EH3RC-Y TaA::'SF0PJ'tA.TI01T3 DUPJITG HORIZOIITAL WALimTCr Benedict and Murschhauser LIBRARY OF THE NEW YORK STATE COLLEGE OF HOME ECONOMICS CORNELL UNIVERSITY ITHACA, NEW YORK ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING BY FRANCIS G. BENEDICT AND HANS MURSCHHAUSER WASHINGTON, D. C. Published by the Carnegie Institution of Washington 1915 Cornell University Library The original of tiiis book is in tine Cornell University Library. There are no known copyright restrictions in the United States on the use of the text. http://www.archive.org/details/cu31924003145848 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING BY FRANCIS G. BENEDICT AND HANS MURSCHHAUSER WASHINGTON, D. C. Published bt the Carnegie Institution of Washington 1915 CONTENTS. Page Introduction 5 Methods of determining the energy transformations during walking 7 Methods of studying the gaseous exchange during walking 8 Fimdamental principle of studying the gaseous exchange incidental to walking. . 9 Units of measurement used in walking experiments 10 Previous researches on the gaseous exchange during walking 12 Summary of results of previous observations 21 Methods and apparatus for studies of muscular work 29 Description of apparatus used in this research 31 Universal respiration apparatus 31 TreadmiU 34 Accessory apparatus 36 Method of recording the respiration-rate 37 Method of recording the pulse-rate 37 Step counter 38 Method of measuring height to which the body is raised 39 Plan of research 42 General routine of the experiments 44 Standing experiments • 44 Sitting experiments 45 Walking and running experiments 46 Experiments with food 46 Subjects 47 Statistics of experiments 48 Discussion of results 61 Basal values 61 Basal metabolism of subject 1 62 Influence of food and body position 65 Basal metabohsm of subject II 66 Metabolism in the lying position 66 Metabolism in the sitting position 67 Comparison of the metabohsm in the lying and sitting positions 69 Metabolism in various standing positions 70 Influence of food upon metabohsm in the standing position 72 Metabolism during walking 76 Walking experiments with subject 1 76 Experiments without food 77 Experiments with food 80 Energy required for the elevation of the body '. 80 Walking experiments with subject II 81 Experiments without food 82 Experiments with food 87 Influence of the character of diet on the heat-output per unit of work 93 Influence of fatigue upon the heat-output per unit of work 94 Comparison of the heat-output per unit of work dtu:ing running with that obtained during walking 96 Analysis of the mechanics of locomotion 98 ILLUSTRATIONS. Fig. 1. General view of apparatus used for walking experiments 32 2. Schematic outline of universal respiration apparatus 34 3. TreadmiU designed by E. H. Metcalf 35 4. Detail of ball bearing for stedl tubes on the treadmill 38 5. Step counter 38 6. Apparatus for recording the height to which the body is lifted, and step counter with connections 40 7. Typical kymograph record showing character of step 41 3 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. INTRODUCTION. This investigation was undertaken after several long conferences with Professor Zuntz of Berlin and Professor Durig of Vienna, whose researches on the work of forward progression are classical. The pre- liminary experiments were made during the sojourn at the Nutrition Laboratory of Dr. Carl Tigerstedt of Helsingfors. Subsequently data were acquired by Messrs. H. L. Higgins and L. E. Emmes of the Laboratory staff. We wish to express our thanks to these gentlemen and particularly to Dr. Tigerstedt for the data regarding Subject I. A certain amount of walking on a level inevitably forms a part of the daily routine of nearly every Uving person, for even those who are designated as sedentary in their habits do a not inconsiderable amount of walking in the house or in short distances upon the street. To one who has not computed the actual distance traversed by the housewife during a day, the sum total of the distance walked is smprising. Such a control may readily be obtained with a simple pedometer, for although a pedometer can not be classified as an instrument of precision and is subject to many errors that are fre- quently overlooked, nevertheless it shows in a striking manner that very few individuals close a day of ordinary fife without having moved in forward progression a distance of not less than 2 or 3 kilometers. Not infrequently this distance is doubled or trebled by those who would ordinarily assume that they had not taken a particularly long walk. The personal experience of one of us while writing a report showed that the walking for a day consisted in going twice to and from the house, which was 400 meters from the laboratory, i. e., a total distance of 1,600 meters, and in walking about the laboratory while engaged in instruction and research. Throughout a period of several months the pedometer, which was carefully controlled and tested, showed that the average distance walked per day amounted to 7 miles (11.27 kilometers).^ Undoubtedly innumerable instances even more striking than this may be cited, which would show that it is reasonable to assume that practically aU persons do considerable walking during the course of 24 hours. Inasmuch as there are many individuals whose habit of life or profession requires a large amount of walking — for example, those walking to and from business, mail carriers, and espe^ 'Benedict, Proc. Am. Phil. Soc, 1910, 49, p. 162. 6 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. daily soldiers — an intimate knowledge of the physiology of walking is obviously of great importance. Practically all of the previous researches on the physiology of walking have been conducted either from a desire to study the conditions inci- dental to walking done in mountain chmbing, the results being of importance to physicians and those taking regular exercise, or, as in the case of the classical researches of Zuntz and Schumburg, from a desire to study the influence of walking on the metabolism, with a view to applying the results directly to army movements. In the present day, when special attention is directed towards efficiency, the minimizing of extraneous muscular movements, and the transportation of material by hand and leg motion with the least possible expenditure of physical energy, we may assume that there is every incentive for studying care- fully the physiology incidental to walking in a horizontal direction. While from the abstract physiological standpoint a study of all of the various factors incidental to walking is of great value, perhaps the most important phase of the investigation is the study of the energy transformations and the determination of the amount of nutrients in the food necessary to provide for such activity. The results of such study are of especial practical value in determining the energy require- ments of an army engaged in marching a certain distance over a level country. As previously stated, this thought dominated the study of Zuntz and Schumburg. A more universal appUcation of the results may be made by the physician who, if he knows the energy involved in walking, is able to prescribe more intelligently a definite amount of exercise for the ambulatory patient. Furthermore, as physical exercise, particularly walking, is an important factor in weight reduction and in athletic training, exact information as to the energy required may be put to practical use in such connection. Theoretically the movement of 1 kilogram 1 meter would call for no positive work other than that in overcoming the resistance of the air; nevertheless a considerable amount of work is required of the human body as a machine in accomplishing this feat of moving the mass in a horizontal direction. The apportionment of the total energy output of the body between that required for the maintenance of the vital functions and that required for walking is not, however, simple. When a person is walking, not only is energy required for the external mus- cular exercise, but a heat production is necessary for the entire main- tenance of the body activities, including muscular tonus, the work of circulation, respiratory muscles, and the external work of balancing the body in an upright position, none of these activities contributing directly to the work required to move the body in a horizontal direction. With only a knowledge of the amount of food eaten, it is impossible to estimate the proportion of food required for the activity of walking and that for vital maintenance. A closer analysis is therefore essential. INTRODUCTION. / METHODS OF DETERMINING THE ENERGY TRANSFORMATIONS DURING WALKING. It is necessary, first of all, to study the energy transformations which are pecuHarly incidental to walking. Thus, in the simplest case, if the person could walk directly in a vertical direction, a definite amount of external muscular work would be performed which would be repre- sented by the product of the weight of the body and the height walked. In walking in a horizontal direction, theoretically no external work is performed and there is no change in the potential energy of the body. We are thus unable to measure the energy output by the kilogram- meter, the unit most commonly used, or by any of the other ordinary work units. Consequently we have very Uttle information regarding the energy output. It is true that we must not disregard the extremely illuminating researches of earlier workers, who have attempted to estabhsh a con- stant, although with wide variations, which would show approximately the amount of energy required to move 1 kilogram 1 meter in a hori- zontal direction. These researches will be referred to in a subsequent section. It is important for us to note, however, that aside from such methods of calculation as are based upon the constant estabhshed by the earUer physiologists, we have no means of calculating the energy output required in walking. A possible method of measurement would be to determine the energy output directly by having the subject walk inside of a calorimeter. This has been attempted, although in an imperfect manner, by certain French investigators, including Hirn^ and Chauveau,^ who used the so-called "emission calorimeter" with a tread wheel. In these studies, however, a large proportion of the work was done in lifting the body, and hence the amount of forward progression, which is of special interest to us, is complicated by the very much greater work involved in the elevation of the body. Finally, it is possible, owing to the valuable computations and methods of research established by Zuntz, to study the respiratory exchange, namely, the carbon-dioxide output and the oxygen intake, and thus compute indirectly the total calorific output. This last method has been adopted by aU physiologists as the most suitable for the purpose. Practically all previous research has therefore been based upon the general principle of determining the total respiratory exchange both while the subject is walking and during rest when lying, sitting, or standing, and noting the increment in the carbon-dioxide output and oxygen intake due to walking. From the heat of combustion and analysis of pure nutrients, such as carbohydrate and fat, it has been computed that for each Hter of 'Hirn, Recherches sur I'fiquivalent m^oanique de la chaleur, Paris, 1858. ^Chaveau, Compt. rend., 1899, 129, p. 249. 8 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. oxygen required in the combustion, there are produced from 4.686 to 5.047 calories. When pure carbohydrate is burned, each Uter of oxygen utiUzed in the combustion corresponds to 5.047 calories, but when pure fat is burned each Uter of oxygen corresponds to 4.686 calories. Zuntz and his co-workers have prepared a table showing the calorific equivalent of the oxygen consumption when fat and carbohydrate are burned.^ This has been most ingeniously elaborated by WiUiams, Riche, and Lusk." By studjdng the output of carbon dioxide and the intake of oxygen, not only is information secured regarding the total oxidation of material, but also some idea is gained of the character of the combustion by noting the relationship between the volume of carbon dioxide given off and the oxygen absorbed. With equal volumes of carbon dioxide and oxygen, this relationship or "respiratory quotient" is 1.0 and indicates that carbohydrate has been exclusively burned. When the respiratory quotient approximates 0.7 the indication is that fat has exclusively been burned. By this means, therefore, it is possible to compute with great accuracy the heat output from the total oxygen consumption and the respiratory quotient. In certain earlier researches, and especially prior to the time when methods were devised by which oxygen absorption could be more easily determined than formerly, the measurements of the carbon- dioxide output alone were used, but a much greater error is introduced into the computations by this method. Unfortunately, of the two factors, carbon dioxide and oxygen, that which is of the greater sig- nificance, namely, oxygen, is the more difiicult of determination, while the measurement of the carbon-dioxide excretion is relatively a simple matter. METHODS OF STUDYING THE GASEOUS EXCHANGE DURING WALKING. Inasmuch as a study of the problem of the energy transformations during walking demands a careful study of the gaseous exchange, we find all the methods used based upon this principle. The simplest is that in which the subject walks inside a closed chamber by means of which the product of respiration — carbon dioxide — ^is easily collected. This method was first employed by Sond^n and Tigerstedt^ in the clas- sical research with their large respiration chamber in the Karolinska Institute in Stockholm. This chamber had a capacity of 100 cubic meters and, unlike any respiration chamber previously used, permitted the subject free movement. When the subject walked back and forth across the room, a considerable distance could be traversed. At that time only the carbon-dioxide output was determined with this method. ^Zuutz and Schumburg, Physiologie des Marsches, Berlin, 1901, p. 361. ^WilUams, Bdche, and Lusk, Journ. Biol. Chem., 1912, 12, p. 357. 'Sond6n and Tigerstedt, Skand. Archiv f. Physiol., 1895, 6, p. 165. INTEODUCTION. 9 A second method involves the attachment of certain breathing apph- ances, either nosepieces or mouthpiece, with an apparatus for meas- uring the volume of the expired air. When the breathing appliances have been adjusted, the subject assumes a certain body position and then walks along a movable path, such as a treadmill. This method was frequently used by Zuntz and Durig and their co-workers. A modification of this method is that in which the apparatus for measuring and samphng the expired air is carried upon the back of the subject, somewhat as a knapsack would be carried. The subject is then no longer confined to walking upon a treadmill, but may walk on level ground anywhere. This method was used extensively by Zuntz and Durig and their associates and by Douglas. Instead of having the apparatus carried by the subject, it may be transported by an assistant walking a suitable distance behind him. The subject thus breathes through the nose or mouth apphance, but is not obliged to support the heavy apparatus. This method was employed by Burgi, Schnyder, and, in certain experiments, by Kolmer and Brezina. Finally, it is possible to have the gas measuring and sampling appa- ratus in a fixed position and the subject attached to it by a long tube. He then walks in a clearly defined path, either back and forth across the room or in a circle of which the apparatus is the center. This method, which obviously limits appreciably the distance to be covered and the general freedom of the subject, has found slight use with certain French observers, particularly Amar. FUNDAMENTAL PRINCIPLE OF STUDYING THE GASEOUS EXCHANGE INCIDENTAL TO WALKING. The method of superimposition is the only one feasible for these studies. By this method the gaseous exchange during walking in dif- ferent positions must be carefully studied by one or more of the methods previously referred to; subsequently, in order to apportion specifically the energy transformation due to walking, it is necessary that a certain part due to the metaboHsm of maintenance be subtracted from the result obtained for the total gaseous exchange. The exact selection of the amount of energy transformed for main- tenance to be deducted from the total energy transformation has been a matter of considerable discussion. We have now come to realize that a human individual may subsist on numerous metabolic levels. The. subject sound asleep, without food in the stomach, has a minimum metaboHsm, but is utterly incapable of intellectual or physical activity. It would obviously be impracticable, if not indeed undesirable, to deduct this minimum metabolism from the measurement of the total metaboHsm and assume that the difference would be wholly due to the energy transformation due to walking. Even when the subject is lying awake, we still have a base-Hne which is far removed from the 10 ENEEGY TRANSFOBMATIONB DUBING HORIZONTAL WALKING. process of walking. To obtain the best expression of the superimposed energy requirement for forward progression, it is necessary to secure, as nearly as possible, the energy requirement of the subject in a position which involves all of the extraneous muscular movements incidental to the waking condition and in a vertical position. This may be obtained by deducting the metaboUsm required for the standing position. And yet the earUer researches have been most unsatisfactory in the attempts to measure the metabolism under these conditions. With the standing position we again have numerous possible varia- tions. The subject may stand in a completely relaxed position; he may possibly eliminate in a large part the effort of balancing by leaning on a staff or lying back shghtly out of the vertical against a support ; or he may stand in a fixed position with rigid muscles, such as that of "atten- tion." We should, theoretically at least, expect to find a considerable difference in the metabohsm necessary for these various upright positions. Durig has clearly pointed out^ that there are numerous arguments against assuming that the metaboUsm while standing in any one of these positions can rightly be deducted from that during walking to obtain the true energy transformation due to the walking, for it is quite possible that certain of the external muscular movements incidental to balancing and sustaining the body in an upright position may be greatly modified, if not indeed dispensed with, in the ordinary motions of for- ward progression. Consequently, one finds that, in previous researches, the base-line used almost universally among physiologists has been the metabolism observed with the subject lying awake without food in the stomach, i. e., in the post-absorptive condition. The assumption is then made that the incrementof metabolism duringwalking over that observed with the subject lying awake is a true measure of the metabolism due to the muscular exercise of moving the body in a horizontal direction. UNITS OF MEASUREMENT USED IN WALKING EXPERIMENTS. While under ordinary conditions the amount of work performed in any inanimate or animate motions is expressed in terms of foot-pounds, kilogrammeters, or calories, it is obvious that no one of these units can be appropriately employed for indicating the energy transformations during walking. In walking, a given weight is carried through a given distance. To be sure, there is inevitably a sUght lifting of the total weight of the body at each step due to the anatomical arrangement of the feet and leg-muscles, but this weight is again immediately lowered to the same position, so that, mechanically at least, there is no effective work accompUshed. The only external evidence of performance is that a given weight is moved forward a given distance. In this discussion we may for the present appropriately eliminate the possible effect of wind resistance produced by the body in walking or the 'Durig, Denksohriften d. math.-natur. Klasse d. kaieerl. Akad. d. Wissensch., 1909, 86, p. 267, INTRODUCTION. 11 external influence of the wind in a direction either favorable or unfav- orable for forward progression. But in practically all walking done by man considerable differences exist in the weight moved forward . With the bather walking on a flat sandy beach we have simply the weight of the body plus the negligible weight of the bathing-suit. On the other hand, with a pedestrian taking his "constitutional," we have the weight of the clothing, amounting to 3 or 4 kilograms, possibly supplemented by the weight of knapsack, camera, and other accessories. Finally we have the exaggerated case of the fully equipped trooper carrying knapsack, emer- gency rations, and a considerable quantity of ammunition. Thus we have the possible necessity of distinguishing between the movement due to the Ufting of body-weight and that due to lifting inert or dead weight. The subject is further comphcated when we attempt to analyze the weight of the living organism. The body is made up of bone, muscle, and fat. Fat may be looked upon as inert body-material, and when there is an excessive accumulation of fat, as with obese persons, it is possible that it may be considered in great part as dead weight. It is thus seen that the problem of studying the physiology of forward progression involves the question of analyzing the character of the weight transported. For the present the unit of accompUshment in the work of forward progression on a horizontal plane must be con- sidered as the transportation of 1 kilogram 1 meter, i. e., 1 "horizontal kilogrammeter . ' ' 12 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. PREVIOUS RESEARCHES ON THE GASEOUS EXCHANGE DURING WALKING. Before reporting and discussing the results of our own experiments, a brief abstract is given of all previous research in which the metabolism during walking has been studied. In thus reviewing the work of other investigators, it has seemed advisable to record the results on the basis of the movement of 1 kilogram over 1 meter of level path, i. e., 1 hori- zontal kUogranmieter, and to compare them in a large summary table rather than to give them under each research. This is done in table 1 (see pages 22 to 27). Observations of Smith, 1859. — The earliest attempt to measure the gaseous metaboUsm during walking was made by Edward Smith, who, in his memoirs entitled "Experimental inquiries into the chemical and other phenomena of respiration and their modifications by various physical agencies,"^ gives the details of two walking experiments. In these experiments Smith collected the products of respiration by attach- ing a mask to the face and forcing the expired air through specially constructed boxes, containing caustic potash to absorb the carbon dioxide. The mask had two openings, one for inspired air and the other for expired air, a valve system providing for the separation of the two currents of air. A dry gas-meter was attached at the intake point. The expired air was first passed through a Woulff bottle containing pumice-stone and sulphuric acid, then into a gutta-percha box contain- ing a solution of caustic potash, and finally through a second Woulfif bottle containing pumice-stone and sulphuric acid. The walking was done inside of a room and covered a distance of approximately 10 meters in each direction. All of the precautions incidental to modern experiments as to recording the barometric temperature and pressure were taken, and a further factor, which is only too frequently neglected in modern work, namely, pulse-rate, was also recorded. The subject carried a spirometer which weighed 7 pounds, but the exact method of transportation is not clear from Smith's description. Both experiments were made in one afternoon, with an intermission of an hour. During the first experiment he walked at the rate of 2 miles an hour and during the second at the rate of 3 miles an hour. For a base-line he deter- mined the metabohsm at rest without food and in a sitting position. Smith's values, which are given in English grains, were recomputed to grams by Sonden and Tigerstedt.'' While walking at the rate of 2 miles an hour, the carbon dioxide per minute was 1.173 grams; at 3 miles an hour, 1.674 grams; and sitting at rest without food, 0.482 gram. The energy per horizontal kilogrammeter computed from these values is given in table 1, page 22. The results obtained by Smith have been criticized by Gruber^ and Voit,* who both consider them 'Smith, Phil. Trans. Roy. Soc, London, 1859, 149, p. 681. ''SondSn and Tigerstedt, Skand. Arohiv f. Physiol., 1895, 6, p. 166 'Gruber, Zeitschr. f. Biol., 1891, 28, p. 470. *Voit, Hermann's Handbuch der Physiologie, 1881, 6, p. 201. PREVIOUS RESEARCHES ON GASEOUS EXCHANGE. 