.100 
 
 Gms.N per day 0.154 
 
 0.021 
 
 0.113 0.199 
 
 0.208 
 
 O.269 
 
 0.026 
 
 
 
 Hair. 
 
 
 
 
 Grams 5.0 
 
 
 130 
 
 
 
 6.0 
 
 Wt. of N 0.575 
 
 
 1-555 
 
 
 
 0.666 
 
 % N 11.50 
 
 
 11.961 
 
 
 
 11 . 100 
 
 Gms.N per day 0.052 
 
 
 0.025 
 
 
 
 0.022 
 
 
 
 Cage washings. 
 
 
 
 
 Wt. of N 0.326 
 
 0.268 
 
 1 . 084 (47 days) 
 
 
 0.189 
 
 Gms.N per day 0.029 
 
 O.OI78 O.O23 
 
 
 
 0.006 
 
 
 Total Nitrogen Excreted. 
 
 
 
 
 Grams 2.592 
 
 
 . . 
 
 . . 
 
 
 
 Gram per day 0.235 
 
 0.064 
 
 0.161 0.247 
 
 0.256 
 
 O.317 
 
 0.054 
 
 muscle was but slightly lowered. This pronounced decrease of creatine 
 found by us in fasting muscle is a most significant fact and shows clearly 
 that in fasting we can not, with accuracy, consider the total amount of 
 excreted creatine as resulting from the complete and permanent disin- 
 tegration of muscular tissue. It appears that the creatine of the urine 
 is derived either from disintegrating muscular tissue or is removed, in 
 some manner from such tissues which are still functioning within the body. 
 
 12. As a result of our experiments we have shown that in repeated 
 fasting there is a slower and less profound tissue disintegration during 
 the second fast, indicating a greater resistance of the body, acquired as 
 a result of the initial fast. This increased resistance, noted in our ex- 
 periments, following fasting may indicate that the “repeated fast,” if 
 properly regulated, may possess important therapeutic properties. The 
 gi eater length of the second fast could not have been due to increased 
 fat stores, for the fat present in the body of the animal at the beginning 
 of the second fast was, according to our calculations, only fifty grams 
 more than at the beginning of the first fast. 
 
 * Total weight of nitrogen fed during the four days. 
 
42 
 
 FASTING STUDIES. 
 
 References. 
 
 References. — i. Brugsch: Oppenheimer, Handbuch der Biochemie, 4, 295. 2. 
 
 Pashutin, Pathological Physiology , 1902. Quoted by Pashutin. 1 3. Albitsky. 4. 
 Tuvim. 5. Manassein. 6. Kagan. 7. Ugrumoff. 8. Sadoven. (9. Schulz and Man- 
 gold, p. 419. 10. Schulz and Stiibel, p. 431. 11. Schulz and Hempel, p. 439. 12. 
 
 Schulz, p. 462;) Arch. ges. Physiol ., 114, 1906. 13. Voit, Z. fur Biol., 2, 307 
 
 (1866). 14. Seeland, Biol. Centr., 7, 145 (1887). 15. Richet, Compt. rend. soc. 
 
 biol., 61, 546 (1906). Quoted by Pashutin. 16. Rucher. 17. Smith. 18. Ranke. 
 19. Pettenkofer and Voit. 20. Oppenheim. 21. Sadoven. 22. Richet. 23. Howe, 
 Mattill and Hawk, Proc.Soc. Biol. Chem., July, 1910. 24. Falk ,Beitr. Physiol. (Stuttgart), 
 1875, quoted by Pashutin. 25. Kumagawa and Miura, Arch. Physiol., 1898, 431. 
 26. Pfliiger, Arch. Physiol., 119, 119 (1907). 27. Daddi, Arch. Hal. biol., 30, 439 
 
 (1899). 28. Grandis, Arch. ital. biol., 12, 237 (1889). 29. Krawkaw, Wratsch, 1891. 
 
 30. L’abb6 and Vitry, Compt. rend soc. biol., 62, 699 (1907). 31. Luciani and Bufalini, 
 
 Arch. sci. med., 5, 338 (1882). 32. Folin, Z. physiol. Chem., 41, 233 (1904). 33. Oster- 
 
 berg and Wolf, Biochem. Z., 5, 304 (1907). 34. Underhill and Kleiner, J. Biol. Chem., 
 
 4, 165 (1908). 35. Schondorf, Arch. ges. Physiol., 117,257(1907). 36. Leffmann, 
 
 Z. physiol. Chem., 57, 476 (1908). 37. Weber, Arch. exp. Path. Pharm., 58, 93 (1907). 
 
 38. Dorner, Z. physiol. Chem., 52, 93 (1907). 39. Bohtlingk, Arch. sci. biol. St. 
 
 Petersburg, 8, 483 (1903). 40. Van Hoogenhuyze and Verploegh, Z. physiol. Chem., 
 
 46, 415 (1905). 41. Benedict, Carnegie Institution of Washington, Publication No. 
 