13 somewhat high. Katzenstein^ likewise considers the values too high and inexact and regrets that Smith did not state whether the walking was done on a level or on an incline. Although Smith gives the weight of the subject and the apparatus in one instance, Katzenstein criticizes the absence of body-weights for each experiment. Katzenstein's further criticism, that the carrying of the apparatus by the subject was a fault in the technique, is of special interest in view of the subsequent use of a portable gas-meter by Zuntz and his co-workers. Observations of Gruber, 1891. — The unusual interest in mountaineer- ing, which is particularly active in Switzerland, has resulted in a large number of physiological observations upon the effect of high altitudes on the human body. One of the earliest of the observations, which included measurements of the gaseous metabolism, was that of Gruber,^ who pubhshed the results of a research carried out in the Physiological Institute at Berne under the direction of Professor Kronecker. Gruber himself was the subject of the study. Instead of using a mask with two valves, Gruber employed a tube in the mouth for expiration, the nose being closed by the fingers of the left hand. During inspiration the rubber tube leading to the mouth was tightly closed by the teeth and air was inspired through the nose. The inspired air was passed into a U-tube containing soda to absorb the carbon dioxide. A rubber air-cushion, which could be compressed by the arm of the subject, permitted the accumulation of the excess of air during the expiration, and dm-ing inspiration this air was forced out through the absorbing vessels by pressure with the arm. Three experiments are reported in which the subject walked without other load than the apparatus. The rate of walking in the first experiment was 80 steps per minute. Two of the three experiments consisted of 10 minutes of walking followed by 10 minutes of sitting, Gruber concluding from his results that apparently as much carbon dioxide is excreted in the 10-minute rest after the short walking-period as during the walking. He also made three experiments while sitting in a chair prior to walking and two experiments after walking. On the basis of the sitting values he contends that with walking on a level the carbon-dioxide production is twice as great as with sitting. Both the rest and walking experiments were made 5 or 6 hours after food. Comparing the results of observations made both in trained and imtrained condition and considering the amount of carbon dioxide produced during rest as 1, he obtained the following figures:^ Rest. Walking. Ascent (untrained). Ascent (trained) . Series 1 . . . Series 2 . . . 1 1 '2.00 1.75 4.1 3.05 3.3 2.42 'Incorrectly stated as 1.89 by Gruber and recalculated by us. 'Katzenstein, Archiv f. d. ges. Physiol., 1891, 49, p. 331. Gruber, Zeitschr. f. Biol., 1891, 28, p. 466. HUd., p. 490. 14 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. From these data he concludes that the carbon-dioxide production of a working man is not a function of his work, since the metabolism decreases with practice. Observations of Katzenstein, 1891. — ^In the walking experiments of Smith and Gruber, only the carbon-dioxide production was deter- mined, and the first investigations, in which the oxygen consumption was also measured, were those carried out by Katzenstein^ in Zuntz's laboratory in Berlin. Emplojdng the Zuntz mouthpiece and valves for separating the inspired air from the expired air, Katzenstein sampled and analyzed the expired air, the measurement of the total amount of air expired being made with a gas-meter. There is consider- able doubt as to the conditions of the experiments. In certain instances it is specifically stated that the subjects were in the post-absorptive state, i. e., without food for 12 hours, but the general impression is given that in the majority of instances the experiments were not made with the subject in the post-absorptive condition. The rate of walking was from 51 to 92 meters per minute. The criticism was subsequently raised by Durig^ that the treadmill was not completely level, there being a sUght elevation of somewhat less than 1 degree. To provide a base-line for these observations, the metabolism of the subject was measured while he stood quietly upon the treadmill. Observations were likewise obtained with the subject in a lying position. The author concludes that the oxygen consumption for the unit of effective work is greater for small amounts of work than for large amounts, and that the respiratory quotient during work remains essentially unaltered. The computation of the values per horizontal kilogrammeter is given in table 1, page 22. Observations of Sonden and Tiger stedt, 1895. — The large respiration chamber in Stockholm afforded suflScient space for experiments in which the work of walking could be measured.* The carbon dioxide alone was determined and with most of the experiments moderate amounts of food were taken. The body-weight with clothing was recorded before and after the experiment. The subjects walked from 3,000 to 6,920 steps per hour without a load. As a base-line the authors used a value found for three resting periods when the subject was sitting, these periods being between periods of walking; large differences are shown in the carbon-dioxide output per hour during the resting periods. The energy per horizontal kilogrammeter, as com- puted from the carbon-dioxide output during the walking experiments, is given in table 1, page 22. Observations of Schumburg and Zuntz, 1896. — In connection with a series of experiments made in the Alps, Schumburg and Zuntz carried ^Katzenstein, Archiv f. d. ges. Physiol., 1891, 49, p. 330. ^Durig, Denkschritt. d. math.-natur. Klasse d. kaiserl. Akad. d. Wiasenaoh., 1909, 86, p 250 'Sonden and Tigerstedt, Skand. Archiv f. Physiol., 1895, 6, p. 1. PREVIOUS RESEARCHES ON GASEOUS EXCHANGE. 15 out a number of observations in Zuntz's laboratory in Berlin, using the treadmill.^ A dry gas-meter weighing 7 kilograms was employed and was carried by the subject. In most of the experiments the inspired and expired air were separated by a valve system and a mouthpiece and nose-clamp were used. It is interesting to note that in the severe work of going uphill in this series of experiments, the authors record that the breathing appliances were very uncomfortable, particularly the nose- clip. They accordingly trained themselves to inspire through the nose and expire through the mouth, a procedure which they satisfied them- selves gave accurate results. To estabhsh a base-hne, numerous exper- iments were made with the subject ia a sitting position, these being carried out in the laboratory in Berlia, in the hotel at Zermatt, and also in the camp and on the glacier in the mountains. Three walking experiments were made with Zuntz as subject in a room of the labora- tory buildiag and two on the treadmill, which was in a practically hori- zontal position. Two treadmill experiments were also made with Schumburg as a subject. The energy per horizontal kilogrammeter, as computed from the difference between the sitting values and the walk- ing values, is given in table 1, page 22. Observations of A. Loewy, J. Loewy, and L. Zuntz, 1897. — In connec- tion with their studies in the Alps, A. Loewy, J. Loewy, and L. Zuntz made several experiments in Berhn on the treadmill in the Landwirt- schafthche Hochschule.^ Two experiments were made with A. Loewy, three with J. Loewy, and five with L. Zuntz. A 6.6 kilogram dry gas- meter was carried by the subject on his back and connected with the Lob valve attached to the mouthpiece. The nose was closed with a clamp. For the experiments made with A. Loewy, a base-hne was used which was founded upon earUer observations. The resting values for J. Loewy and L. Zuntz were determined presumably with the subject sitting. No statement is made as to whether the subjects were in a post-absorptive condition or not. Certain experiments were also made at Col d'Olen, in which the subject likewise carried the dry gas-meter, but for these experiments the resting values were obtained with the subject in a lying position. The results for the observations with the subject walking on a level are summarized in table 1, page 22. Observations of L. Zuntz, 1899.— In his observations regarding the gaseous-exchange of bicycle-riders, Leo Zuntz included a number of experiments made on the treadmill in the Landwirtschafthche Hoch- schule.^ The statement is inade that the treadmill had an incHnation of approximately 1 degree. Of special interest in these observations is the fact that Leo Zuntz paid particular attention to the influence of speed upon the gaseous metaboHsm, varying the rate of walking from 'Schumburg and Zuntz, Arohiv f. d. ges. Physiol., 1896, 63, p. 461. ^A. Loewy, J. Loewy, and L. Zuntz, Archiv f. d. ges. Physiol., 1897, 66, p. 477. 'L. Zuntz, Untersuohungen ilber den Gasweohsel uud Energieumsatz dea Rad ahresHirsch- wald, Berlin, 1899. 16 ENERGY TRANSFORMATIONS DURING HORIZOIWAL WALKING. 53.45 meters per minute to 145.53 meters per minute. In the treadmill experiments the stationary wet gas-meter was used and consequently the subject was not obUged to carry an excessive load. The author discusses the absence of wind-resistance and points out that at the forced speeds, the treadmill ran so irregularly that it was frequently necessary for him to run two or three steps. The research is especially worthy of note as representing an attempt to secure unusual accuracy, although one could wish that the experiments had been made with the subject in the post-absorptive condition. An abstract of the results obtained at the different speeds is recorded in table 1, page 22. Observations of Setschenow and Schatemikow, 1900. — Setschenow and Schatermkow^ report a research made at the physiological institute of the University of Moscow, which included six waUdng experiments, the length varying from 64 to 75 minutes. Three of these experiments were preceded by a preliminary period of 6 to 9 minutes in which the subject stood. Basal values were obtained in two additional experi- ments with the subject at complete rest. In the walking periods the subject, who was the fireman of the institution, walked back and forth in the courtyard, the speed averaging 62.47 meters per minute; the distance walked in each experiment was usually 4,455 meters. All of the experiments were made 3 to 4 hours after a Ught breakfast of white bread and tea. As the data given are unfortunately insufficient for computing the values per horizontal kilogrammeter and do not lend themselves to tabular presentation, this research is not included in table 1.' Observations of Frentzel and Rea^h, 1901 . — In studying the influence of an unbalanced diet upon muscular work, Frentzel and Reach made a number of observations with the subject walking on the Zuntz treadmill in the level position in the LandwirtschaftUche Hochschule in Berlin.^ The rate of walking varied from 31.39 to 80.15 meters per minute. The resting metabolism was determined with the subject lying upon a couch, but both the resting and walking experiments were made with food in the stomach. Resting experiments were made on 16 days with subject F and on 12 days with subject R. A considerable number of resting experiments were made prior to the walking period and on days •other than the walking days. Durig has criticized the experiments as showing imusuaUy large fluctuations in the energy required per hori- zontal kilogrammeter. A summary of the results is given in table 1, page 22. Observations of Zuntz and Schumburg, 1901. — In their extensive research on the physiology of walking, Zuntz and Schumburg made 'Setschenow and Schatemikow, Le Physiol. Russe, 1900, 2, p. 44. *War conditions make it impracticable for me to commvmicate with my personal friend. Professor Schatemikow, regarding the figures in this research. As they stand, if a body-weight is assumed and » resting base-line deducted, values representing only about 50 per cent of those found in other published researches are obtained. — F. G. B. •Frentzel and Reach, Archiv f. d. ges. Physiol., 1901, 83, p. 477. PREVIOUS RESEARCHES ON GASEOUS EXCHANGE. 17 numerous observations of the respiratory exchange of two students, B and P.^ The experiments were made in Zuntz's laboratory and the treadmill was used. In most of the experiments the subjects carried the German army equipment with a considerable load, the maximum weight of the load being 31.5 kilograms. In some of the experiments they carried no load, and it is these latter experiments with which we are chiefly concerned. The basal values were obtained with the sub- ject lying upon a sofa. Both the resting and walking experiments were made after the subject had taken a Ught breakfast. The usual correction for the sUght elevation of the treadmill is considered. The experiments were so adjusted that both subjects could walk at the same time upon the treadmill and two complete sets of respiration apparatus, including gas-meters, were employed. The energy per horizontal kilogrammeter is given in table 1, page 22. Observations of Dwig and Zuntz, 1904- — Observations on the metabo- lism during walking on a horizontal level were carried out in Vienna, Col d'Olen, and Capanna Margherita by Durig and Zuntz, and their results were pubUshed in 1904.^ The dry gas-meter and the mouthpiece, valves, and nose-clamp were employed. The resting experiments were made every morning with the subject in bed, except when a study was made of the after-effect of work. Some of the resting vahies used for experiments with Zuntz as a subject were taken from the published results of a previous research. Both the resting and walking experi- ments were without food. After the basal values obtained with the subject lying in bed had been deducted, the energy per horizontal kilo- grammeter was computed and is given in table 1, page 22. Observations of Caspari, 1905. — In his study of vegetarianism, Cas- pari^ had an opportunity of studying the metaboUsm of two compet- itors in a walking-match from Dresden to Berhn, one of whom was a vegetarian and the other subsisted on a mixed diet. The experiments were made upon a treadmill in Zuntz's laboratory and presumably the Zuntz technique was carried out in all details. Striking differences in the gait of the two subjects were noted. One of the subjects, K. M., won the match, proving himself a particularly efficient walker. The basal values were obtained while the subject was lying in absolute rest. The walking experiments were made with food, and probably the rest experiments also, although no statement is given in regard to the food with the rest experiments. After the basal values had been deducted from the values found with the treadmill, the energy per horizontal kilogrammeter was computed and is given in table 1, page 24. Observations of Zuntz, Loewy, Mvller, and Caspari, 1906. — In con- nection with the classical research on the physiology of man in high 'Zuntz and Schumburg, Studien zu einer Physiologie des Marsches. Hiraohwald, Berlin, 1901. *Durig and Zuntz, Travaux de I'annfee 1903, Laboratoire soientifique international du Monte Rosa, Turin, 1904, p. 65; also Archiv f. Anat. u. Phyeiol., Physiol. Abth., 1904, Suppbd., p. 417. "Caspari, Archiv f . d. ges. Physiol., 1905, 109, p. 473. 18 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. altitudes, which was made m the Alps, Zuntz, Loewy, Muller, and Caspari^ carried out a series of resting and horizontal walking experi- ments in Berlin. The resting experiments were all made with the sub- ject lying in bed in the morning before taking food. The experiments on horizontal walking were presumably without food and hence it is permissible to deduct the basal values directly to compute the energy per horizontal kilogrammeter. (See table 1, page 24.) The large wet gas-meter was doubtless employed in the Berlin experiments, together with the rest of the Zuntz technique. The walking experiments on the treadmill were as follows: With Waldenburg, 9; with Kolmer, 4; with Caspari, 5; with Muller, 6. The especially high values found with Kolmer in this research are commented upon at length by Durig.^ Observations ofDurig, 1906. — In connection with an expedition in the Alps, Durig and his wife were the subjects of a number of resting and horizontal walking experiments.^ Both the basal and walking experi- ments were made with the subject in the post-absorptive condition, that is, several hours after the taking of food. The dry gas-meter was used and the Zuntz technique. 18 resting experiments and 8 horizontal walking experiments were made with Durig and 11 resting experiments and 6 walking experiments with Frau Durig, all of these being in the mountains. In addition the average value is given for a number of horizontal walking experiments with Frau Durig in Vienna. Special comment should be made of the extraordinarily heavy appa- ratus — 16.5 kilograms — carried by Frau Durig, who weighed but 45.7 kilograms. The basal values have been deducted from the values obtained in the walking experiments and the energy per horizontal kilogrammeter has been computed; this is given in table 1, page 24. Observations of Durig, Kolmer, Rainer, Reichel, and Campari, 1909. — The extraordinary care which characterizes all of the researches of Durig is manifest in his classical contributions on the physiology of man in the Alps; of particular value are his keen criticisms and summation of earlier research.* Indeed, nowhere do we find so sharp a recognition of aU of the fundamental tenets of careful experimentation in gaseous metabohsm, and particularly in the physiology of walking with special reference to the physiology of man in high altitudes, as in this series of contributions from Durig's laboratory. The observations were all made with the portable dry gas-meter which, with the equipment, weighed 11 kilograms. The experiments were made in the morning, either without food or after taking a cup of weak tea. 32 observations were made on a level road 200 meters long in Vienna and 12 observa- 'Zuntz, Loewy, Muller, and Caspari, Hoheuklima und Bergwanderungen in ihrer Wirkung auf den Mensohen, 1 Aufl., Berlin, 1906. ^Durig, Denkschriften d. math.-natur. Kl. d. kaiserl. Akad. der Wisaensoh., 1909, 86, pp. 253 and 254. »D>irig,.Arohiv f. d. gea. Physiol., 1906, 113, p. 213. *Durig, Kolmer, Rainer, Reichel, and Caspari, Denkachrift. d. math.-natur. Kl. d. kaiserl. \.kad. der Wiasenaoh., 1909, 86, p. 242. PREVIOUS RESEARCHES ON GASEOUS EXCHANGE. 19 tions were made on the Semmering. Durig discusses carefully all of the factors which should be taken into consideration in experiments of this nature and in the interpretation of their results. Of especial inter- est is his discussion of the suitable base-line to be deducted from the values found in the walking experiments. It should be noted that he adheres to the basal values found with the subject lying, as he considers the evidence in regard to the standing position wholly inadequate. His final computations for the energy required per horizontal kilo- grammeter are given in table 1, page 24. Observations of Amar, 1910. — In a research on the metaboUsm of Arabs/ Amar made numerous walking experiments with 15 subjects, who ranged in weight from 59 to 78 kilograms. The experiments were not made with the subject in the post-absorptive condition and loads weighing from 45 to 60 kilograms were carried. The Thiry metaUic valve and a dry gas-meter were used. The author gives a very inade- quate description of his technique. It appears, however, that Amar probably calculated the heat output of his subjects from the energy of the food and obtained an approximate control of his results by com- puting the heat output from the actual determinations of the oxygen intake and the calorific value of oxygen. Averages computed from the individual figures given by the author are included in table 1, page 24. Observations of Amar, 1911. — In a series of walking experiments reported in 1911, Amar^ used Thiry respiration valves which were con- nected with a dry gas-meter. The apparatus was placed upon a table, which was pushed behind the subject as he walked, and samples were withdrawn and analyzed, apparently for oxygen alone. The experi- ments were made in the morning, 10 or 12 hours after the taking of food. Control experiments were made with the subject standing and also with the subject sitting, Amar noting an increase in the metabo- Hsm during standing as compared with sitting. His computations are based upon the increase in energy expenditure while walking above that in a state of repose. Especial attention is given in his discussion to the variations in the load and the economic value of the rate of walking. The experiments have been criticized by Brezina and Kol- mer,' who protest against the lack of information as to the details of the experiments and the technique and state that the barometric pressure, temperature, and carbon-dioxide output were apparently not taken into consideration. An abstract of Amar's results is given in table 1 , page 24. Observations of Brezina and Kolmer, 1912. — ^A series of walking exper- iments was made by Brezina and Kolmer in Durig's laboratory in Vienna.* In one set of experiments the subject carried on his back a 'Amar, Le rendement de la machine humaine, Paris, 1910; also, Le moteur humain, Paris, 1914, pp. 493 and 494. 'Amar, Joiiru. de Phyaiol. et de Pathol, gen., 1911, 13, p. 212. 'Brezina and Kolmer, Biochem. Zeitschr., 1912, 38, p. 132. ^Ibid, p. 129. 20 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. dry gas-meter and the ordinary equipment was used, including the mouthpiece and valves. This, with additional weights, made a load of 11 Idlograms. In one set of the experiments in which heavier loads were used, the gas-meter was carried by the subject, but in later experi- ments the meter was carried by an assistant who walked behind the sub- ject. All of the experiments were made on a path near the laboratory and after the subject had taken a cup of tea with sugar. The authors maintain that the last heavy meal was taken 12 hours previous to the experiment, and that the influence of the ingestion of food was there- fore ehminated. In computing the results, a basal value of 1,083 gram- calories per minute was assumed for the subject. The authors point out that this is, to a certain degree, arbitrary. It should be noted that some of the experiments are considered less accurate than the others, owing to difficulties with the gas-meter. Special emphasis is laid upon variations in load and in the rate of walking. In a subsequent paper Brezina and ReicheP review the earher paper of Brezina and Kolmer and make certain corrections in the values. These corrected values are used by us in our summation of the results of Brezina and Kolmer in table 1, pages 24 and 26. Observations of Douglas, Haldane, Henderson, and Schneider, 191S. — In connection with researches carried out on Pike's Peak by Douglas, Haldane, Henderson, and Schneider, a number of observations were made on Douglas, both at Oxford and on Pike's Peak.^ The special form of respiration apparatus devised by Douglas was used, consisting of a mouthpiece and a pair of valves connected by tubing with a large rubber bag carried on the back of the subject. The experiments were made with the subject hpng in bed, in a standing position, and walking on a horizontal plane at the rate of 2 to 5 miles an hour. The basal values were assumed to be those measured when the subject was stand- iag quiet with the muscles relaxed. Apparently some of the lying experiments were made with the subject in a post-absorptive condition, while the standing and walking experiments were made after food had been taken. No attention was paid to the character or amounts of food eaten and the possible influence upon the measurements, as the observa- tions were not made primarily for the purpose of studying the absolute metabolism. A summary of the results is given in table 1, page 26. Observations of Brezina and Kolmer, 1914- — To carry out the plan conceived by Durig of studying the metabohsm during wa lkin g under every possible condition and to complete their own earher experiments, in which they studied the influence of speed and load during walking on a horizontal path, Brezina and Kohner made a second research, in which they studied the influence of the work of ascent upon the metabo- ^Brezina and Reichel, Biochem. Zeitschr., 1914, 63, p. 170. ^Douglas, Haldane, Henderson, and Schneider, Phil. Trans. Roy. Soc. London, 1913, ser B 203, p. 185. PREVIOXTS RESEARCHES ON GASEOUS EXCHANGE. 21 lism.^ Numerous experiments were carried out in the Hoschschule f. Bodenkultur in Vienna, with the treadmill both horizontal and incUned, and with and without load. Presumably the wet gas-meter was used. In the load experiments the subject carried a knapsack with weights in it. In the 15 experiments with the treadmill horizontal, the rate of walking ranged from 30.1 to 55.2 meters per minute, the speed being low to correspond with the rate of walking in the experi- ments with the treadmill inclined. Brezina himself was the subject. The experiments were made in the forenoon, 1| hours after the taking of a cup of sweetened tea. The base-line used was probably the values obtained with the subject lying down, and the Zuntz method for deter- mining the respiratory metabohsm was employed. The caorlies required per horizontal kilogrammeter are given in table 1, page 26. Observations of Galeotti, Barkan, Giuliani, Higgins, Signorelli, and Viale, 1914- — On an expedition to Col d'Olen on Monte Rosa in 1913 Galeotti, Barkan, Giuhani, Higgins, Signorelli, and Viale made a number of observations on the gaseous metabohsm of four individuals while the subjects were walking on a level.^ Since basal values were obtained in only one instance, it was necessary to assume these for the other subjects. The Douglas bag and the Siebe-Gorman valves were used, except in one experiment when the Tissot valves were substituted. Strict attention was given to the use of food, certain tests being specifi- cally made after breakfast. The experiments which are reported here, however, were made with the subjects in the post-absorptive condition. An abstract of the values for the horizontal kilogrammeter is given in table 1, page 26. SUMMARY OF RESULTS OF PREVIOUS OBSERVATIONS. In an attempt to arrange in chronological order a mathematical expression of the values on a comparable basis, we have gathered together all of the hterature available on horizontal walking and sum- marized the results of previous researches in table 1. In some of the work, particularly in the earher observations, certain assumptions were essential. These assumptions, which were based upon careful analyses of all of the experiments, upon deductions drawn from the experience of this laboratory in metabolism experiments, and upon known and recog- nized errors in technique are, we believe, justifiable and in all proba- bility are not greatly in error. The table gives the name of the author and the date of reporting the results of the research; the name or initials of the subjects of the walking experiments; the conditions under which the experiments were made; the method of measuring the respiratory exchange; the kind of walking, i. e., in a room, out of doors, or on a 'Brezina and Kolmer, Biochem. Zeitschr., 1914, 65, p. 16. 'Galeotti, Barkan, Giuliani, Higgins, Signorelli, and Viale, Reale Accademia dei Lincei, Rome, 1914, and Arch. d. Pisiol., 1914, 12, p. 277. 22 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. Table 1.— Historical summary of walking Author, date of publica- tion, and subject. Smith, 1859: Smith... Gruber, 1891: Gruber . . Katzenstein, 1891: Kohansky .... Krzywy Wellnitz Zimm Sond^n and Tigerstedt, 1895: F. A. W G.J L. B F Condition during walking. 6 hours after food (?). 5 to 6 hrs. after food. >With food (?) . . Schumburg and Zuntz, 1896: N. Zuntz Schumburg N. Zuntz A. Loewy, J. Loewy, and L. Zuntz, 1897: A. Loewy' J. Loewy' L. Zuntz' A. Loewy' J. Loewy' L. Zuntz' L. Zuntz, 1899: L. Zuntz Frentzel and Reach, 1901: Frentzel Reach With food . Method of measuring respura- tory exchange. Mask for resting; mouth- piece for walking; spirome- ter carried. Tube in mouth; absorption apparatus in knapsack car- ried on breast. Zuntz and Lehmann . Respiration chamber . Not stated . .do. Zuntz and Schumburg, 1901: B P Durig and Zuntz, 1904 : Durig*' Durig' N. Zuntz' Durig" N. Zuntz" A few hours after ■ light, liquid breakfast. Moderate break- fast. •With food . After a light breakfast. Without food (?). Zuntz and Geppert; dry gas- meter on back. Kind of walking. Indoors, 11 yds. in each direc- tion. In room, back and forth. Zuntz treadmill. In chamber. ...do. ...do. Zuntz; wet gas-meter near treadmill. Zuntz . Zuntz and Geppert. Zuntz and Schumburg; dry gas-meter on back. do Zuntz treadmill. ....do In room. Zuntz treadmiU. . Small plateau in front of hoteL Zimtz treadmill. .do. .do. Free path, out- doors (?). In hut 'The summarized data in this table are based on published material cited on pp. 11 to 21. 'Calculated from the data available, assuming a respiratory quotient of 0.85 and 1,609.35 meters per mile. 'Computed with several necessary assvraiptions, i. e., length of step, 0.680 meter; number of steps, 80 per minute for second figure as well as for the first; respiratory quotient, 0.85. 'Values computed by Duiig (Denkschrift. d. math.-natur. Kl. Akad. Wissensch., 1909, 86, p. 242), including correction for angle of ascent as indicated by Katzenstein. 'All heat values computed by Durig (loc. cit., p. 256), assuming respiratory quotient of 0.85. Corresponding figures assuming a quotient of 0.7.«i T»pro- n .i.'io n .q'ra n kik ^^a n quo StTMMAKY OF EESULTS OF PREVIOUS OBSERVATIONS. experiments on horizontal plane.^ ' 23 Resting value. Average weight moved. Distance per minute. Range. Average. Heat computed per hori- zontal kilogrammeter. Range. Average. Sitting, 6 hrs. after food Sitting, 5 to 6 hrs. after food . Standing, leaning against tread- miU; with food (?). Sitting, with food . Sitting . Lying, without food . do kilos. 92.08 72.00 meters. "Ruhewert" Sitting .....do Lying, without food "Ruhewert" Lying, with food . . . Lying, after light breakfast . . . 62.73 78.25 70.09 68.77 80.00 88.20 80.00 72.60 81.10 80.00 72.60 81.10 80.30 72.94 87.10 66.40 67.92 72.93 f"91.80 \ "73.80 ["82.50 /"74.90 1 "78.70 56.00- 92.00 51.00- 75.00 58.00- 71.00 64.00- 66.00 58.70- 62.40 62.20- 67.90 47.80- 52.40 61.60- 62.40 59.00- 62.80 64.50- 59.20 61.80- 70.21 59.40- 62.04 63.21- 67.33 r 53.45- 59.35 92.41-103.40 135.74-145.53 57.80- 80.15 31.39- 39.91 60.28- 66.88 31.58- 37.23 70.66- 89.21 68.21- 82.38 95.70-111.50 57.34^ 96.42 78.66- 80.66 66.26- 91.69 60.03- 66.67 meters. ' 53.64 , 80.46 ■ 54.40 54.40 74.48 61.80 63.50 66.05 60.60 60.10 65.10 32.00 61.20 42.00 50.10 62.00 60.90 66.40 67.10 60.54 66.24 66.76 98.67 140.11 66.94 35.92 63.95 34.68 76.60 73.50 99.60 85.56 79.93 84.21 62.86 gm.-cals. 0.506-0.506 .603- .427 «.606- .803 .636- .706 .511- .560 .446- .636 .661- .724 .769- .877 .548- .712 '.538- .561 9.621- .693 n. 015-1. 168 .442- .626 .511- .620 .507- .696 .464^ .676 .474r- .571 .448- .681 .574- .714 .633- .699 .742- .796 gm.-cals. ^0.407 ^468 3.360 ^356 *.526 •.786 «.564 '426 '.606 '.337 '.465 '.324 '.678 '.616 '.718 .670 .635 .570 .681 .821 .616 ».664 9.663 91.072 .627 .660 .553 .558 19.627 '9.609 .527 .584 .663 .668 .774 •Values given by Durig (loc. cit., pp. 248 and 261). Figures for treadmill experiments were corrected by Durig for angle of ascent. 'Experiments conducted at Berlin. 'Experiments conducted at Col d'Olen. 9A11 heat values calculated by Durig (loc. cit., p. 270). '•Corrected by authors for slight elevation of treadmill. Values for load experiments may be found on pages 247, 265, and 278 of the original publication, but not corrected for angle of ascent "Experiments conducted at Vienna. "Not given by authors; these weights were accordingly computed from other data which had been obtained by means of the body weight. ''Experiments conducted at Capanna Margherita. 24 ENERGY TKANSFORMATIONS DURING HORIZONTAL WALKING. Table 1. — Historical summary of walking Author, date of publica- tion, and subject. Condition during walking. Method of measuring respira- tory exchange. Kind of walking. Caspari, 1905: K. M With food . Zuntz . Zuntz treadmill. J. B. .do. .do. .do. Zuntz, Loewy, Miiller, and Caspari, 1906: Waldenburg Kolmer Muller Caspari Durig, 1906: Frau D.2 Frau D.' Durig' Durig,' Kolmer, Rainer, Reichel, and Caspari, 1909: Durig* [without food . Without food . Without food, or after cup of weak, sweet- ened tea. Zuntz andGeppert; presum- ably wet gas-meter. Zuntz and Schumburg; dry gas-meter on back. 'Zuntz ; dry gaa-meter on back . Zuntz treadmill. Free path . Outdoors . Durig' . . Kolmer' . Rainer^ . Reichel'' . Reichel* . .do. .do. .do. .do. .do. .do. .do. .do. .do. .do. .do. .do. .do. .do. do. Amar, 1910: Arabs . . With food. Amar, 1911: S. L. . . Brezina and 1912: Brezina . Kolmer,' 10 to 12 hrs. after food. 12 hrs. after food, or at least 2 hrs. after cup of sweetened tea. Chauveau valve; dry gas- meter. Chauveau valve; dry gas- meter; apparatus on table moved with subject. Dry gas-meter carried by assistant in first aeries of experiments, and in last 3 series; in 2d and 3d series, subject carried meter. Circular path outdoors. In room, 11 me- ters in each direction. Level path out- ■ doors, involv- ing turns. •Average values as corrected by the authors. 'Experiments conducted at Vienna. 'Experiments conducted at Spomer Alps. •Not given by the author, but computed from other data in which the body weight was involved There is a possible error of about 2 kilograms in this weight. 'The data from these experiments have been grouped for this summary, with special attention to place and rate of waUdng. 'Experiments conducted at Semmering. SUMMARY OF RESULTS OF PREVIOUS OBSERVATIONS. 25 experiments on horizontal plane — Continued. Resting value. Average weight moved. Distance per minute. Range. Average. Heat computed per hori- zontal Mlogrammeter. Range. Average. Lying. . do. Lying, without food . Lying, without food. Lying. .do. .do. .do. .do. .do. 'Repos" with food. Standing or sitting. I " Erhaltungsumsatz ' I gram-calories. of 1083 kilos. meters. 63.18 134.90-142.90 63.47 172.00-182.60 65.35 122.20-135.70 74.08 44.14- 78.85 85.10 37.00-48.45 87.40 73.29- 86.52 81.95 67.47- 82.69 *57.50 63.38 66.07- 79.63 79.25 88.49- 99.63 76.20 72.30-102.30 116.6 126.0 76.10 1 141.8 152.5 76.30 100.70-103.20 76.50 105.40-110.30 49.2 94.20 -I 62.40- 69.20 100.00-105.20 47.2 75.40 <, jj^g g 129.5 100.30 59.30-100.80 95.45 . 95.45 113.80 .134.10 ■ 66.00 73.30 70.10 70.90 69.90 70.50 70.50 93.20- 99.20 103.50-103.90 72.00- 90.00 72.00- 80.40 32.40- 89.83 30.33- 78.08 46.10- 72.20 81.80- 98.60 103.10-115.80 120.10-131.20 140.60-141.00 meters. 139.40 177.30 131.70 60.20 43.16 81.17 76.76 65.00 71.90 95.44 90.10 116.60 126.00 141.80 152.50 102.20 107.90 49.20 66.30 102.50 47.20 87.20 115.80 129.50 88.20 96.30 103.70 79.20 74.80 59.43 55.06 62.40 88.90 111.20 125.20 140.80 gm.-ecUs. .952-1.018 1 . 154-1 . 184 .906-1.020 gm.-caU. .979 1.169 .972 1.636 1.845 1.613 1.643 .629- .691 .620- .660 .517- .559 0.628 .735 .854 1.023 0.566-0.587 .622- .636 0.542 0.535-0.591 .635- .659 0.538 0.533-0.584 0.772 .954 0.510-0.583 .563- .591 .641- .659 .257- .402 .310- .335 .315- .577 .365- .535 4844- .6033 5615- .5985 7266- .7684 8172- .8777 .9347-1.0070 .604 .668 .641 .539 .628 .735 .854 1.023 0.573 .629 .542 .562 .648 .538 .567 .772 .954 .548 .573 .650 '.308 '.323 8.410 8.422 '.534 .574 .743 .846 .971 'Averages computed from results given by the author for 15 different subjects. The results are grouped for the two loads carried. 'AU heat values computed from the oxygen increment during walking as given by the author and with an assumed respiratory quotient of 0.85. Assuming a quotient of 0.75, the results for the two series are: range 0.307 to 0.562; average, 0.400; range 0.356 to 0.521, average 0.411. •The results in each of these series have been grouped for this summary according to the rates of walking, the Umit of the first group being set at about 80 meters in agreement with the treat- ment given the material in the original publications. The values for distance and calories are given as corrected by Brezina and Reichel. 26 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. Table 1. — HisUrrical summary of walking Author, date of publican tion, and subject. Condition during walking. Method of measuring respira- tory exchange. Kind of walking Brezina and Kolmer,' 1912 (continued): Brezina Do. 12 hrs. after food, or at least 2 hrs. after cup of sweetened tea. .do. Dry gas-meter carried by assistant in first series of experiments, and in last 3 series; in 2d and 3d series, subject carried meter. .do. Do. .do. .do. Do. .do. -do. Do. Douglas, Haldane, Hen- derson and Schneider, 1913: Douglas Do. Do. Brezina and Kolmer, 1914: Brezina Galeotti, Barkan, Giu- liani, Higgins, Sig- norelli, and Viale, 1914: Barkan Higgins Signorelli Viale .do. .do. With food Dougli .do. .do. lyi hrs. after cup of sweetened > Without food . .do. Level path out- doors, involv- ing turns. .do. .do. .do. .do. .do. Zuntz . Douglas bag ....do ....do ....do f Room in labora- tory at Ox- ford; circular path. Railway track. Pike's Peak, back and forth. {Grass track, Ox- ford, back and .forth. Zuntz treadmill. Half indoors, half outdoors, with a turn (all indoors in bad weather.) 'The results in each of these series have been grouped for this summary according to the rates of walking, the limit of the first group being set at about 80 meters in agreement with the treat- ment given the material in the original publications. The values for distance and calories are given as corrected by Brezina and Beichel. ^The body-weights in all the series with Douglas have been computed from the distance and the total oxygen and the oxygen per kilogram per meter as given by the authors. The subject's naked weight at Oxford was about 63.5 kilograms; at Pike's Peak, 60.8 kilograms. StTMMAKY OF EESTJLTS OF PEEVIOUS OBSERVATIONS. 27 experiments on horizontal plane — Continued. Resting value. Average weight moved. Distance per minute. Range. Average, Heat computed per hori- zontal kilogrammeter. Range. Average. "Erhaltungsumsatz" of 1083 gram-calories. .do. .do. .do. .do. ^Standing, with food . .do. ....do. 'Ruhewert". Lying, without food . 103.50 103.60 103.20 .103.00 112.60 113.00 113.00 ri23.00 \123.00 "72.45 72.77 73.76 72.73 72.78 270.71 70.48 70.70 70.88 70.89 272.94 72.59 73.77 74.36 73.42 meters. 37.40- 73.30 78.20- 84.20 92.80- 94.50 105.7 118.1 125.60-129.00 43.50- 77.70 84.60- 94.90 104.70-105.10 115.80-115.80 32.80- 74.00 86.00- 90.60 100.0 111.4 37.00- 76.70 78.10- 82.20 89.90- 94.70 48.60- 68.90 90.9 '52.03- 54.18 77.25- 82.88 102.73-107.83 119.90-121.77 134.65-137.87 351.77- 54.98 77.52- 81.55 102.73-111.32 112.67-116.15 132.77-132.77 '53.12- 54.98 79.13- 84.50 105.42-111.58 120.43-126.33 130.63-139.48 31.00- 51.70 51.40- 55.20 49.40- 52.40 30.1 meters. 53.10 81.50 93.40 105.70 118.10 127.30 62.00 91.00 104.90 115.80 54.80 88.30 100.00 111.40 55.00 80.20 92.30 58.80 90.90 '53.38 81.00 105.95 120.97 136.26 '53 . 64 79.66 108.09 114.53 132.77 '53.91 82.88 108.36 123.38 136.26 46.10 53.80 51.10 30.10 86.70 65.00 86.70 86.70 gm.-cals. 0.4709-0.5278 .5040- .5411 .6088- .6213 0.7107 .9281 0.9626-0.9720 .4502- .5716 .5651- .5933 .7232- .7982 .9121- .9122 .4766- .5883 .5439- .6436 0.7256 .9143 0.5433-0.6391 .5450- .6778 .8143- .8796 .5858- .5884 0.7747 '0.366 .440 .523 .638 .889 '.524 .568 .713 .768 .919 '.547 .588 ■ .704 .770 .978 ■ -0.448 - .490 - .549 - .682 - .907 - .607 - .634 - .862 - .897 - .964 - .609 - .600 - .873 -1.009 -1.126 .50 - .56 .48 - .56 .42 - .55 0.51 0.570 -0.395 .374 - .398 gm.-cals. 1.489 .520 .616 .711 .928 .967 1.506 .581 .761 .912 1.529 .594 .726 .914 '.577 .611 .847 1.587 .775 '.402 .460 .531 .656 .898 '.562 .600 .789 .833 .942 '.571 .595 .776 .897 1.066 <0.53 * .50 * .50 ' .51 «0.448 ' .550 » .386 » .375 'It was necessary to convert the distances from miles to meters and to compute the heat from the oxygen consumption. 'Averages of individual heat values given for the groups in the original publication. 'In computing all heat values for Barkan and Viale, the value for lying without food was assumed to be represented by 255 c.c. of oxygen; for Signorelli, 225 c.c. of oxygen. In one experiment with Higgins, a corresponding value of 203 c.c. of carbon dioxide and of 265 c.c. of oxygen was obtained. The weight of the clothing and the Douglas bag was about 7 kilograms in each case. 28 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. treadmill; the basal or resting value for the individual subjects; the total weight of the material moved in a horizontal direction, including the weight of the body, clothing, apparatus, and any supplementary weight which the subject may have carried; the distance walked per minute; and finally, the calories per horizontal kUogrammeter. The speed of the walking, which is shown by the distance walked per minute, is a factor of considerable importance; the range and average are therefore both given. Whenever there is a great difiference in the speed for the same subject, a special line is assigned to it. Wide varia- tions are shown in the average distance walked per minute in the several researches, the figures ranging from 32 meters in the Sonden and Tigerstedt experiments pubUshed in 1894 to 177.3 meters in an experi- ment by Caspari reported in 1905. As a result of a careful analysis of the earlier literature of Durig and of Zuntz and his co-workers, the unit adopted for expressing the work of forward progression is the calories required to move 1 kilogram 1 meter or, as we shall here designate it, the calories per horizontal kilo- grammeter. The range and average are both given for these values, but the average values have here the greater significance, and it is the last colunm of the table which we shall particularly consider. Examin- ing these data, we find that they range from 0.308 gram-calorie in the series of experiments made by Amar and reported in 1910 to 1.169 gram-calories in the experiment by Caspari pubUshed in 1905. Con- sidering the varied technique, there is, on the whole, a rather remark- able agreement of the values found for the movement of 1 kilogram of body-weight per meter in a horizontal direction, particularly when we consider those values obtained at moderate walking speed. Comparing the data for the average distance walked per minute and those for the average calories per horizontal kilogrammeter, we find that the rate of walking has a considerable influence upon the results and that the higher values are invariably found with the greater speeds, although the reverse is by no means true. In general, when the rate of walking does not exceed 80 to 90 meters per minute, the values lie between 0.3 and 0.7 gram-calorie, with a distinct tendency for them to approach 0.55 gram-calorie. This value has been especially con- sidered by Durig in his admirable review of the literature of this subject. It is clear, then, that we have here to do with an intimate relationship between the rate of walking and the expenditure of energy required to move 1 kilogram 1 meter in a horizontal direction, and that the superimposed load has relatively little influence upon the results. METHODS AND APPARATUS. 