 77, 386 (1907). 42. Benedict and Diefendorf, Am. J. Physiol., 18, 362 (1907). 43. 
 
 Cathcart, Biochem. Z., 6, 109 (1907). 44. Gies, Am. J. Physiol., 5, 235 (1901). 45. 
 
 Hawk, University of Pennsylvania Medical Bulletin, Dec., 1905. 46. Gies, Am. 
 
 Physiol., 14, 403 (1905). 47. Hawk and Grindley, Proc. Am. Soc. Biol. Chem., 1908. 
 
 9. Gill and Grindley, J. A. C. S., 31, 695 (1909). 48. Marcuse, Arch. Physiol., 
 
 59, 232 (1896). 49. Benedict and Gephart, J. A. C. S., 30, 1760 (1908). 50. 
 
 Folin, Z. physiol. Chem., 32, 504 (1901); 36, 333 (1902); 37, 548 (1903). 51. Osterberg 
 
 and Wolf, J . A. C. S., 31, 421 (1909). Gill, Allison and Grindley, J. A. C. S., 
 
 31, 1078 (1909). Benedict, Proc. Am. Soc. Biol. Chem., July, 1910. 52. Folin, Z. physiol. 
 
 Chem., 37, 161 (1902). 53. Benedict and Meyers, Am. J. Physiol., 18, 397 (1907). 
 
 54. Poduschka, Arch. exp. Path. Pharm., 44, 59 (1900). 55. Lusk, The Science of 
 
 Nutrition, 1909, 46. 56. Folin, Am. J. Physiol., 13, 66 (1905). 57. Fowler and 
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 Path. Therapie, 1, 419 (1905); Baumstark and Mohr, Ibid., 3, 675 (1906); Lehman, 
 Zuntz, et. al., Virchow's Arch., 131 (suppl.), (1893). 61. Klerker, Biochem. Z., 3, 45 
 
 (1907). 62. Closson, Am. J. Physiol., 16, 252 (1906). 63. Shaffer, Am. J. Physiol ., 
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 Festschrift, p. 15. 
 
 1 These references were obtained from the English translation of Pashutin’s 
 book, prepared under the direction of Francis G. Benedict of the Carnegie Institution. 
 
VITA 
 
 The author graduated from the University of Illinois in 1906 with 
 the degree of Bachelor of Science in Chemistry. He received a scholar- 
 ship in chemistry, in the University of Illinois for the year 1906 to 1907, 
 and obtained the degree of Master of Arts in 1907, from that institution, 
 with a thesis in physical chemistry. During the following year he was 
 assistant chemist in the Laboratory of Physiological Chemistry of the 
 University of Illinois and did graduate work in physiological chemistry. 
 As assistant in physiological chemistry in the University of Illinois he 
 pursued * graduate courses in physiological chemistry, under Professor 
 Philip B. Hawk, and in physiology and histology. 
 
 He is a member of Phi Lambda Upsilon, Sigma Xi, the Gamma 
 Alpha Graduate Scientific Fraternity, the American Chemical Society, 
 the American Association for the Advancement of Science, the American 
 Society of Animal Nutrition, the Illinois Academy of Science, the Ameri- 
 can Society of Biological Chemists, and the American Health League. 
 
 PUBLICATIONS 
 
 1. “The Electrolytic Corrosion of Brasses” (with Azariah T. Lincoln 
 and David Klein): Journal of Physical Chemistry , 11, 501 (1907). 
 
 2. “Comparative Tests of Spiro’s and Folin’s Methods for the Deter- 
 mination of Ammonia and Urea” (with P. B. Hawk) : Journal of Biolog - 
 ical Chemistry , 5, 477 (1909). 
 
 3. “On the Preservation of Feces” (with T. A. Rutherford and P. B. 
 Hawk): Proceedings of the American Society of Biological Chemists , 
 
 1909, xlix. 
 
 4. “A Study in Repeated Fasting” (with P. B. Hawk) : Proceedings 
 of the American Society of Biological Chemists , July, 1910. 
 
 5. “Fasting Studies on Men and Dogs” (with H. A. Mattill and P. B. 
 Hawk) : Proceedings of the American Society of Biological Chemists , 
 
 July. I91O. 
 
 The author wishes to express his sincere thanks to Professor Philip 
 B. Hawk, under whose direction this investigation was conducted, for 
 the kindly counsel and unerring advice, which were stimulating factors 
 during the progress of this research and through the author’s graduate 
 work in general.