29 METHODS AND APPARATUS FOR STUDIES OF MUSCULAR WORK. Judging from the results as published in the earlier Uterature and summarized in the preceding section, there is apparently httle choice as to the various methods of study. A large majority of the observa- tions have been made with the Zuntz respiration apparatus or some of its modifications; it is obvious that any criticisms applying to this apparatus affect greatly the averages of the values obtained with it. While the construction of the Zuntz apparatus and the various types of valves, particularly the later form of valve used by Durig, is such as to ehminate resistance as far as possible, nevertheless the employment of this type of apparatus for studying muscular work is distinctly open to criticism. The use of a wet gas-meter of the size ordinarily employed with the Zuntz respiration apparatus may be seriously objected to for muscular- work experiments, since in these experiments the volumes of expired air may exceed 80 or 90 hters per minute. The caUbration of such a meter at the rate of ventilation used in severe-work experiments is by no means a simple matter, the ordinary method being to allow the water in a 10-Uter spirometer to flow out and draw the air in after it. On the ordinary Elster gas-meter 10 hters correspond to one or at most two revolutions of the drum, and the possibihties of error at the begin- ning and the end in tests of this kind are obvious. Indeed, both Durig and Zuntz have recognized this difficulty and, to avoid errors in the measurement of the large volume of air expired from the lungs, have "made experiments with a very large gas-meter formerly used in studies with horses. If the value obtained for the volume of air be too small (and the error, if any, would be in this direction), obviously that for the total metaboUsm will likewise be too small, and hence the values com- puted for the horizontal kilogrammeter will be too low. The importance of knowing the energy requirement of the body for direct forward progression is so great as to justify further observations upon it, particularly if an apparatus is used in which an entirely dif- ferent principle is employed for measuring the metabohsm. One of the main lines of research planned for the Nutrition Laboratory is a study of muscular work in its various phases. Obviously innumerable problems, of both physiological and economic importance, can be studied with suitable apparatus. A special calorimeter has been constructed in this laboratory for research on muscular work, but it is not yet ready for actual use. Prior to experimentation with this calorimeter, it seemed important to make a number of preliminary studies of the metabolism during muscular work with the universal respiration apparatus. This apparatus had previously been used in a study of the metabohsm incidental to riding a bicycle ergometer, in 30 ENEEGY TRANSFORMATIONS DURING HORIZONTAL WALKING. which it was shown that it was capable of measuring the metabolism with an oxygen consumption as high as 3 liters per minute.'^ Careful control tests of the apparatus have proved conclusively that, so far as the mechanical construction is concerned, the measurement of the carbon- dioxide production and the oxygen consumption is accurate. As a part of the plan for studying the metaboUsm during the various phases of muscular work, a special type of treadmill was devised and constructed in this laboratory several years previous to the present research. Certain features of this treadmill, which will be described subsequently, make it especially apphcable for use in experiments with men. Having, therefore, an exceptionally satisfactory treadmill and a respiration apparatus capable of measuring the metabohsm incidental to severe muscular work, it was possible, in 1913-14, to make a study of the metaboUsm of a subject while he was walking on a level. As soon as the respiration calorimeter for muscular work is ready for use, the treadmill wiU be placed inside this apparatus and certain of the experiments repeated. The indirect calorimetry computed from the gaseous exchange as measured by the respiration apparatus can thus be controlled by direct determinations of the energy output. This control is of particular importance, since the use of any form of respiration appUance, such as a mouthpiece or nosepieces, is open to legitimate criticism in that the use of such artificial methods of breath- ing, particularly with a very great ventilation of the lungs, may lead to a disturbance in the mechanics of ventilation, thus causing a distur- bance in the respiratory exchange. Although the universal respiration apparatus has been carefully controlled as to the measurement of both carbon dioxide and oxygen, nevertheless if there is a disturbance in the mechanics of ventilation, as, for instance, an over-ventilation of the lungs, there may be a considerable amount of carbon dioxide exhaled that was not simultaneously produced. The retention of carbon dioxide by altered respiratory mechanics is also not improbable. By means of the studies with the respiration calorimeter, therefore, it will be possible not only to determine the metabohsm by direct calorimetry, but also to obtain the gaseous exchange with free breathing without the use of either mouth or nose apphance. It should furthermore be borne in mind that while the method of computing the calorimetry from the gaseous exchange has, according to our experience up to the present time, given highly satisfactory results with muscular repose, there is always the possibihty of a dis- turbance in the character of the metabohsm during severe muscular work and a consequent disturbance in the relationship between the gaseous exchange and total heat production. It is thus seen that the final tests with the respiration calorimeter are essential for a complete understanding of this problem. On the other hand, the previous ^Benedict and Cathcart, Carnegie Inst. Waah. Pub. No. 187, 1913. DESCRIPTION OF APPARATUS. 31 research with the professional bicycle rider, in which the universal res- piration apparatus was employed, was so successful and the problem of the metabolism during the work of forward progression is so important that, pending the completion and testing of the special calorimeter, the study of this problem with the universal respiration apparatus and the treadmill has seemed entirely justifiable. DESCRIPTION OF APPARATUS USED IN THIS RESEARCH. UNIVERSAL RESPIRATION APPARATUS. The researches of Zuntz and his co-workers have shown that in walk- ing at a somewhat high rate of speed and carrying a load, the amount of muscular exertion required, even on a level path, is very considerable, approximating that required for bicycle riding under severe stress. Obviously, therefore, any respiration apparatus used in a study of the metaboUsm under these conditions must be capable of accurately measuring a maximum oxygen consumption per minute of 3,000 c.c. and a maximum carbon-dioxide production of 2,500 c.c. Fortunately the modified form of the universal respiration apparatus employed by Cathcart had already demonstrated its abihty to fulfill these conditions. The form of apparatus used in this later research was therefore essen- tially that employed for the earher study.^ A general view of the apparatus with its relations to the subject and the accessory apparatus is shown in figure 1. Siace the completion of the research on the bicycle rider, various minor modifications have been made in the apparatus, chiefly with a view to f acihtating operation and contributing to its accuracy. Certain other changes were also necessary to adapt it to the type of experiment planned for this research, namely, walking upon a treadmill at varying rates of speed. These changes were considered of sufficient importance for us to give here a schematic outhne of the apparatus as actually employed in the research. (See figure 2.) As will be seen from the diagram in figure 2, by turning a 3-way valve N, the subject, breathing through a mouthpiece P, can be con- nected with the ventilating air-current, which is kept in motion by a rotary blower. The air leaving the blower is first passed through a glass WilUams bottle which serves as a safety trap against back-suction of acid. The air is next forced through two Wilhams bottles, each having a capacity of 2.5 liters and half filled with sulphuric acid, which removes the moistiu-e from the air. The ventilating current is then deflected by means of the 2-way valve V^ into either one of two 'Brezina and Kolmer in a recent paper from Durig's laboratory (Biochem. Zeitschr., 1914, 65, p. 33) have questioned the accuracy of the respiratory quotients obtained with this apparatus by Benedict and Cathcart in their study of a bicycle rider. Durig in a private communication says that the method used by Benedict and Cathcart for obtaining the respiratory quotients was accu- rate and that the criticisms of Brezina and Kolmer were founded upon an insufficient study of the results. 32 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. carbon-dioxide absorbing systems. Each series of absorbers include two bottles filled with moist soda lime for absorbing the carbon dioxide, and also a small WiUiams bottle, containing sulphuric acid to absorb the moisture given up to the dry air by the soda lime. The air, which is at this point free from carbon dioxide and water, passes through the 2-way valve 7^ and after being moistened in a small WiUiams bottle to a comfortable degree for respiration, it continues until it passes Fig. 1. — General view of apparatus used for walking experiments, with subject in position. A, absorption apparatus; B, treadmill; C and D, oxygen supply; E, step counter and accessory apparatus. by the opening of the tube connected with the mouthpiece P and into the tension equalizer. This tension equalizer is a large copper can, over the top of which is fitted a rubber bathing-cap. From this copper can the air finally returns to the intake side of the blower, oxygen being admitted as required from a cylinder of the gas. Thus we have a com- plete closed-circuit system, with the rubber bathing-cap acting as a tension equalizer to compensate for changes in the volume of air as the subject breathes in and out. DESCRIPTION OF APPARATUS. 33 The algebraic increase in weight of the soda-Ume bottles and the following Williams bottle in the carbon-dioxide absorbing system gives the weight of the carbon dioxide absorbed during the experimental period. The volume of oxygen introduced may be determined from the loss in weight of a small cylinder or, as indicated in figure 1, meas- ured by conducting it from a large cyUnder through a carefully calibrated gas-meter which is immersed in water to prevent sudden fluctuations in temperature. To prove the efficacy of the soda lime as an absorbent, a part of the air current may be deflected through two petcocks and passed through a solution of barium hydroxide in a small Erlenmeyer flask. The sUghtest trace of carbon dioxide will produce a turbidity in the solution. To ehminate the effect of the long dead space between the mouth- piece, P, and the main ventilating air-pipe, a supplementary pipe E is connected from a point near the mouthpiece to the main air-pipe. By this means the air-current may be deflected so that It will pass directly by the mouthpiece and within a few centimeters of the lips of the sub- ject. The rubber tubes connecting the subject with the ventilating system are of sufficient length and flexibihty to permit considerable lateral and vertical head-motion by the subject while walking, this beiag essential to his comfort. In actual experimenting, before the experimental period is begun, the 3-way valve N is so turned that the subject breathes room air. Then, at the end of a normal expiration, the 3-way valve is again turned, connecting the subject with the main ventilating air-current as seen in the diagram. Immediately afterwards the valve M (which has previ- ously been open, giving free passage of the air into the metal tee leading to the tension equahzer) is also turned, thus deflecting the air-current through the supplementary pipe E, so that it passes near the mouth of the subject. Before connecting the subject with the ventilating system, the motor is started to equalize pressure throughout the system and then stopped. Sufficient oxygen is next admitted to distend the rubber bathing-cap until a slight positive pressure is observed on the manometer. On starting the blower again, the rubber bathing-cap sinks somewhat, due to the compression of the air in forcing its way through the sulphuric acid and the various absorbing-vessels. At the first inspiration the rubber cap sinks down into the can, rising again with each expiration. As the oxygen is consumed, the height to which the rubber cap fills decreases and oxygen is admitted to keep the fluctuations of the bathing-cap to approximately the same range. At the conclusion of the experiment, the valve M is first turned and shortly afterwards the valve N is also turned, always at the end of a normal expiration. At this point the experimental period is concluded so far as the subject is concerned. The motor is next allowed to run for a minute or two to 34 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. sweep out completely the carbon dioxide in the system. When the motor is stopped sufficient oxygen is admitted to bring the tension to the original point; the volume of air in the system is then exactly the same as at the beginning. The carbon-dioxide absorbers and the accom- panying WilUams bottle are weighed and a reading taken of the amount of oxygen which has passed through the gas-meter, the data being used subsequently for computing the carbon dioxide produced and the oxygen consumed during the experimental period. H" ^^fr^iT HjSOt HjSO,. TRAP ^CnS-' METER Fig. 2. — Schematic outline of vmiversal respiration apparatus. The general course of the ventilating current is shown by arrows. P, mouthpiece; N, 3-way valve connecting subject with ventilating current; V- and Y^, 2-way valves for carbon- dioxide absorbing system; M, valve connecting with supplementary pipe E for eliminating the dead-air space; S, petcock connection for tambour and kymograph for recording the respiration. TREADMILL. In many of the earlier iavestigations for measuring the carbon- dioxide output and oxygen consumption of a walking man, the subject has been obhged to carry a respiration apparatus upon his back or drag behind him a long tube attached to the breathing appliance. In other studies the apparatus has been pushed along behind the subject by an assistant. Obviously none of these methods are suitable for experiments in which there is rapid movement. Furthermore, whUe we have no desire to minimize the value of the Zuntz apparatus — an apparatus which has contributed largely to our knowledge of the physiology of man, particularly for high altitudes — it should be said that the transportation of an unwieldy gas-meter and accessory appa- ratus by a subject is open to serious objection. It has been shown DESCRIPTION OF APPARATUS. 35 by Zuntz and his co-workers that a subject may be trained in a rela- tively short time to carry such an apparatus successfully; nevertheless its transportation is at best difficult, and the method is more fitted for studies of that type of walking in which loads would normally be carried upon the back. Both Zuntz and Durig have shown that they recognize this fact, as they have made studies in which the gas-meter was not carried by the subject, and the walking was done upon a treadmill. Several years previous to our research, Mr. E. H. Metcalf, then a member of the staff of the Nutrition Laboratory, designed a tread- mill and had it constructed under his supervision in the machine-shop of the Laboratory. This treadmill was used for the first time in the research carried out in 1913-14 and the fact that it satisfactorily sus- tained a severe test at this time testifies to the designing abihty of Mr. Metcalf and the constructive skill of Mr. W. E. CoUins, the mechan- ician of the Laboratory. A general view of the treadmill and its dispo- sition with regard to the other apparatus is shown in figure 1. A more detailed perspective drawing is given in figure 3. Fig. 3. — Treadmill designed by E. H. Metcalf. The endless leather belt travels over two wooden pulleys A and B, the actuating] mechanism being an electric motor. The tension on the belt may be adjusted by the bolt and nut C. A revolution counter is shown at R. The treadmill is provided with an endless belt, 60 cm. wide, 435 cm. long, and approximately 10 mm. thick. This belt travels over two wooden pulleys, A and B, having a width of 60 cm. and a diameter of 41 cm. The pulleys are supported on ball bearings at the ends of a wooden frame. On the rear pulley is attached a sprocket- wheel, which connects with the reducing-gear actuated by a one-half horsepower electric motor. The tension on the belt may be easily adjusted by means of the screw C, thus preventing the belt from sUp- ping. To support the belt between the pulleys and provide a surface for the man to walk upon, steel tubes, 46 in number, with an external diameter of 25 mm. and a length of 61 cm., are set into a steel frame- work, the distance between the centers of the tubes being 27 mm. These steel tubes are fitted at each end with annular steel ball bearings. 36 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. The detail of the bearmgs is given in figure 4, which shows the method of support on the angle frame of the treadmill. The thick belt thus rests upon a roUing, frictionless surface throughout the whole length of the space over which the man is to walk, providing a substantial, perfectly smooth path for the subject. It is impossible to feel the numerous steel rolls, as the space between them is so small. The leather belt gives a particularly satisfactory footing. The speed of the motor may be modified at will and tests may be made at rates of speed varying from less than 50 meters per minute, the equivalent of a very slow walk, to 150 meters per minute, or a rapid walk, if not, indeed, running. Since the driving of the treadmill belt is from the rear wheel, all tend- ency to slip is at this point. Consequently, by counting the revolutions of the forward wheel, where no sUp can possibly take place, we have an accurate measure of the total distance traveled. Upon the periphery of the forward wheel are attached two revolution counters (one on each side)^ which can be read intermittently. One runs continuously; the other may be connected or disconnected at will. In use, the subject walks upon the treadmill for several minutes before the experi- mental period commences. At the exact moment of beginning the measurements of the respiratory exchange, one of the revolution counters is put in action, so that the distance covered during the period is accurately known. It is thus unnecessary to assume con- stancy in the revolution of the motor or to apportion the distance walked during the period from the total measurement of the revolutions of the front wheel during the entire time the subject is walking. Considering the number of the various bearings and the size of the apparatus it produces much less noise and vibration than any treadmill that we have thus far seen. By means of adjusting screws at the for- ward end of the treadmill, which are not shown in figure 3, any desired elevation can be readily secured. In this research, however, the tread- mill was invariably used in a perfectly level position. At the date of writing (June 1915), after two years of use, the apparatus shows no signs of wear and gives most satisfactory service. ACCESSORY APPARATUS. The universal respiration apparatus and the treadmill provide accu- rate measurements of the total carbon-dioxide production and the oxygen consumption and of the distance walked by the subject. In addition to this, certain observations are important, particularly in studying the physiology of walking, and it was therefore necessary to have some means of recording accurately the respiration, the pulse, the number of steps taken by the subject, and the height to which the body was raised with each step. 'But one of these (R) is shown in figure 3. DESCRIPTION OF APPARATUS. 37 METHOD OF RECORDING THE RESPIRATION-RATE. The rise and fall of the rubber bag or tension equaUzer with each expiration and inspiration can be readily counted by an observer, thus giving an admirable index of the respiration. As a matter of fact, during the conduct of a severe-work experiment of this type, the other observations are so numerous that it is at best very difficult for an observer to concentrate his attention upon such counting. A tambour has therefore been connected to the petcock S (see fig. 2) which is moved by the difference in pressure existing in the connection between the mouthpiece and the tension equalizer. With each exhala- tion a shght pressure is exerted upon the tambour which records upon a kymograph drum. Subsequently an accurate count of the number of respirations per minute and for the entire experimental period may be made from the tambour record and that of a time marker. METHOD OF RECORDING THE PULSE-RATE. The important relationship between total metabohsm and the pulse- rate, which has been so frequently observed in this Laboratory^ made it desirable to secure records of the pulse-rate of the subject during the walking periods. The difficulties experienced in securing counts of the pulse-rate in the earher research with the bicycle rider have been freely commented upon in the publication giving the report of that study, and it was hoped that some graphic method could be found which would give accurate records of the pulse-rate of a walking man. Unfortunately such a method was not available at the beginning of our research, and counts were therefore made at the wrist or with a stethoscope. Even Yirith a stethoscope at some distance, as employed by Krogh,^ the noise of the treadmill was such as to make the counting of the pulse-rate extremely difficult. Variousforms of tambours^were also experimented with in an attempt to register the pulse graphically, but without result. Finally, by using body leads and the Bock-Thoma oscillograph or the Einthoven string galvanometer, we were able to secure graphic records of the pulse-rate of the subject while he was walking upon the treadmill. Great difficulty was experienced in such photographic registration with the dehcate oscillograph and string galvanometer, for not only were there leakages of the 220-volt current used for the motor, but also a static charge on the rapidly revolving belt. Various devices were used for obviating these difficulties, such as having the subject wear rubber-soled shoes and even rubbers over the shoes, but with Uttle success. Finally a metalUc brush was made of a piece of brass tubing, 69 cm. long, to which pieces of brass chain, 7 to 9 cm. long, were attached at intervals of approximately 1.5 cm. This brush was so attached to the treadmill that the ends of the chains lay upon the 'Krogh, Skand. Aichiv f. Physiol., 1913, 30, p. 375. ^Bowen, Contributions to Medical Research dedicated to Victor Clarence Vaughan, June 1903, p. 462, Ann Arbor, Mich. 38 ENERGY TRANSFORMATIONS DTTRING HORIZONTAL WALKING. surface of the leather belt. As one end of the chain brush was grounded to the brass tube, when the ends of the chain dragged over the surface of the revolving belt the stray electric currents were picked up and grounded. Satisfactory records could thus be obtained. Unfor- tunately the conditions of the experimental work in the Laboratory at that time prevented our freeing a galvanometer and oscillograph for the particular purpose of determining the pulse-rate; consequently records could not be regularly secured. Indeed, the few observations obtained were of so striking a character that we can not adequately discuss them in this monograph.^ STEP COUNTER. With a steady electric current, the rate of revolution of the treadmill per minute is presumably fairly constant and the length of the steps in ordinary walking is also doubtless approximately the same; neverthe- less it is important to show in these tests the exact number of steps taken, so as to indicate the distance per step and thus furnish an added factor for analyzing the mechanics and physiology of walking. To count the steps automatically was imperative, as even with several assistants an accurate record of the steps during exceedingly rapid Fig. 4. Fig. 5. Fig. 4. — Detail of ball bearing for steel tubes on the treadmill. Fig. 5.— Step counter. A pulley A is attached to the extended shaft of a revolution counter; a small cord passes over this pulley and is protected by a guard C. The shaft automat- ically stops at B and B'. walking would be very difficult to obtain. A small device was there- fore employed which counted automatically each vertical motion of the body produced in walking. To a belt about the waist of the subject was attached a cord leading over two pulleys on the ceihng to a small spring which maintained a tension on the cord. The general arrangement of this device is shown in figure 1 and the details in figures 5 and 6. At each step the sub- ject lifted the body a certain distance and lowered it again. Advan- tage was taken of this lifting and lowering of the body to raise and lower a small weight (.B,fig. 6) at the end of a cord which rested upon a pulley (A, fig. 5, and D, fig. 6) on the shaft of a revolution counter (see fig. 5). 'See pp. 85 and 92. DESCRIPTION OF APPARATUS. 39 Automatic stops, B and B', controlled the distance traveled by the pulley. At each upward movement of the body the cord was lowered and the counter pulled in one direction to the stop, the cord simply sUpping over the pulley if the movement continued. As the body returned to its ordinary position, the weight drew the counter back to the second stop. In this way each upward movement of the body was accurately recorded. Records obtained with this counter of the number of steps taken by one of our subjects in several walking experiments are given in table 2. These are compared with the number of steps counted for the same periods from kymograph records obtained by a method subsequently described.^ A large number of controls, which were made by counting the movements of the body from the kymograph records and by obser- vations of the counter, showed that this step-counter could be relied upon. Table 2. — Comparison of steps recorded by step-counter with those counted from kymograph records. Date. Period No. Distance (meters) per minute. Steps per minute (counter). Total number of steps. Counter record. Counted from kymo- graph record by Observer I. Observer II. 1914 Apr. 24.... Apr. 27.... May 4 . . . . May 5 il I 1 2 I 3 f 2 { 3 I 4 76.4 78.4 73.5 78.1 114.5 109.1 102.0 106.0 102.6 103.6 116.2 117.1 114.8 116.0 138.3 133.9 127.8 133.1 126.5 126.3 1,687 1,452 1,596 1,442 1,470 1,491 1,459 1,338 1,402 1,269 1,700 1,454 1,588 1,418 1,472 1,508 1,476 1,340 1,428 1,274 1,704 1,444 1,596 1,400 1,472 1,516 1,458 1,356 1,420 1,262 METHOD OF MEASURING HEIGHT TO WHICH THE BODY IS RAISED. One factor of the mechanics of walking has hitherto been neglected for the most part by writers, particularly when studies of the gaseous metaboUsm have been made, that is, the height that the body is raised during the process of walking. At each step the body is raised and lowered a distance of approximately 25 to 50 mm., and even higher in running.^ In any careful research on the physiology of walking in which the efficiency of the body for moving a certain weight a certain distance forward is studied, it is necessary to note the height of this upward movement. For this purpose Dr. Carl Tigerstedt used a device 'See p. 40. ^See table 20, p. 98. 40 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. which, in principle, was much like that used by him in studsdng mus- cular work done with the foot.^ This apparatus, shown in figure 6, consists of a small pointer A, travehng on a brass block between two upright guides. The cord G, attached to the pointer, passes over a pulley to the belt of the subject, as shown in figure 1. The spiral spring F, previously referred to in the description of the step-counter, keeps a tension upon the cord. Obviously as the body is raised the tension of the spiral spring draws the pointer down and as the body is lowered the cord draws the pointer up. Each upward and downward movement of the body- may be recorded on a rotating kymograph drum, thus giving a graphic record of the character of each step. A typical record is given in figure 7. It can be seen that the height to which the body is lifted may be obtained from the kymograph record by measuring each downward stroke of the pointer and adding the sum of the values. Experimental conditions prevented us from using a kymograph with a rapidly rotating drum or a long paper kymograph, and the con- sequent superimposing of the records shown in figure 7 obviously makes the counting of the number of steps somewhat difficult; neverthe- less it is possible and has been done in a number of instances (see table 2). By measuring several points in the curve, an approximate average height of step may easily be obtained; these averages may then be multiplied by the total number of steps recorded on the step-counter and the distance to which the body is raised computed. In an attempt to sum up these upward move- ments by means of some automatic arrangement, a special form of work adder-wheel was devised for the purpose. This work adder-wheel, which is shown as B in figure 6, relied upon the friction Fig. 6. — ^Apparatys for recording the height to which the body is lifted, and step-counter with connections. The cord G, connected with the body of the subject, passes over a grooved wheel B attached to the shaft of a revolution-counter C, a pawl p, preventing backward motion. It then passes to a pointer A (the marker of Dr. Carl Tigerstedt), tension being supplied by the light spring F. An upright cord travels over a grooved pulley on the step-counter D, with the tension produced by the small weight traveling in a tube E. The height to which the body is raised at each step is thus recorded in a dual manner, first, by the excursions of the pointer A over a kymograph dram and second by the accumu- lated movement of the wheel B. 'C. Tigerstedt, Skand. Archiv f. Physiol., 1913, 30, p. 299. Fig. 7. — Typical kymograph record showing character of step. The upper line shows the admission of oxygen to the apparatus, each section representing 1 liter; the second line the steps taken by the subject; the third line the respiration; and the lowest line the time in minutes. DESCRIPTION OF APPABATUS. 41 of a cord passing over a groove in the wheel to rotate it in one direction as the body was hfted, a pawl -p preventing any backward motion. When the body returned to the normal position, the cord slipped over the pulley without producing any movement of the wheel. Each upward movement of the body accordingly resulted in a forward motion of the wheel. The work adder-wheel was directly connected to the shaft of a revolution-counter, and a record of the total move- ment of the wheel was thus obtained. (See C, fig. 6.) The total dis- tance the body was raised would theoretically be that found by multi- plying the number of revolutions of the wheel by its circumference. At the very rapid rate of walking used in some of these experiments, great difficulty was experienced in the earlier development of the appa- ratus in securing an accurate record. The cord would frequently slip and there was considerable reverse movement, for it was not easy to find a pawl arrangement which would work perfectly. This reverse movement or "back-lash" was not overcome until practically the end of the series of observations; recently, however, a very thin laminated spring has been used as a pawl, which has practically ehminated this lost motion. The apparatus as described represents the completed form. As a matter of fact, in many of our experiments we were unable to use the records obtained from this wheel. The height to which the body was raised was therefore determined by the method previously referred to, i. e., measuring on the kymograph curve the general average dis- tance to which the body was raised and multiplying this by the record of the number of steps on the step-counter. In order to have these measurements of absolute value, the distance traveled by the pointer on the kymograph or that traveled by the cord passing over the work adder-wheel must represent exactly the distance over which the body is elevated or lowered. This assumes that there is no disturbance in the attachment of the cord to the body and that the tension of the spring is such as to prevent any diag or inequality in the movement of the pointer over the drum. As may be seen from the kymograph curve reproduced in figure 7, the constancy in the movement of the pointer was very satisfactory. On the other hand, the method of attaching the cord to the subject was certainly open to serious criticism. Wire was used for the most part, vioHn string being substituted for such portions as passed over the flexible parts, thus minimizing the tension and the danger of alterations in the length. There was, however, opportunity for considerable flexibility in the play of this cord at the point where it was attached to the belt holding up the trousers of the subject ; accordingly, the records obtained with the kymo- graph and with the work adder-wheel are without doubt invariably some- what too small and the body was in all instances raised to a higher point than that indicated in table 2. The expediency of attaching a wire direct to the body by means of sxu-geon's plaster was not resorted to, although the omission was undoubtedly an error in our observations. 42 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. PLAN OF RESEARCH. While this investigation was undertaken primarily with the object of studying the metabolism of walking on a level, certain preliminary observations were necessary, particularly for the purpose of estabhshing a base-Une for comparison with the metaboUsm during walking. As has previously been stated, certain investigators, in studying the metabohsm during walking, have been incUned to employ as a basis of comparison the metaboUsm determined with the subject in the lying position and without food in the stomach, i. e., maintenance metabo- hsm. Many attempts have also been made to study the metabohsm with the subject in the standing or the sitting position, these values being deducted from the total metaboUsm obtained with the subject walking, the increment being considered as due to the work of forward progression. It was therefore necessary in our research to study the metaboUsm not only during walking but also under other conditions, thus increasing the number of problems to be studied. A considerable number of experiments were carried out with the subject standing in different positions, those used being (1) with the subject standing with the body relaxed, as one would stand quietly without external support; (2) leaning against a support at the back; (3) leaning upon a staff; and (4) standing with muscles tense in the position of "attention." By determining the metaboUsm in these various resting or standing attitudes, all conceivable base-lines could be obtained. Furthermore, it was found that during experiments with rapid walking there was considerable lateral motion of the arms, as is the habit with many walkers, particularly professional pedestrians. Consequently certain experiments were made with the subject standing and swinging the arms from side to side as in a fast walk, but without moving the feet. A number of observations were also made of the metaboUsm with the subject sitting, with the idea that the values thus obtained might be used for the basal metaboUsm. As a matter of fact, only a few of the sitting experiments were made with the subject in the post-absorptive condition and the values secured have not been used for actual comparison purposes. The main object of the research was, of course, the study of the metaboUsm during walking and specificaUy the study of the increase in the metaboUsm due to walking at increasing speeds. In the series of walking experiments, therefore, the subject was required to walk at a very slow speed, then at a medium speed, and finaUy at a very fast speed. In a few experiments the subject actuaUy ran, thus giving data for comparing the work of forward progression while the subject was waUdng with that while he was running, two entu-ely distinct methods of forward progression. Certain observations were also made regard- ing the effect upon the metaboUsm of fatigue due to long-continued is PLAN OF RESEARCH. 43 walking, the subject being required on several days to walk for a con- siderable length of time. While, in common with several other investigators, we have beheved that the most sharply defined results can be obtained in experiments without food, nevertheless the experimental conditions were such as to make it relatively simple for us to obtain values after the ingestion of food. With this end in view, we made specific studies of the metabo- lism under the various conditions of standing, walking at various speeds, running, and after fatigue, not only when the subject was with- out food in the stomach but also after food had been taken. The experiments carried out after food duphcated the experiments with the subject in the post-absorptive condition; it was therefore possible to note whether or not there was a summation effect on the metaboHsm due to the ingestion of food and to the work. While in some of the experiments the diet was uncontrolled, in a number of them it was prescribed. The constituents of the meal varied widely in these latter experiments, a special protein diet being supphed on some days, on others a diet containing an excess of fat, and again a diet with a large proportion of carbohydrate. The results of these experiments accordingly gave data as to the effect of a special diet upon the metaboUsm during walking. This, in brief, was our plan of research at the beginning of the series of observations. Incidentally a number of other important physio- logical details were developed as the research progressed. These will be taken up specifically in the discussion of the results of the experi- ments. 44 ENEEGY TRANSFORMATIONS DURING HORIZONTAL WALKING. GENERAL ROUTINE OF THE EXPERIMENTS. It will be seen from the foregoing that the experiments in this research fall naturally into four classes: standing experiments, sitting experi- ments, walking and running experiments, and experiments with food. From the earUer experience of Cathcart with a subject riding a bicycle, it seemed inadvisable to attempt the estabhshment of a standard test- ing base-Une:4;o be deducted from the total metaboHsm of the subject while walking. It was therefore decided that each morning, prior to the walking experiment, measurements should be made of the standing resting metabolism; consequently the research included a large number of standing experiments. STANDING EXPERIMENTS. The respiration apparatus used for this research and previously described was designed more especially for measuring the large amounts of carbon dioxide exhaled during severe muscular exercise, and was, in consequence, fitted with two soda-lime bottles, each of which must be weighed. If only the resting metabolism had been determined, a slightly different arrangement would have been preferred, with the use of but one soda-lime bottle, a spirometer form of apparatus being substituted for the rubber bathing-cap and tension-equaUzer. Never- theless, since the resting metaboHsm with the subject standing was to be used primarily as a base-hne to be deducted from the greatly increased metabolism during walking, it was considered that this method of measuring the metaboHsm during standing would be sufficiently satisfactory for the purpose, and hence the standing metaboHsm was determined on exactly the same apparatus as that used in the walking experiments. As a matter of fact, the experiments made by Cathcart, and more especiaUy those of Carpenter,^ have shown that the tension- equaHzer form of the unit apparatus gives as accurate measurements of the metaboHsm during rest as does the spirometer type. In this par- ticular apparatus, however, we could readily have dispensed with the second soda-Hme bottle. After arriving at the Laboratory, the subject arranged his clothing for an ordinary walking experiment and assumed the standing position on the treadmiU, this position being maintained for some time prior to the actual experimentation. After the effect of the sHght exertion of coming to the Laboratory and of ascending the stairs had passed away, the mouthpiece was inserted, the nosecHp attached, and at the proper time, i. e., at the end of a normal expiration, the 3-way valve connecting the subject with the ventilating system of the respu-ation apparatus was turned. The experiment was then continued in the usual manner ^Carpenter, Carnegie Inst. Waah. Pub. 216, 1915, pp. Ill to 118. GENERAL ROUTINE OF THE EXPERIMENTS. 45 for approximately 15 minutes. At the end of a normal expiration the valve was again turned and the experiment was completed. During the entire experiment oxygen was admitted regularly from a cylinder, the gas being passed through a carefully cahbrated Bohr gas-meter immersed in water. At the conclusion of the experiment the ventila- tion was continued for a few minutes to insure the thorough sweeping out of all the carbon dioxide; finally the tension-equahzer was filled with oxygen to the same tension that existed at the beginning of the experiment. Owing to the greatly increased carbon-dioxide production during severe muscular work, the ventilating current was so adjusted that the ventilating pump would cause 80 to 90 Uters of air to pass by the mouth of the subject per minute, thus minimizing the danger of the rebreathing of the air. Furthermore, by means of the supplementary valve M (see fig. 2, page 34), the dead space in the rubber tube leading from the mouthpiece to the main air-pipe was wholly eliminated, for when the valve N had been turned after the beginning of the experi- ment, all of the air passed immediately by the mouthpiece. While this deflection of the air-current was unnecessary in the experiments with the subject standing, nevertheless it was also used in these experi- ments in order that the procedure might be the same in both series. Since it is important to note whether or not the subject remained in essentially the same degree of muscular repose throughout the standing experiments, a graphic record was obtained of the degree of movement by connecting the subject with a cord attached to a movable pointer traveling over a smoked-paper kymograph. In an ideal experiment the course of the pointer would be a straight hne. While it was impos- sible for the subject to remain as perfectly quiet as he would when lying upon a comfortable couch, nevertheless the regularity and constancy of the muscular repose of the subject from experiment to experiment was, to say the least, very striking. We are therefore safe in assuming that in practically no experiment were the irregularities in the metabo- lism for the standing position attributable to changes in the degree of muscular repose of the individual, except in those tests in which special positions while standing were assumed, such as the position of "attention." SITTING EXPERIMENTS. A few sitting experiments were also made with one of the subjects. These were not used as a base-Une, but are recorded simply to show that an endeavor was made to secure the best possible base-line for deduc- tion from the total metabolism obtained during walking. The incre- ment in the metaboUsm for the sitting position was unfortunately not studied with sufficient sharpness to make a definite conclusion possible. 46 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. WALKING AND RUNNING EXPERIMENTS. In the walking and running experiments the routine was essentially that previously outlined for the standing experiments, save that during the entire period and for several minutes before the experiment began the subject was walking upon an electrically driven treadmill which was kept in motion at a definite rate of speed. These rates of speed varied from 53 meters per minute to 149 meters per minute. The flexibility of the mouthpiece and its attendant by-pass made it possible for the subject to walk with perfect comfort at these varying speeds and the vertical movement of the body with each step did not interfere in the shghtest with the correct measurement of the gaseous exchange. With a slow speed, there was no noticeable extraneous muscular effort other than that of walking; with a high speed the subject made vig- orous lateral motions of the arms, as is quite common with profes- sional pedestrians when walking at high speed. As previously noted, an attempt was made to measure the influence of this lateral motion by studjdng the metaboUsm with the subject standing and swinging the arms. In a number of the experiments the subject ran, although the rate of progression was but little higher than in the experiments with fast walking. EXPERIMENTS WITH FOOD. , A considerable number of experiments were made after a breakfast selected by the subject; a few were also made after dinner, the diet being uncontrolled. In the experiments in which a study was made of the possible influence of diets containing a preponderance of protein, fat, or carbohydrate, the first meal of the day was given to the subject at the Laboratory after a walking experiment in the forenoon in which he was in the post-absorptive state. This meal consisted of steak, rice, potatoes, or various fats. A second walking experiment was then made under otherwise identical conditions. The exact weight of food was not recorded, since our only aim was to make sure that the body was plentifully suppHed with the special food constituent being studied. In general, the subject ate aU that he possibly could and as a con- sequence was frequently disincUned to walk in the afternoon. A careful study of the effect of muscular work upon the urine was impracticable, hence no analyses of the urine accompany these observa- tions. Previous tests with one of the subjects have shown, however, that the accompUshment of a large amount of severe muscular work was not accompanied by an excessive excretion of nitrogenous products in the urine.^ 'Benedict and Cathoart, Carnegie Inst. Wash. Pub. 187, 1913, p. 98. SUBJECTS. 47 SUBJECTS. In this series of observations two subjects were used, both of whom were more or less trained to severe muscular activity, as one (A. J. O.) was a semi-professional baseball player and the other (M. A. M.) a professional bicycUst. The preUminary observations in this research were carried out by Dr. Carl Tigerstedt with the subject A. J. O. (sub- ject I), a man of athletic buUd, who readily adapted himself to the observations on the treadmill and with the respiration apparatus. He was 29 years of age, 180 cm. in height, and had a body-weight with clothing varying from 72.1 kilograms to 74.8 kilograms and without clothing of 69.7 kilograms. Since in this study one of the prime objects was to note the energy required to move 1 kilogram of material in the forward direction 1 meter, i. e., 1 horizontal kilogrammeter, the body- weight with clothing is of importance in each experiment. These weights are given in detail in table 13 (page 78). The major portion of the research was devoted to a study of forward progression at varying speeds with the subject M. A. M., who had previously served as the subject of Cathcart in the study of the mus- cular work of bicycle-riding. This subject, who in this pubhcation is designated as subject 11,^ was 31 years of age, 177 cm. in height, and had a body-weight with clothing varying from 69.9 to 72.4 kilograms. The average body-weight without clothing during this series of experi- ments was 68.3 kilograms. The body-weight with clothing is given for each experiment in tables 14 and 16 (pages 83 and 88). 'Further data regarding the body measurements, etc., of this subject may be found in the report of the previous study on muscular work. See Benedict and Cathcart, Carnegie Inst. Wash. Pub. 187, 1913, p. 35. 48 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. STATISTICS OF EXPERIMENTS. The data regarding the experimental conditions, the respiratory exchange, and the mechanics of walking for the experiments with sub- ject I (A. J. O.) are given in table 3; the data for subject II (M. A. M.) appear in table 4. Table 3. — Summary of results obtained with subject I in experiments withovi food, during the period from Nov. 26, 1913, to Dec. Z7, 191S. [Observations made by Dr. Carl Tigerstedt. Values per minute.] Date and condition. Dura- tion. Nmi. se.' Standing,'' 10'' 10" a. m. . Standing,* lO*" 45™ a. m. . Food about 1 p. m. Standing, 2'' 25" p. m . . . Standing, 2'' 45" p. m . . . Walking,^ S"" 24" p.m... Nov. S8. , Standing, lO*" 29" a. m . . Standing, ll"" 03" a. m . . Standing, ll"" 30" a. m . . Nov. B9. Standing, Oi" 16" a. m . . . Standing, g"" 50" a. m . . . Walking,' lO"" 22" a. m . . Dec 1. Standing, Q* 30" a. m . . . Standing, O"" 57" a. m . . . Walking,' lO^" 51" a. m. . Walking,' 11'' 34" a. m. . Dec. 2. Standing, S'' 54" a. m . . . Standing, Q"" 17" a. m . . . Standing, lO'' 15" a. m . . Walking,' 10'' 30" a. m. . Walking,' 11'' 18" a.m. . Dec. S. Standing, O"" 10" a. m . . . Standing, Q"" 38" a. m Standing, lO"- 06" a. m . . , Walking,' 10'' 45" a. m . . , Walking,' 11'' 05" a. m. . . Walking,' 111" 35" a. m. . . Standing, ll"" 56" a. m . . . Standing, 12'' 20"p. m . . . mm. sec. 15 17 15 45 15 04 15 35 15 15 IS 07 15 06 15 48 15 18 15 19 15 02 Car- bon diox- ide. 15 11 15 12 9 59 9 57 15 28 15 22 IS 25 4 44 10 16 16 07 15 07 15 05 10 47 10 17 10 17 13 55 15 09 c.c. 283 278 277 268 943 226 222 218 228 219 886 222 217 692 731 Oxy- gen. c.c. 343 321 341 328 1015 284 291 279 282 282 873 276 282 852 904 Respi- ratory quo- tient. 234 290 223 289 214 280 630 853 695 849 226 286 214 269 209 270 685 845 701 842 697 872 215 270 206 267 0.82 .86 .81 .82 .93 .80 .76 .78 .81 .78 1.02 .81 .77 .81 .81 .81 .77 .77 .74 .82 .79 .80 .78 .81 .83 .80 .80 .77 Aver- age pulse- rate. 80 87 85 96 95 91 94 90 94 90 88 90 99 97 85 89 Average respi- ration- rate. Dis- tance, Rais- ing of body. 18.3 18.0 18.4 27.9 16.3 17.2 17.7 18.6 18.3 25'. 7 19.0 19.5 25.4 24.5 19.4 21.1 21.5 26.8 25.3 19.5 21.8 21.9 27.3 26.7 25.5 23.0 23.0 meters. 75.3 meters, 3.57 75.1 3.15 76.0 76.6 76.2 76.8 76.5 77.2 77.5 3.26 3.36 3.20 4.62 3.57 3.62 3.68 'The subject had breakfast at about 7 a. m. 'Throughout this summary of resets, the designation "standing" signifies standing in a relaxed position unless otherwise specified. --/iiuuius m a 'Subject I walked at a rate of not far from 105 steps per minute th7-niiir>' 18" p.m... 20 08 719 905 .79 26.5 78.6 4.15 Dee. 18. Standing, 9^ 03" a. m 25 05 238 272 .88 88 21.1 Standing, Q"" 54" a. m . . . . 25 33 230 263 .87 88 21.3 Standing, Vf" 34" a. m . . . 25 13 220 255 .86 83 21.6 Walking,! ll* 08" a. m... 20 04 726 811 .89 23.9 75.3 3.52 Walking,! 11'' 40" a. m . . . 20 28 740 833 .89 25.2 76.4 3.68 Walking,! 12>> 10" p. m. . . 20 04 730 864 .84 27.7 76.7 3.68 Walking,! 12'' 39" p. m. . . 20 12 734 867 .85 27.3 76.9 3.73 Dee. 19. Standing, gi" 43" a. m 25 03 238 279 .85 96 22.5 Standing, Itf^ 16" a. m . . . 25 06 239 282 .85 97 23.3 Standing, Itf- 49" a. m . . . 25 06 241 275 .88 100 23.1 Walking,! iiii25"a. m. . . 20 06 789 869 .91 23.9 76.1 3.83 Walking,! ll"" 54" a. m... 20 06 772 860 .90 28.1 77.9 3.99 Walking,! 12'' 21" p. m . . . 20 10 755 871 .87 29.0 78.3 4.10 Walking,! 12'' 52" p. m. . . 10 40 759 872 .87 29.8 78.7 4.20 Dec. SO. Standing, 8'' 55" a. m 25 06 237 272 .87 99 23.3 Standing, 9'' 29" a. m 25 17 271 106 22.5 Standing, 10'' 06" a. m . . . 25 10 224 273 '.'82 106 23.2 Standing, 11'' 10" a. m . . . 24 56 216 263 .82 100 22.5 Walking,! nk 44m a. m. . . 20 04 690 825 .84 26.1 76^2 S.6S Walking,! 12'' ll"p. m. . . 20 07 735 884 .83 28.5 77.7 4.04 Walking,! 12'' 40" p. m . . . 20 02 731 924 .79 30.1 78.6 4.20 Walking,! l*" 02" p. m 19 59 747 934 .80 31.3 78.9 4.25 Dec. SS. Walking,! ^Qk 17" a. m. . . 20 15 718 873 .82 27.4 76.3 4.07 Walking,! iQh 47m a. m. . . 20 06 610 790 .77 28.6 65.9 3.17 Walking,! 11'' 20" a. m. . . 20 07 641 901 .71 30.2 77.4 4.28 Walking,! ll''48"a. m. . . 20 06 600 788 .76 29.7 67.5 3.41 Dec. gS. Walking,! 9'' 25" a. m... 20 29 695 833 .83 25.7 75.9 3.97 Walking,! gh 59m a. m 20 47 589 740 .80 24.2 66.4 3.07 Walking,! 10'' 38" a. m... 20 18 694 863 .80 27.2 76.9 4.08 Walking,! n'' I2"a. m. . . 20 06 602 764 .79 26.0 66.4 3.30 Dec.g7. Walking,! 9'' 56" a.m.... 21 53 685 846 .81 27.0 78.4 4.06 Walking,! 10^41" a. m. . . 20 43 563 725 .78 26.9 65.2 3.02 Walking,! 11'' 15" a. m. . . 20 52 664 852 .78 28.6 78.6 4.13 3.02 Walking,! 11'' 48" a. m... 20 11 560 732 .76 29.1 65.1 !Subject I walked at a rate of not far from 105 steps per minute throughout the waUdns series except on Dec. 22, 23, and 27. The total distance walked on Dec. 17 was 8 798 meters- Dec 18 8,435 meters; Dec. 19, 7,581 meters; Dec. 20, 8,142 meters; Deo. 22, 10 075 meters-' Dec' 23* 10,338 meters; Dec. 27, 11,908 meters. The subject walked between periods. The stera ne^ minute in the last three periods on Deo. 22 were 95, 101, and 95; during the periods on D^ 9" on uec. -Sd, STATISTICS OF EXPERIMENTS. 51 Table 4. — Summary of results in experiments vdth subject II, during the period from Mar. 16, 1914, to May 15, 1914. [Values per minute.] Date, condition, and time. Dura- tion. Car- bon diox- ide. Oxy- gen. 1^ >-3 24™p. m 17 12 743 759 .98 24.5 59.3 96 2.12 Mar. SO. Food, 7'' 30™ a. m. : Standing, attention, 9'' 36™ a. m . 15 54 261 271 .96 76 20.2 Standing, staff, lO"" 15™ a. m . . . 14 51 261 296 .88 71 18.9 Walking,* ll"- 17™ a. m 17 15 658 744 .88 ■ ■ 19.2 68^5 ioo 2.26 Food, 12''45™p. m.: Standing, staff, l"" 50™ p. m 15 8 318 339 .94 95 21.3 Walking,' 2'' 27™ p. m 16 15 56 11 681 666 722 709 .94 .94 24.8 21.9 58,5 57.3 92 99 2!40 2.24 WaUdng," 3'' 26™ p. m Mar. 31. Without food: Standing, support, &' 38™ a. m . . 17 48 214 239 .89 77 18.4 Standing, relaxed, 9^ 21™ a. m. . . 16 40 209 260 .80 71 17.8 Standing, attention, 10^ 02™ a. m 13 15 9 59 233 571 272 673 .86 .85 75 19.4 21.6 57.2 98 2.01 Walking,' ll"" 47™ a. m 'Leaning comfortably against support, hands on back. 'Total distance before lunch, 1,286 meters; after lunch, 1,345 meters. Subject held his hands on his legs as he walked. 'Total distance before lunch, 1,184 meters; after lunch, 1,266 meters. Subject held his hands at his sides as he walked. *Total distance before lunch, 1,158 meters; after lunch, 1,210 meters. In the morning, subject walked with hands held on his legs; in the afternoon his hands were swinging at his sides. 'Total distance before lunch, 1,198 meters, subject walking with hands swinging. After lunch distance was 2,229 meters, hands being on support in the first period and swinging at his sides in the second period. Most of the time between periods he was not walking. 'Total distance before lunch, 1,085 meters; after lunch, 1,020 meters. 54 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. Table 4. — Summary of results in experimerds with subject II, during the period from Mar. 16, 1914, to May IS, 1914 — Continued. [Values per minute.] Date, condition, and time. Dura- tion. Car- bon diox- ide. Oxy- gen. ■g a & 2 O p. Dis- tance. Steps. Rais- ing of body. Mar. SI — Continued. Food, 1 p. m.: Walking,' 2'' OS" p. m Standing, relaxed, 2'' 55" p. m . . Standing, attention, S"" 40" p. m. Apr. 1. Without food: Standing, 9^ 50™ a. m Walking,* 11'' 21"° a. m Food, 12 noon: Standing, l"" 42°> p. m mm. 14 15 11 15 15 sec. 57 36 52 16 30 Avr. S. Food, 7''15"a. m.: Standing, support, 8^ 44" a. m . Standing, relaxed, 9*' 53" a. m . Standing, attention, lO*" 46" a. m 15 12 Apr. S. Without food : Sitting, g"" 46" a. m Standing, staff, 9'' 26" a. m . . Walking,' Itf- 33" a. m Food, 12'' 45" p.m.: Walking,' l"" 28" p. m Walking,' 2'' 28" p. m Walking,' S"" 35" p. m Standing, 4'' 28" p. m Apr. 4. Food, 7''15"a. m.: WaUdng,* lO"" 11" a. m Walking,'' lO"" 56" a. m Walking,* 11'' 29" a. m Food, l''15"p. m.: Walking,* 2 p. m Walking,* 2'" 65" p. m Apr. 6' Without food : Standing, 9^ 04" a. m Walking,' 9^ 41" a. m Walking,' 10'' 22" a. m Walking,' 111" 13" a. m Protein, 12'' 55" p. m.: Standing, l"" 24" p. m Walking,' 1'' 46" p. m Walking,' 2'' 24" p. m Walking,' S*" 05" p. m 12 18 15 20 15 7 13 43 14 18 14 7 14 52 16 24 15 35 15 42 14 18 13 57 15 46 16 9 16 11 15 41 17 14 13 16 15 59 14 36 16 49 c.c. 686 277 265 213 563 283 273 253 248 219 224 771 667 894 860 244 608 573 554 657 705 218 586 567 563 241 631 641 617 c.c. 706 345 326 266 699 319 306 281 268 246 278 903 686 938 895 293 651 620 669 707 755 246 681 668 659 296 726 712 708 0.97 80 81 .80 .81 .90 .93 .89 .81 .86 .97 .96 .96 .83 81 23.9 20.6 20.7 17.9 21.5 20.2 20.0 19.7 .93 '73 .92 '79 .83 93 93 89 674 86 '72 86 =74 84 '77 82 87 '78 90 87 '86 meters. 57.7 69.3 lO.O 19.4 19.1 20.6 76.2 23.4 67.0 22.2 77.0 22.8 78.9 22.0 20.7 55.1 22.6 66.8 21.3 66.2 23.3 54.4 24.6 58.1 19.8 17.3 57.8 20.9 56.8 21.6 56.0 19.2 19.4 67.5 21.6 57.2 23.7 56.3 99 100 114 100 110 112 100 100 99 101 101 104 106 102 mstert. 1.72 2.22 2.38 2.02 2.84 3.38 1.91 1.98 1.87 2.05 2.21 1.78 1.83 1.91 105 2.21 105 1.94 99 1.81 'Total distance before lunch, 1,085 meters; after lunch, 1,020 meters. "Total distance preliminary to and during period, 1,164 meters. 'Total distance before lunch, l,244meters; after lunch, 7,296 meters. During about 43 minutes between the first two periods of the afternoon subject was not walking. *Total distance before lunch, 6,207 meters; after lunch, 4,132 meters. The walking was continuous. 'Electrocardiograms were obtained during these periods by means of the string galvanometer. •On and after Apr. 6, subject's food was prepared and eaten at the Laboratory. 'Total distance before lunch, 6,140 meters; after lunch, 5,433 meters. The walking, as was usually the case in the days that follow, was continuous. STATISTICS OF EXPERIMENTS. 56 Table 4. — Summary of results in experiments urith subject II, during period from Mar. 16, 1914, to May 15, J9i.^— Continued. [Values per minute.] Date, condition, and time. Dura- tion. Car- bon diox- ide. Oxy- gen. "I Average respira- tion-rate. Dis- tance. Steps. Rais- ing of body. min. sec. c.c. c.c. meters. Tneters 14 29 214 257 0.83 =^82 19.2 15 30 606 729 .83 275 20.1 59.6 109 i.98 15 59 544 660 .82 '79 22.1 58.2 105 1.95 14 8 198 235 .84 ■^0 21.4 13 23 217 289 .75 !'88 19.6 16 19 643 732 .88 =^81 20.4 68.8 107 1.78 13 36 671 772 .87 288 25.1 58.7 107 1.83 13 59 593 701 .85 24.5 56.9 99 1.82 14 6 204 265 .77 18.4 12 15 761 922 .82 21.0 76.1 121 3.01 14 22 662 826 .80 23.0 77.6 117 2.81 13 29 660 807 .82 24.4 78.3 113 2.85 12 54 230 275 .83 19.1 13 22 785 869 .90 , . 22.6 76.9 120 2.50 13 47 792 892 .89 25.7 78.0 117 3.01 13 1 782 917 .85 25.6 79.2 118 2.70 14 21 202 228 .89 82 18.7 15 15 590 701 .84 21.0 59.3 107 1.92 14 23 546 663 .82 22.5 58.8 105 2.05 14 34 568 697 .81 25.1 59.0 107 1.98 12 42 245 295 .83 76 19.4 15 32 649 704 .92 22.9 59.4 107 i.95 13 48 646 691 .93 26.2 59.0 106 1.94 15 37 651 715 .91 26.1 58.8 104 1.94 13 39 212 257 .83 78 19.3 14 22 795 814 .98 22.6 77.3 120 2.77 12 46 712 841 .85 24.4 78.9 114 3.21 13 13 264 301 .88 86 21.1 13 23 879 948 .93 23.1 78.0 lis 3.01 14 31 857 926 .93 26.0 79.1 116 3.17 14 25 823 881 .93 26.8 79.7 116 3.35 15 52 223 245 .91 94 19.0 13 51 630 695 .91 20.7 61.7 104 2.45 14 35 597 666 .90 23.6 62.1 103 2.34 14 59 563 672 .84 24.4 62.4 102 2.37 Apr. 7. Without food: Standing, 9'' 18™ a. m Walking,' 9^ 56°' a. m Walking,! 10'' 58" a. m. . . Standing, ll"" 27°' a. m . . . Protein, 1'' 03" p. m. : Standing, l"" 31'° p. m Walking,' 2'' 07"° p. m WaUdng,' 3 p. m Walking,' 3'' 42" p. m Apr. 8. Without food : Standing, 9^ 10" a. m . . . . Walking,' 9'' 40" a. m Walking,' lO"" 14" a. m. . . Walking,' 1 1'' 01" a. m... Protein, 1 p. m.: Standing, li- 23" p. m Walking,' 2'' 01" p. m ... . Walking,' 2'' 44" p. m.... Walking,' 3'' 30" p. m ... . Apr. 9. Without food : Standing, 8'' 42" a. m Walking,* 9'' 22" a. m Walking,* lO"" 07" a. m . . . Walking,* lO*" 47" a. m . . . Carbohydrate, 1 p. m.: Standing, l"" 19" p. m Walking,* l"* 45" p. m Walking,* 2'' 26" p. m Walking,* S^" 14" p. m Apr. 10. Without food : Standing, 8'' 54" a. m Walking,' 9'' 34" a. m Walking,' lO*" 57" a. m . . . Carbohydrate, 12'' 08" p. m. Standing, 12'' 31" p. m... Walking,' l"" 16" p. m Walking,' l''50"p. m Walking,' 2'' 30" p. m Apr. 14. Without food : Standing, 9'' 02" a. m Walking,' 9'' 48" a. m Walking,' 10* 22" a. m . . . Walking," 11'' 06" a. m . . . 'Total distance before lunch, 5,007 meters; after lunch, 6,560 meters. Electrocardiograms were obtained during these periods by means of the string galvanometer. 'Total distance before lunch, 7,287 meters; after lunch, 7,933 meters. *Total distance before lunch, 5,868 meters; after lunch, 6,141 meters. Total distance before lunch, 8,372 meters; after lunch, 7,221 meters. «Total distance before lunch, 6,157 meters; after lunch, 6,062 meters. 56 ENERGY TEANSPORMATIONS DURING HORIZONTAL WALKING. Table 4. — Summary of results in experiments with subject II, during the period from Mar. le, 1914, to May IS, iSi^— Continued. [Values per minute.] Date, condition, and time. Dura- tion. Car- bon diox- ide. Oxy- gen. as (O .SS-.B P. o Da ^ !>3 Dis- tance. Steps. Rais- ing of body. Apr. 14 — Continued. Carbohydrate, 1 p. m. ■} Standing, l"" 23° p. m Walking,'' l"" 59"° p. m Walking,' 2'' 37"» p. m Walking,=i S"" IS™ p. m Apr. IB. Without food: Standing, S"- 48"° a. m Walking,' Q"" 30° a. m Walking,' 10'' 08° a. m Walking,' lO* 53° a. m Carbohydrate, 12'' 52° p. m.: Standing, l"" 14° p. m Walking,' l*" 52° p. m Walking,' 2'' 29° p. m Walking,' 3'' 05° p. m Apr- IS. Without food : Standing, %^ 32° a. m Walking,* 9'' 14° a. m Walking,* 9'' 53° a. m Walking,* 10'' 36° a. m Fat, 11'' 30° a. m.: Standing, 12'' 03-" p. m Walking,* 12'' 44° p. m Walking,* l"- 20° p. m . . . Walking,* 2'' 14° p. m . . . Walking,* 3'' 10° p. m . . . Apr. 17. Without food : Standing, 8'' 31° a. m Walking,'^ 9'' 08° a. m Walking,^ 9'' 50° a. m Walking,' lO"" 37° a. m Fat, 11'' 30°' a.m.: Standing, 12'' 09° p. m Walking,' 12'' 48° p. m Walking," l*" 34° p. m Walking,' 2'' 18° p. m Walking,' 3'' 17° p. m Apr. m. Without food : Standing, 10'' 25° a. m Walking,' 11'' 16° a. m Fat, 12''30°'p. m.: Standing, l''03°p. m Walking,' l''45°p. m Walking,' 2'' 25° p. m Walking,' 3'' 10° p. m Walking,' 4'' 01° p. m mt/n. 14 14 15 14 15 12 15 14 14 14 14 17 14 15 12 14 14 14 15 15 18 14 13 14 14 16 14 13 14 16 13 15 12 14 14 14 16 c.c. 242 681 672 671 214 818 696 690 267 875 859 898 568 544 553 239 600 593 611 586 214 773 672 693 231 715 735 799 721 205 562 220 590 624 608 592 c.c. 295 674 711 693 252 934 829 837 277 902 878 924 258 644 668 661 292 683 635 701 707 265 895 816 862 281 832 883 938 856 242 639 301 683 699 705 709 0.82 1.01 .94 .97 .85 .88 .84 .82 .96 .97 .98 .97 .81 .86 .82 .80 .82 .86 .83 .85 .84 .85 .73 .86 .89 .86 .83 90 74 83 90 85 99 78 91 19.5 23.4 25.6 25.9 17.7 22.7 25.7 25.4 20.9 26.4 28.9 28.9 19.5 21.0 24.4 25.1 19.8 23.4 25.7 26.2 25.0 20.9 23.8 24.6 26.8 20.5 23.5 26.8 27.9 27.2 19.1 22.8 19.7 21.8 25.2 25.8 25.4 metere. 6i!2 62.0 62.2 77.2 79.4 80.2 78.2 80.1 80.6 61.9 62.3 61.8 61.6 62.3 62.2 61.9 76.6 78.1 79.3 77.6 79.5 80.0 80.0 60.8 60.4 60.6 60.5 60.5 102 102 104 119 115 114 117 115 117 105 106 106 107 105 105 103 118 117 117 118 117 119 115 105 104 104 103 105 meters. 2.38 2.20 2.27 3.33 3.52 3.49 3.88 3.72 3.88 2.38 2.37 2.20 2.49 2.48 2.24 2.21 3.36 3.45 3.80 3.96 3.78 3.85 3.67 2.32 2.45 2.56 2.34 2.23 'Subject did not eat so much as on other days. 'Total distance before lunch, 6,157 meters; after lunch, 6,062 meters. 'Total distance before lunch, 8,306 meters; after lunch, 7,657 meters. *TotaI distance before lunch, 6,350 meters; after lunch, 10,547 meters. 'Total distance before lunch, 8,680 meters; after lunch, 13,566 meters. 'Total distance preliminary to and during period before lunch, 1,651 meters; after lunch 1,494 meters. ' ' STATISTICS OF EXPERIMENTS. 57 Table 4. — Summary of results in experments with subject II, during period from Mar. 16, 1914, to May 16, iSi^— Continued. [Values per minute.] Date, condition, and time. Dura- tion. Car- bon diox- ide. Oxy- gen. 03 to & 2 '2^ Dis- tance. Steps. Rais- ing of body. Apr. gS. Without food: Standing, S"- 39"° a. m Walldng,' 9^ 27™ a. m Walking,! Kfi 09" a. m Walking,! loi" 44"" a. m Fat, 11'' 40" a. m.: Standing, 12'' 14'° p. m Walking,' 12'' SS"" p. m Walking,' 1'' 34"" p. m Walking,! g"" 20" p. m Walking,! 3I1 oi" p. m Apr. S3. Without food : Standing, 8'' 40" a. m Walking,' 9'' 34" a. m Walking,' 10'' 15" a. m Protein, 11'' 10" a.m.: Standing, 11'' 40" a. m Walking,^ 12'' 24" p. m Walking,* li" 03" p. m Walking,* l''47"p. m Walking,' 2'' 28" p. m Apr. Z4. Without food : Standing, 9'' 14" a. m Walking,' 9'' 55" a. m Walking,' 10'' 26" a. m Protein, 11'' 15" a. m.: Standing, 11'' 58" a. m Walking," 12'' 35" p. m Walking,' l"" 07" p. m Walking,' l"" 42" p. m Walking,' 2'' 15" p. m Apr. SB, Without food : Standing, 9)^ 30" a. m Walking,* 10 a. m Walking,* 10'' 33" a. m Carbohydrate, 11'' 20" a. m.. Standing, 11'' 49" a. m . . . . Walking,* 12'' 25" p. m Walking,* 12'' 56" p. m Walking,* l"" 40" p. m Walking,* 2'' 30" p. m mvn. sec. 14 20 15 17 13 26 13 48 14 11 14 30 13 37 13 18 14 03 15 57 15 23 16 37 14 7 13 13 18 13 56 15 5 12 43 14 31 12 24 14 19 14 21 13 9 13 54 14 12 59 13 24 13 25 12 21 14 37 17 17 15 23 15 13 c.c. 207 687 655 647 227 725 729 737 739 208 561 528 223 644 607 606 605 198 635 636 219 699 696 704 712 192 543 531 274 668 662 688 641 c.c. 251 804 829 822 299 833 877 877 858 252 675 660 254 722 705 707 707 790 828 299 835 853 890 880 264 665 695 324 708 711 694 0.83 .85 .79 .79 .76 .87 .83 .84 .86 .82 .83 .80 .88 .89 .86 .86 .86 .80 .77 .73 .84 .81 .79 .81 .73 .82 .76 .85 .94 .93 .99 84 93 74 81 82 94 79 83 18.0 23.5 26.1 25.7 20.8 26.0 28.7 28.4 28.4 19.3 20.5 24.2 20.1 19.7 24.6 25.0 24.3 18.7 21.4 25.8 20.3 22.9 26.3 26.3 25.4 19.6 20.2 23.3 20.6 22.2 24.4 25.7 25.2 meters. 76.8 78.6 79.2 77.3 78.7 79.5 79.7 61.1 61.2 61.6 61.6 61.7 76.4 78.4 77.0 78.8 80.0 80.0 63.0 64.0 63.1 63.8 63.7 63.3 118 118 115 116 118 118 116 105 105 106 105 104 103 116 117 114 115 115 115 105 107 104 105 106 104 meters. 3.33 3.44 3.42 3.86 3.86 3.92 3.84 2.53 2.46 2.74 2.57 2.48 2.63 3.60 3.69 3.74 3.88 3.84 3.88 2.65 2.69 2.88 2.70 2.85 2.64 'Total distance before lunch, 7,548 meters; after lunch, 11,466 meters. 'Total distance before lunch, about 4,300 meters; after lunch, 8,908 meters. Total distance before lunch, 3,888 meters; after lunch, 9,414 meters. *Total distance before lunch, 3,216 meters; after lunch, 9,089 meters. 58 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. Table 4.— Summary of results in experimenis viith subject II, during period from Mar. 16, 1914, to May 16, iSi^— Continued. [Values per minute.] Date, condition, and time. Dura- tion. Car- bon diox- ide. Oxy- gen. i>3 ,a » <".S Dis- tance. Steps. Rais- ing of body. Apr. 27. Without food : Standing, 9'' 10"° a. m Walking,' Q"" 48°" a. m Walking,! lo* 20°' a. m Carbohydrate, ll"" lO"" a. m.: Standing, ll"" 34"' a. m Walking,' 12'' 12'° p. m Walking,' 12'' 45'» p. m Walking,' 1'' 21'» p. m Walking,' 2'' 08"" p. m Apr- ^8. Without food: Standing, 8'' 39" a. m Walking,^ gi" 12'° a. m Walking,^ 9'' 48'» a. m Walking.ii 10'' 26'" a. m Walking,* 11'' 07" a. m Walking,2ll''51'»a. m.... Walking,* 12'' 33'° p. m Walking,* 1'' OG"" p. m Walking,* l'' 49" p. m Apr. 29. Carbohydrate, 8'' 35" a. m.: Walking," 9'' 08" a. m Walking,' 9'' 45" a. m Walking,' 10'' 24" a. m Walking,' 11'' 04" a. m Walking,' 11'' 40" a. m Walking,' 12'' 18" p.m.... Walking,' 12'' 58" p. m Walking,' l"" 36" p. m May 4- Without food: Walking,* 9'' 27" a. m Walking,* 10'' 02" a. m Walking,* 10'' 35" a. m Carbohydrate, 11'' 19" a. m: Walking,* 12'' 02" p. m ... . Walking,* 12'' 43" p. m Walking,* l''16"p. m mm. 13 13 12 13 14 13 15 15 12 15 13 13 13 15 14 14 IS 14 14 14 14 14 13 14 15 10 11 11 10 11 10 c.e. 217 681 679 294 845 837 886 850 221 713 699 680 689 716 657 682 659 817 789 817 797 777 696 704 666 1183 1052 934 1212 1237 1177 c.c. 245 767 821 326 854 797 840 801 274 828 820 846 858 925 854 865 859 864 863 839 810 823 790 848 830 1377 1175 1117 1336 1295 1221 0.89 .89 .83 .90 .99 1.05 1.05 1.06 .81 .86 .85 .80 .80 .77 .77 .79 .77 .95 .91 .97 .98 .94 .88 .83 .80 .86 .90 .84 .91 .95 .96 77 75 82 20.5 23.3 26.5 21.0 24.7 27.5 28.6 27.6 19.6 23.0 26.4 26.1 26.5 27.4 27.2 28.1 27.8 27.1 27.2 27.8 26.9 26.6 27.2 26.9 27.1 25.1 27.7 26.9 27.1 28.6 28.6 meters. 73.5 78.1 77.5 78.9 79.8 80.2 78.2 79.8 80.0 80.7 80.4 80.2 80.2 80.4 76.9 78.2 79.5 80.9 81.1 80.9 81.0 81.1 114.5 109.1 102.0 113.4 112.3 110.3 115 116 116 115 116 116 118 116 116 118 121 116 119 116 114 114 114 117 116 115 120 116 138 134 128 134 131 128 meters. 3^25 3.64 3.60 3.56 3.88 3.74 .63 .56 .52 .74 .76 .29 3.73 3.69 6.43 6.16 5.88 6.84 7.88 7.41 'Total distance before lunch, 3,793 meters; after lunch, 10,612 meters. *Total distance in about 5 hours of continuous walldng, 23,697 meters. Pulse-rate of subject standing during 5 consecutive minutes immediately after walking, 100, 98, 96, 96, and 98 per minute. 'Total distance in 4 hours and 52 minutes of continuous walking, 22,919 meters. Pulse-rate during 4 minutes soon after the carbohydrate breakfast averaged 73 per minute with the subject sitting; during 4 consecutive minutes immediately after walking it was 80, 79, 80, and 79 per minute, with subject standing. *Total distance before lunch, 6,949 meters; for about 20 minutes after the first period subject was not walking. After lunch, the distance was 9,362 meters. STATISTICS OF EXPERIMENTS. 59 Table 4. — Summary of results in experiments with subject II, during the period from Mar. 16, 1914, to May IS, iSi^— Continued. [Values per minute.] Car- II ^1 Ssl Rais- Date, condition, and time. Dura- tion. bon diox- Oxy- gen. ii era espi on-ri Dis- tance. Steps. ing of ide. <- ft > u-9 < body. May 5. Without food: min . sec . c.c. c.c. meters. meters. Standing, 8'' 24=' a. m 13 7 229 277 0.83 83 19.7 Walking,' gk 42°' a. m 10 3 1011 1124 .90 25.5 106.6 133 4.86 WaUdng,' 10'' 11" a. m 11 5 915 1101 .83 28.0 102.6 127 5.82 WaHring,' 10^ 44'» a. m 10 3 909 1151 .79 28.0 103.6 126 6.06 Protein, 11'' SI" a. m.: Walking,' 11'' ST" a. m 9 25 1136 1298 .87 28.8 112.6 133 7.69 Walking,' 12'' 30'° p. m 10 39 1137 1318 .86 29.0 113.7 133 7.83 Walking,' 1'' IS" p. m 10 32 1035 1208 .86 28.5 106.0 128 6.40 Walking,' 1'' 46" p. m 9 31 1076 1307 .82 29.2 111.3 129 7.76 May 6. Without food: Walking," 9'' 08" a. m 8 65 1902 2068 .92 31.1 140.7 149 9.76 Walking,^' Itf' 31" a. m 9 30 1848 2125 .87 33.4 139.6 148 8.90 Walking.Ml'' 15" a. m 8 12 1796 2279 .79 36.1 142.9 152 8.79 Carbohydrate, 11'' 40" a. m.: Walking,* 12'' 45" p. m 8 36 2058 2147 .96 35.5 143.0 146 7.82 May 10. Without food : Standing, 8'' 34" a. m 11 5 216 246 .87 74 18.4 Walking,' 9'' 44" a. m 8 22 2017 2232 .90 32.8 146.6 153 5.37 Walking,' 10'' 47" a. m 8 14 2101 2384 .88 38.6 146.8 157 6.59 Walking,' 11'' 33" a. m 9 2 2006 2234 .90 37.1 146.6 154 8.02 Protein, 12'' 15" p.m.: Walking,' 12'' 43" p. m 8 14 2017 2244 .90 36.0 146.5 157 7.83 Walking,' 1'' 20" p. m 8 16 2020 2249 .90 40.1 146.9 156 8.28 May 11. Without food: Standing, 8'' 37" a. m 12 47 259 278 .93 85 20.9 Standing, swinging arms,* 9'' 16" a. m 10 8 52 19 456 1860 516 1886 .88 .98 20.9 35.7 146! 6 184 13.45 Rimning,* 10'' 01" a. m Running,* 10'' 49" a. m 8 55 1765 1938 .91 36.8 146.7 183 13.89 Rxmning,' 11'' 35" a. m 8 24 1793 1964 .91 37.9 148.3 181 14.80 Carbohydrate, 12 noon: Running,' 12'' 44" p. m 8 23 1795 1910 .94 36.6 148.7 181 13.95 'Total distance before lunch, 8,126 meters; after lunch, 9,193 meters. During about 40 minutes of the intervals between periods in the afternoon, subject was not walking but sitting. ^otal distance before lunch, 5,571 meters; after lunch, 1,543 meters. Subject perspired very much with this rate of walking and worked with arms and hands much more than at the other speeds. For about 1 hour and 40 minutes of the intervals between periods in the morning he was not walking. 'Total distance before lunch, 6,307 meters ; after lunch, 3,468 meters. During about 1 hour and 20 minutes of the intervals between periods in the morning and about 25 minutes in the afternoon, subject was sitting or walking a little in the room. *rhe subject swung his arms as vigorously as in the fastest walking. 'Total distance before lunch, 6,696 meters; after lunch, 1,729 meters. In running, the subject held arms up in front of body and the only movement of the arms was that of the swaying body. During about 1 hour and 10 minutes of the intervals between periods in the morning subject was not running. 60 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. Table 4 — Summary of results in experiments mth avbjed II, during period from Mar. 16, 1914, to May IS, 19U— Concluded. [Values per minute.] Date, condition, and time. Dura- tion. Car- bon diox- ide. Oxy- gen. ^■ fi' S ^ l-i m 09 >1 oi. 0) " 9 Z "> 2 Dis- tance. Steps. Rais- ing of body. May 12. Without food: Standing, g"" 01™ a. m Running,' Q*" 45"' a. m Running,' lO* 44™ a. m Running,' 11'' 38™ a. m ... . Running,' 12'' 26™ p. m. . . . Running,' l"" 10™ p. m May IS. Without food : Standing, lO"* 20™ a. m Walking,' 11'' 08™ a. m Walking,' 11'' 47™ a. m Walking,' 12'' 31™ p. m Walking,' l"" 16™ p. m Walking,' 2'' 18™ p. m Walking,' 3'' 06™ p. m May 14. Without food : Standing, 8'' 32™ a. m Walking,' 9'' 33™ a. m Running,' 10'' 28™ a. m Running,' 11'' 20™ a. m Running,' 1 1"" 58™ a. m . . . . Running,' 12'' 35™ p. m May 15. Without food : Standing, swinging arms. Si" 52™ a. m Running,' 9'' 57™ a. m Running,^ 10'' 37™ a. m Running,' 111" 21™ a. m. . . . mm. sec. 13 15 8 19 8 55 8 28 10 12 10 6 12 27 13 30 12 49 12 1 13 10 12 29 13 2 13 53 7 41 7 58 8 51 9 2 9 41 12 8 9 40 8 45 8 20 c.c. 226 1983 1758 1723 1752 1734 213 711 734 639 597 767 753 209 2046 1997 1646 1625 1701 554 1735 1727 1647 c.c. 293 2194 1843 2011 2050 2080 257 851 942 826 789 998 980 257 2281 2272 1867 1900 1982 643 1967 1978 1947 0.77 .90 .95 .86 .85 .83 .83 .84 .78 .77 .76 .74 .77 .81 80 .90 .88 .88 .86 .86 .86 .88 .87 .85 82 79 20.4 34.9 37.8 39.2 39.2 40.8 20.2 25.9 27.9 27.0 25.4 28.0 28.5 20.0 38.6 37.5 38.3 38.8 40.1 22.3 38.6 37.3 38.9 meters. 147. i 148.4 148.1 148.1 148.7 82.6 89.7 80.1 76.6 93.3 91.9 146.4 146.5 147.2 147.6 148.1 144.7 148.4 147.7 191 183 182 184 176 117 120 112 106 117 118 153 168 183 182 183 182 183 184 meters. 15.40 14.09 15.40 14.64 15.37 3.76 4.51 3.29 2.84 5.25 5.15 7.59 15.44 12.47 12.34 11.09 12.48 12.14 13.13 'The total distance was 10,111 meters. During 2| hours of the intervals between periods subject was sitting or walking a Uttle in the room. 'Total distance, 21,724 meters, the walking being continuous. •Total distance, walking and running, 9,432 meters. During about 2 hours and 11 minutes of the intervals between periods, subject was sitting or standing. 'Subject swung his arms as vigorously aa in the fastest walking. 'Total distance, 5,711 meters. During about 58 minutes of the intervals between periods, subject was not running or walking. DISCUSSION OF RESULTS. 61 DISCUSSION OF RESULTS. BASAL VALUES. In any study of walking in a horizontal direction, two main prob- lems present themselves: first, the variations in the energy requirement of different individuals for walking varying distances at varying velocities, and second, the actual energy requirement for transporting the body-weight or a superimposed load in a horizontal direction, i. e., the calorie output per horizontal kilogrammeter. As has already been noted, the mechanical processes incidental to walking, even at a moderate pace, usually involve some extraneous muscular movements apparently not directly connected with the motion of forward progression, such as the more or less vigorous swinging of the arms and a not inconsiderable raising of the body-weight with each step. Since these extraneous muscular movements do not necessarily have an effective value in transporting the body-weight or the super- imposed load in a horizontal direction, the problem of finding the calorie requirement for such transportation of weight becomes an exceedingly complicated one. If the principle of the deduction of basal values for determining the energy required to move 1 kilogram 1 meter in a horizontal direction may be legitimately employed, and this has been the method adopted by all investigators, it becomes an important point as to what should properly be considered as the basal maintenance metaboUsm in experi- ments when the subject is walking in a horizontal direction. As will be seen by reference to the summary of previous researches given in table 1, investigators have varied considerably in their usage in this respect. Certain members of the Zuntz school have almost invariably employed the resting (lying) metaboUsm obtained with the subject in the post-absorptive condition. The values found with the subject sitting or standing quietly have also been used as a basis in determining the metabolism due to the muscular activity of walking. Still another basal value which may be considered is that found when the subject is standing and moving his arms in a manner similar to that employed in more or less rapid walking. Finally it may even be possible to use the metaboUsm determined during slow walking for a basal value to be deducted from the metaboUsm found while the subject was walking at a rapid rate. The vaUdity of this assumption will be examined later. Bearing in mind the experience of Benedict and Cathcart in attempt- ing to secure a suitable base-line for their bicycle rider, we considered it desirable to study our subjects in varying positions. The positions selected have been outlined in a preceding section. Since we were to make a large number of observations upon each subject, we concluded that the period of adjustment to the type of the experiment would be so short that a moderately constant value could be found in the standing 62 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. position, and hence the greater part of our control data was obtained in experiments with the subject standing in a relaxed position. Although values for comparison are available for other standing posi- tions and for the subject sitting, no studies were made by us with the subject in complete muscular repose, i. e., lying upon a couch, in the post-absorptive condition. Obviously it is first necessary to examine the data thus secured to determine the suitable basal values to be deducted from the total metaboUsm while walking, such a critical examination being of funda- mental importance. Since the total metaboUsm is made up of that for maintenance, i. e., the basal value plus that for forward progression, it is evident that the larger the value deducted for maintenance — as, for instance, the value obtained with the subject standing as com- pared with the subject lying on a couch — the smaller will be the result obtained for the energy requirement for the work of forward pro- gression. BASAL METABOLISM OF SUBJECT L As a result of a consideration of the experimental experience of others, it was decided that in the preUminary observations in this investigation, which were made with subject I by Dr. Carl Tigerstedt, the basal value selected should be that necessary for maintenance while the subject was standing quietly without support. Accordingly, during November and December 1913 many observations were made of the subject in this position. Subsequently it appeared that more information was desirable with regard to the normal increment in metabolism due to a change from the lying position to the standing position, but as a study of this phase of the question was wholly incidental to the main problem of the metabolism during forward progression, the experimental routine did not permit such observations. We have, however, the results of a study made in December 1914 by Mr. H. L. Higgins, of the Laboratory staff, upon the lying and standing metabolism of subject I, which were kindly supplied us for comparison with our values. Unfortunately the data thus obtained are more or less fragmentary and must therefore be considered as only subsidiary to the results of the larger investigation. The Thiry valves and Tissot spirometer and mouthpiece were used in this study, both the resting metaboUsm and the standing metaboUsm being observed with the subject in the post-absorptive condition. We have no data as to the influence of food upon the resting metaboUsm of this subject. The results" obtained by Mr. Higgins are recorded in table 5. These values are typical of a large number secured by this observer in another research with subject I, who shows a remarkably uniform basal resting metaboUsm. As will be seen from the table, the heat output per minute of this individual in the lying position was on December 2 and 5, 1914, 1.23 and 1.22 calories respectively. DISCUSSION OF RESULTS. 63 The values found by Dr. Carl Tigerstedt for subject I in the standing position, which was the only position used by Dr. Tigerstedt in deter- mining the basal metaboUsm of this subject, are given in table 6. Since the walking experiments were made in the morning and in the afternoon and with and without food, the basal values were also obtained in both the post-absorptive condition and after the taking of food. The appa- ratus used for determining the respiratory exchange was the same as that employed for the walking experiments. Table 5. — Metdbolism of subject I in the lying and standing positions in experiments vrithmii food, on Dec. S and Dec. 6, 1914. [Observations made by Mr. H. L. Higgins with Thiry valves and Tissot spirometer. Values per minute.] Date and time. Dura- tion. Carbon dioxide. Oxygen. Heat output (com- puted). Pulse- rate. Respira- tion-rate. Dec. 2. Lvine 8** 53™ a. m mins. 10 8 8 10 10 c.c. 210 230 224 208 196 c.c. 249 297 287 256 262 cats. 62 84 85 61 61 22.0 25!7 24.8 24.1 Standing, 9'' IS"" a. m Standing, Q"" 29™ a. m Lying, lO"" 15"" a. m Lvine 10^ 35™ a. m Average, lying Average, standing .... Dec. 5. Lvine 9** 02™ a. m 205 227 256 292 1.23 1.39 61 84 23.6 25.7 10 8 8 9 219 220 217 221 246 269 268 249 62 95 90 57 23.4 25.8 26.4 24.8 Standing, 9'' 27™ a. m Standing, 9^ 44™ a. m Ljring, 10'' 25™ a. m Average, lying Average, standing .... 220 219 248 269 1.22 1.29 60 93 24.1 26.1 As will be seen from table 6, the average post-absorptive values are reasonably constant throughout the entire period from November 28 to December 20. A striking exception is shown on December 5, when the average carbon-dioxide excretion was 236 c.c. per minute and the oxygen consumption 334 c.c. per minute. The protocols for this experiment show that "there was no sleep the night before, but much alcohol." The pulse-rate was especially high on this particular day. The general average for the observations without food shows a carbon- dioxide excretion of 223 c.c. per minute and an oxygen consumption of 280 c.c. per minute. This corresponds to a heat output of 1.34 calories per minute. The pulse-rate of this man was extremely high, averaging 94 beats per minute; the average respiration rate was 21.2 per minute. All of these experiments were made with subject I standing with muscles relaxed. The general uniformity of the data shows that for the most part the subject must have assumed a standing position with a relatively constant muscle strain, although variations are found in 64 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. the experiments without food from 1.28 to 1.57 calories per minute. Excluding the extraordinarily high value of December 6, which is at least in part explained by the alcohoUc excesses of the preceding night, the uniformity in the average values is for the most part striking. It is also of interest to note that the average value obtained for all of the standing experiments without food during December 1913, i. e., 1.34 calories, is the same as that of the average of two standing experiments made by Mr. Higgins in December 1914. (See table 5.) It will be seen, therefore, that for many purposes an average basal value may be used for this subject with propriety and that it may be assumed that when standing quietly and in the post-absorptive condition, his average heat output would correspond to 1.34 calories per minute. Table 6. — Metabolism of subject I in the standing position. [Observations made by Dr. Carl Tigerstedt. Average values per minute.] Date. Carbon dioxide. Oxygen Heat^ output (com- puted). Pulse- rate. Respira- tion-rate. 1913. Without food: Nov. 28 . . C.C. 222 224 220 224 214 214 236 206 222 223 223 229 239 226 c.c. 285 282 279 286 272 267 334 271 275 286 271 263 279 270 cals. 1.36 1.35 1.34 1.37 1.30 1.28 1.57 1.29 1.32 1.37 1.31 1.29 1.36 1.31 92 93 92 89 92 90 105 91 89 101 91 86 98 103 17.1 18.5 19.3 20.7 21.8 22.6 23.2 21.2 22.2 22.2 21.3 21.3 23.0 22.9 29 Dec. 1 2 3 4 5 8 15 16 17 18 19 20 Average, without food .... After breakfast: Nov. 26 223 280 1.34 94 21.2 281 248 273 332 341 335 1.61 1.61 1.61 112 84 22.9 18.2 Dec. 9 After dinner: Nov. 26 Average, with food 267 336 1.61 98 19.8 When the superimposed factor in a metaboUsm experiment is suf- ficiently great to increase the metabohsm several hundred per cent, as is especially the case in muscular-work experiments, the use of a pre- viously estabhshed average basal value is least hable to objection. Although with this particular subject we may assume that we have a fairly well-estabUshed basal value, as a matter of fact, to eliminate possible wide variations we rarely used this average figure and the absolutely determined basal value for each day, which was usually available, was given the preference. DISCUSSION OF RESULTS. 65 INFLUENCE OF FOOD AND BODY POSITION. In this preliminary study of subject I, we have two main factors to be considered, namely, the influence of food and of body position. Influence of food. — We have only three experiments in which obser- vations were made of the standing metabohsm of subject I after the ingestion of food. Singularly enough, the average on each of these three days shows that the heat output after the ingestion of food when measured with the subject in the standing position was the same in all of the experiments, i. e., 1.61 calories. (See table 6.) As we have no basal value without food which was determined on these three days, we must be content to note that the average basal value of 1.34 calories was increased 0.27 calorie per minute as the result of the ingestion of food, an increment of approximately 20 per cent. It is thus clear that prior to the actual walking tests after meals, the subject has a noticeably larger basal metabolism than prior to the walking experiments in the morning in the post-absorptive condition. The intelhgent use of these basal values after food is a subject of subsequent discussion. Influence of position. — In his experiments with subject I in December 1914, Mr. Higgins also obtained data regarding the metabohsm of this subject in the standing position. Any comparison of these values with those obtained by us a year earlier for the same position would be a violation of a fundamental principle in the computation of metabo- hsm experiments, notwithstanding the facts that the difference in the body weight is but 2 kilograms and that the values obtained for the metabohsm in both cases were extraordinarily constant. Neverthe- less, in the hope of throwing some Ught on the probable increment in the metabohsm to be expected from this man, when standing, we have included in table 5 (page 63) Mr. Higgins's results of December 1914 for the standing position. The observations on December 2 show an average heat output for the standing position of 1.39 calories per minute, but three days later (December 5), when the conditions were apparently identical, the heat output was but 1.29 calories per minute. Using the average values for the resting (lying) metabohsm on the same days, we note an increase in the heat output on December 2 from 1.23 calories to 1.39 calories and on December 5 from 1.22 calories to 1.29 calories, the average increase being from 1.225 calories to 1.34 calories. This is equivalent to an average increment of 0.115 calorie, or approximately 9 per cent, due to the changing of the position from lying to standing. Since the average value of 1.225 calories for the metabohsm in the lying position is true not only for the two days referred to but also for an extensive series of experiments made by Mr. Higgins on this subject in 1914, and since the average of the two standing experiments made by the same observer is identical with the average of the long series of standing experiments made one year previous by Dr. Tigerstedt, we may properly maintain 66 ENERGY TRANSFORMATIONS DURING HORIZONTAL "WALKING. that with this subject a change in the body position from lying to standing is accompanied by an increase of approximately 9 per cent in the total metaboUsm. BASAL METABOLISM OF SUBJECT IL With the beginning of the prolonged research on the professional athlete subject II (the bicycle-riding subject M. A. M. of Benedict and Cathcart), we planned definitely to secure much more extended data as to the metaboUsm during various standing positions, to supplement the preliminary data obtained with subject I by observations made on subject II after the ingestion of food, and to determine in addition the metaboUsm of the subject in the sitting position. These studies were whoUy incidental to the main problem of studying the motion of forward progression and were made primarily with a view to iUum- inating the possible selection of the most advantageous and the most scientifically sound base-Unes. At the time the observations were made, we were not certain of their ultimate use in the final computations, so they may properly be considered as a study of the metaboUsm in the standing and sitting positions, with and without food. METABOLISM IN THE LYING POSITION. The observation of the post-absorptive metaboUsm of this subject in the Isang position was confined to one experiment which was made through the kindness of Mr. L. E. Emmes, of the Laboratory staff, but a large number of basal values for this position were also available as a result of the previous study of the subject by Cathcart. The results obtained in the experiment made by Mr. Emmes and an average value for the results obtained by Cathcart are given in table 7. Table 7. — Metabolism of subject II in the lying position in experiments mthout food. [Observations made by Mr. L. E. Emmes. Values per minute.] Date and time. Carbon dioxide. Oxygen. Heat- output (com- puted). Pulse- rate. Respiia^ tion-rate. Total. Per kilo- gram. Total. Per kilo- gram. Apr. 18, 1914:1 8''24'»a. m C.C. 198 186 196 193 c.c. c.c. 226 229 229 232 c.c. cals. 57 56 58 56 15.5 15.6 16.6 16.3 8 54 a. m 9 17 a. m 9 48 a. m Average 193 205 2.86 3.12 229 242 3.39 3.67 1.11 1.17 57 63 15.9 20.0 Dec. 7, 1911, to Apr. 16, 1912^* 'The duration of each period on April 18 was about 15 minutes and 3 seconds. 'Average of experiments made by Benedict and Cathcart. See Carnegie Inst. Wash. Pub. 187, 1913, p. 78. DISCUSSION OF RESULTS. 67 The four periods of the respiration experiment made on April 18, 1914, by Mr. Emmes show an excellent agreement which is character- istic of this subject. Although his nude body-weight was 1.6 Mlograms greater than at the time of the study of Cathcart, being 67.5 kilograms on April 18, 1914, and averaging 65.9 kilograms during the previous research, the heat output per minute was but 1.11 calories per kilogram per minute as compared to the average of the earher observations of 1.17 calories per kilogram per minute. While it is possible that direct averaging in this instance is a mathematically unsound procedure, we may state that the resting basal heat output of subject II in the post- absorptive state may be considered to be, in round numbers, 1.14 calories. METABOLISM IN THE SITTING POSITION. In a further effort to secure all possible data for establishing various scientifically sound base-Unes, we made a special study of the metabo- lism of subject II while he sat comfortably in a chair. Three observa- tions were made with the subject in the post-absorptive condition, four after a Ught breakfast, and five after a heavier midday meal. The data are sujHiciently extended to permit their classification accord- ing to the character of the meal and such a classification is used in presenting the results of the study of the sitting metabolism in table 8. Table 8. — Metabolism of subject II in the sitting position. [Values per minute.] Date. Carbon dioxide. Oxygen. Heat- output (com- puted). Pulse- rate. Respira- tion-rate. 1914 Without food: Mar. 25 27 Apr. 3 Average After light meal : Mar. 16 16 18 19 Average After heavy meal: Mar. 16 17 18 19 25 Average c.e. 192 195 219 C.C. 260 234 245 cats. 1.23 1.13 1.20 59 58 67 18.0 17.6 19.4 202 246 1.19 61 18.3 218 203 233 242 281 256 286 282 1.34 1.23 1.38 1.37 67 18.7 18.3 19.2 18.7 224 276 1.33 18.7 250 213 248 259 272 304 272 297 322 316 1.47 1.30 1.44 1.55 1.54 77 73 19.9 18.1 19.3 21.0 19.6 248 302 1.46 19.6 68 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. With subject I no such classification could properly be made and, indeed, the observations after the ingestion of food were but few in number. The metabolism with and without food are further compared in table 9, to which is added the actual increase in the units of measure- ment commonly employed for the various factors and the percentage increase. From this table we see that as the result of a light meal the carbon dioxide was increased 10.9 per cent, the oxygen consumption 12.2 per cent, and the heat output 11.8 per cent. After the heavy meal, the percentage increase was essentially the same for all three factors, i. e., 22.8 per cent. This percentage increment in the metabolism of subject II after a heavy meal is almost identical with that noted for subject I in the few experiments made with him. Table 9.— Metabolism of subject II in the sitting 'position with and without food [Average values per minute.] Condition. Carbon dioxide. Oxygen. Heat output (computed). Pulse-rate. Respira- tion-rate. Without food After light meal Increase -with food .... After heavy meal Increase with food .... C.C. 202 224 (22 \ 10.9 p. ct. 248 f46 1 22.8 p. ct. c.e. 246 276 r 30 I 12.2 p. ct. 302 r 56 1 22.8 p. ct. cals. 1.19 1.33 r .14 111.8 p. ct. 1.46 r .27 122.7 p. ct. 61 •67 I ^ \ 9.8 p. ct. >75 f 14 \ 23.0 p. ct. 18 18 19 3 7 6 'The pulse-rate was obtained in only one of the four periods after the light meal; in but two of the five periods after the heavy meal. (See table 8.) One serious objection to this type of comparison, however, and an objection that measurably lessens the mathematical value of the incre- ment due to food, is the fact that the periods after the midday meal followed the walking experiment in the morning, in which there was considerable muscular activity. As a matter of fact, the long-con- tinued walking experiments, in which the walking was done at a rapid rate and the muscular activity was very severe, were not carried out until later in the spring of 1914, and hence in all probability we have not here to consider any great after-effect of the muscular activity of the morning which persisted throughout the experiment in the after- noon. Nevertheless, it is quite possible that at least a part of the excess metaboUsm after the midday or "heavy" meal may be explained by the fact that the subject had done considerable walking in the fore- noon, while no such muscular activity preceded the after-breakfast experiments. Of special significance here is the apparent striking relationship between the percentage increment in the total metaboUsm and the percentage increment in the pulse-rate. The increase in the heat- DISCUSSION OF RESULTS. 69 output after a light meal was 12 per cent and that in the pulse-rate 10 per cent, while after a heavy meal the increase in the heat output was 22.7 per cent and in the pulse-rate 23 per cent. This Laboratory- has frequently emphasized in its pubUcations the important relation- ship between the pulse-rate and the metabohsm, the statements usually implying that an increase in the pulse-rate is simultaneous with an increase in the metabolism. Recently, in pubHshing the results of observations on a man who fasted 31 days,^ it was pointed out in com- paring the metabohsm of the subject asleep and awake that the incre- ment in the pulse-rate for the subject awake was directly proportional to the percentage increase in the metabohsm. While it is by no means maintained that the increment in the pulse-rate for subject II was in direct proportion to the increase in the metabohsm, it is of special interest to point out this second striking percentage relationship between the pulse increment and the metabohsm increment, for such a quantitative relationship, if not partaking of the nature of a physio- logical law, at least serves to emphasize in a striking manner the im- portance of records of the pulse-rate in all metabohsm experiments. Occasion is here taken to note that a common error is found with writers who misapply the measurements of the pulse-rate in attempting to compare the pulse-rate of one person with that of another. Our experience in this Laboratory points to no relationship between the pulse- rate of two individuals, although the records for a single individual are frequently surprisingly proportional to the metabohsm. COMPARISON OF THE METABOLISM IN THE LYING AND SITTING POSITIONS. Although observations were made of the metabohsm of subject II in the post-absorptive condition for both the sitting and the lying posi- tions, unfortimately the studies of the metabohsm in the lying position were made some three weeks later than those of the metabohsm in the sitting position, hence a comparison of the results is not free from criti- cism. If, however, we compare the average values found for the metabohsm in the lying position on April 18, 1914 (see table 7), with the average of the values for the metabohsm in the sitting position obtained on March 26 and 27 and April 3, 1914 (see table 8), we find a percentage increase in the carbon-dioxide output with the subject sit- ting over that with the subject in the lying position of 4.7 per cent, in the oxygen consumption of 7.4 per cent, in the heat production of 7.2 per cent, and in the pulse-rate of 7.0 per cent. According to these incom- plete data, therefore, the increase in the metabohsm with the subject sitting over that with the subject in the lying position is approximately 7 per cent.^ We may also note here the agreement between the incre- ment in the pulse-rate and that of the total metabohsm. iBenedict, Carnegie Inst. Wash. Pub. 203, 1915, p. 351. ^See, also, Emmes and Riche, Am. Journ. Physiol., 1911, 27, p. 406. 70 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. METABOLISM IN VARIOUS STANDING POSITIONS. With subject I no attempt was made to vary the character of the standmg experiment, but with subject II numerous observations were made both with and without food and with the subject standing in various positions. A large majority of the experiments, i. e., 53 experi- ments, were made when the subject was standing quietly in a relaxed position. Eight experiments were made with the subject standing in a relaxed position, but resting the hands upon a staff. Eight experiments were made with the subject leaning against a support, and finally 10 experiments were carried out with the man standing in the position of "attention." Since later we noted that there was considerable extraneous muscular activity incidental to walking at a rapid rate, which consisted for the most part of vigorous arm-motion, two experi- ments were made on May 11 and 15 in which the post-absorptive metaboHsm was measured with the subject standing still but moving his arms vigorously, simulating as nearly as possible the movement of the arms in rapid walking. The results of the standing experiments with the subject in the post- absorptive condition will first be considered and are given in table 10. Since our interest is chiefly with the total heat-output, we may properly compare the metaboUsm in the various standing positions upon this basis. When the subject stood in a relaxed position, his heat output varied from the very low value of 1.12 calories to 1.40 calories per minute, the average value being 1.25 calories per minute. When he stood in a relaxed position, but with his hands resting upon a staff, the average heat-output was essentially the same as in the first position, namely, 1.26 calories per minute; but when he leaned against a support there was a perceptible diminution in the heat-output, which then averaged but 1.18 calories per minute. It is, however, somewhat irregular to compare an average value for the relaxed position obtained from 26 experiments with the average values for other positions obtained from only 3 experiments; the sHght changes in the metabohc level shown by these averages are therefore probably without significance. Essen- tially the same may be said of a comparison of the values obtained for the relaxed position with those obtained for the position of attention when the heat output increased only to 1.30 calories. Our general impression was that this subject did not maintain, to any great degree, a rigid position of "attention." From the foregoing comparison it is clear that the experimental evidence does not warrant the deduction that there is a measurable difference in the metabohsm for any of the various standing positions in which the metabolism was studied, such as standing in a relaxed position, with the hands on a staff, with the subject leaning against a support, or standing in the position of "attention." DISCUSSION OF RESULTS. 71 When instead of the muscular repose of these standing positions the subject swings his arms vigorously, as would be done in rapid walking, we note a great increase in the metabohsm. Unfortunately there is not a particularly satisfactory agreement in the values obtained for the two experiments under these conditions, for on May 11 the heat-output per minute was 2.53 calories, or essentially 100 per cent greater than when the subject stood quietly, while on May 15 the metabohsm was 3.13 calories or approximately 150 per cent greater than the metab- ohsm of the subject when standing at rest. Table 10. — Metabolism of subject II standing in different positions in experiments without food. [Values per minute.] Date. 1 S o P4 Date. 1 1 1 1914 Standing, relaxed: Mar. 20 31 Apr. 1 6........ 7 8 9 10 14 15 16 17 21 22 23 24 25 27 28 May 5 10 11 12 13 14 Average ex. 218 209 213 218 r214 1198 204 202 212 223 214 214 205 207 208 198 192 217 221 229 215 259 226 213 209 c.c. 275 260 266 246 257 236 265 228 257 245 252 258 265 242 251 252 264 245 274 277 246 278 293 257 257 cats. 1.32 1.25 1.28 1.21 1.24 1.14 1.26 1.12 1.24 1.21 1.23 1.251 1.28 1.18 1.21 1.22 1.15 1.24 1.20 1.32 1.34 1.20 1.38 1.40 1.24 1.24 71 71 75 74 82 90 82 78 94 74 90 85 78 84 74 82 79 77 82 83 74 85 82 79 80 1914 Standing, staff: Mar. 24 26 Apr. 3 Average Standing, support: Mar. 24 26 31 Average Standing, attention: Mar. 25 27 31 Average Standing, swinging arms ■? May 11 15 c.c. 217 223 224 c.c. 250 253 278 cats. 1.22 1.24 1.34 82 80 77 221 260 1.26 80 196 201 214 256 238 239 1.22 1.15 1.17 79 79 77 204 244 1.18 78 206 219 233 265 272 272 1.27 1.31 1.33 72 71 75 219 270 1.30 73 455 554 516 643 2.53 3.13 214 258 1.25 80 ^The average respiratory quotient of 0.83 is assumed in computing heat-output on this day. ^he subject stood swinging his hands and arms as vigorously as in the most rapid walking. In the subsequent computations several basal values may be em- ployed. The most obvious are those obtained with the subject in the lying position and, as has aheady been seen, we have the results of two series of measurements which were made with an interval of approxi- mately two years. We have also used the value obtained with the subject in the standing position. With subject I, three periods with 74 ENBBGY TRANSFORMATIONS DURING HORIZONTAL WALKING. of "attention." All of these values were obtained after a heavy meal, with the diet uncontrolled. Recognizing again the probable influence of the preceding meal upon the individual values, we see, nevertheless, that the evidence points towards a practically constant metabolism independent of the variations in the standing position under approxi- mately similar conditions. Tablb 12. — Metabolism of svbjecl II standing in digereni positions after a heavy meal. [Values per minute.] Date and conditions of experiments. f 1 1^ II M-2. 1 f Date and conditions of experiments. 1 r 6 IS II 1 1914 Diet uncontrolled : Standing, relaxed: Mar. 16 19 20 21 31 Apr. 1 3 Average Standing, staff: Mar. 24 26 30 Average Standing, support: Mar. 24 26 28 Average Standing, attention : Mar. 21 25 27 31 Average Average of all periods c.c. 261 247 274 275 277 283 244 c.c. 319 303 318 355 345 319 293 cats. 1.54 1.46 1.55 1.70 1.66 1.57 1.42 86 87 96 89 86 81 1914 Diet controlled : Standing, relaxed : Protein — Apr. 6 7 8 23 24 Average .... Carbohydrate — Apr. 9 10 14 15 25 27 Average Fat- Apr. 16 17 21 22 Average .... c.c. 241 217 230 223 219 c.c. 296 289 275 254 299 caZs. 1.43 1.37 1.33 1.24 1.41 81 94 226 283 1.36 88 266 322 1.56 88 245 264 242 267 274 294 295 301 295 277 324 326 1.43 1.47 1.42 1.38 1.58 1.61 76 86 90 83 83 75 289 274 318 356 317 339 1.71 1.55 1.69 95 94 95 294 337 1.65 95 254 254 280 301 289 314 1.46 1.42 1.54 92 93 85 264 303 1.48 82 239 231 220 227 292 281 301 299 1.41 1.36 1.42 1.42 98 99 91 93 263 301 1.47 90 270 292 294 265 295 309 298 326 1.46 1.54 1.50 1.57 94 95 95 89 229 293 1.40 95 280 307 1.52 93 274 317 1.55 91 During the observations made with the specially controlled diets, the subject stood in the relaxed position, without leaning upon a staff or against a support. These show an average value of 1.36 calories for the protein diet, 1.48 calories for the carbohydrate diet, and 1.40 calories for the fat diet, values not strikingly unlike those found on the days with an uncontrolled diet. We may then with propriety average DISCUSSION OF RESULTS. 75 the results obtained for the four groups of experiments in the relaxed position, i. e., one group with an uncontrolled diet and three with con- trolled diets, and state that after the midday meal our subject showed with considerable constancy an average heat-output of 1.45 calories per minute. From the values obtained in the experiments with the subject stand- ing after a heavy meal, basal values were computed for use in calculat- ing the increment due to walking under the same conditions of diet. Considering first the results obtained with an uncontrolled diet, we averaged the values determined with the subject standing in the four positions, i. e., relaxed and unsupported, relaxed with the hands resting upon a staff, relaxed and leaning against a support, and the position of "attention." The averages found were 274 c.c. for the carbon-dioxide production, 317 c.c. for the oxygen consumption, and 1.55 calories for the heat-output. These averages were used for basal values in calculat- ing the increase in the metaboUsm due to walking for all of the experi- ments following a heavy meal with an uncontrolled diet. The average value found on the days with a diet rich in protein, namely, 1.36 calories, was used as a base-line for the walking experi- ments on May 5 and 10, as the standing metabolism was not determined on those days after the protein meal. A heavy protein meal was also taken on April 6, 7, 23, and 24, and both the standing metabolism and the metaboUsm during walking were determined. The standing values for the individual days were therefore used as the basal values for com- puting the increment due to walking. The average value for the carbohydrate days was not used as a basal value for any of the walking experiments. For the walking experi- ments following the midday meal on May 4, 6, and 11, an average of the values obtained on April 25 and 27, i. e., 1.59 calories, was used as a base-line in the computations, as the diets were considered to be more nearly comparable on those days. As has previously been stated, this basal" value was also used for the walking experiment following the carbohydrate breakfast on April 29. For the experiments on April 9, 10, 14, 15, 25, and 27, the standing metabolism determined for that day was used as a basal value for the respective walking experiments which were subsequently made on that day. On days when the diet was excessively rich in fat, the standing metabolism was always determined before the walking experiment and the values found were used as base-Unes for the individual days, no use being made of the average value. Conclusions Regarding Influence op Food upon the Standing Metabolism. The average value for the metaboUsm in the standing experiments without food, as shown in table 10, was approximately 1.25 calories. After the light morning meal, the metaboUsm was 1.43 calories (see 76 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. table 11) and after the heavy noon meal 1.45 calories. There was, therefore, an average increment of approximately 0.2 calorie or 16 per cent, due to the ingestion of the food. This value is strikingly m agree- ment with that found with subject I, and we may accordingly state that the basal metaboUsm of both subjects used in the research was increased approximately 16 per cent by the ingestion of the kinds and amounts of food consumed. This has an important bearing upon the subsequent calculations, for it is obviously impossible to make an intelli- gent comparison of the standing values obtained with individuals in the post-absorptive condition with those obtained after the ingestion of food. Finally, it should not be forgotten that a part, at least, of the incre- ment ascribed here to the ingestion of food, particularly after the noon meal, may be due to a shght after-effect of the muscular activity of walking in the forenoon experimental periods. On the other hand, there is such a close agreement between the increment in the metabolism after the Hght meal, which was invariably taken before the morning walking periods, and the increment in the metabolism after the midday meal, that it would imply that the influence of the morning work must have been extremely small. METABOLISM DURING WALKING. The preliminary study of the mechanics of position incidental to walking was obviously only for the purpose of throwing light upon the various mechanical processes involved in walking and the metabolism essential thereto. The primary purpose of the research was to study the actual metabolism during the motion of forward progression. WALKING EXPERIMENTS WITH SUBJECT I. The first series of walking experiments was made by Dr. Tigerstedt with subject I in the month of December 1913. These experiments were all carried out under practically the same conditions, — that is, following two or three periods when the subject stood in a relaxed position during which time the basal standing metaboUsm was meas- ured. The body-weight of the subject was a practically constant factor, as it did not alter materially throughout the month. An attempt was made to maintain an essentially constant speed of the treadmill, the rate averaging not far from 76 meters per minute. In a few of the experiments, the speed was lowered to about 65 meters per minute, but in no instance did it exceed 80 meters per minute. Under these conditions we may consider that the study was based upon a constant speed and that accordingly all values are strictly comparable so far as the velocity is concerned. The walking experiments con- sisted of two or three and occasionally fo\ir periods per day. On two days walking experiments were made after the ingestion of food. DISCUSSION OF RESULTS. 77 The results of all of the walking experiments with subject I are given in abstract in table 13. Inasmuch as the experience of previous investigators has shown that the variations in velocity play an impor- tant r61e in the total energy transformations, we have recorded here the distance per minute which the subject walked and have likewise computed the number of horizontal kilogrammeters by multiplying the distance per minute by the body-weight. From the kymograph records it was possible to find the average height to which the body was raised at each step by obtaining the average height of each recorded movement; by multiplying this average height by the number of steps, the total distance that the body was moved in a vertical direction could be computed with reasonable accuracy, thus giving a component for the analysis of the various factors involved in walking. The measurements of the distance that the body was elevated per minute are recorded in column d. The total heat-output was calculated from the gaseous exchange not only for the walking periods but also for the standing periods preceding the walking; these values are recorded in column e. The increase due to walking was obtained by difference and the energy required to move 1 kilogram 1 meter in a forward direction, i. e., 1 horizontal kilo- grammeter, is recorded in gram-calories in column g. These latter values are obviously the most important result of this study. EXPERIMENTS WITHOUT FOOD. Considering first the values without food, it will be seen that the heat-output per horizontal kilogrammeter varies from a minimum of 0.446 gram-calorie to a maximum of 0.637 gram-calorie, but the large majority of the values lie close to the average value of 0.507 gram- calorie. A general inspection of these results shows indications of a periodicity or rhythm in the efiiciency of this subject in walking. Thus, low values are found on December 2 and 3, which are followed by a group of high values continuing until December 22, while the last three experimental days that the subject was without food give dis- tinctly lower values than those found in the preceding group. There is not, however, sufficient regularity in this rhythm to indicate the effect of training of the subject or in the use of the treadmill. To be sure, the first value, that of November 29, is considerably higher than the average and it is probably fair to assume that the high value of 0.637 gram-calorie found on December 15 was more or less accidental. On the other hand, we see no reason why these values should be rejected and the figures are sufficiently numerous to make it immaterial whether or not these or any other particular values are discarded so far as the influence upon the general average is concerned. While it may appear that the low values found on the last three days would imply the effect of training, it should be stated at the outset that this subject was a 78 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. Table 13. — Increase in heat-output during walking in experiments with subject I. [Observations made by Dr. Carl Tigerstedt.] Date and condi- tions of experi- ments. (o) Body- weight with clothing. (6) Distanof per minute. (c) , Horizonta] kilogram- meters. (0X6). (d) Raising of body per minute. Heat-output (computed). (e) 1 Total, per Increase over standing. (/) Total. (0) Per horizontal, kilogrammeter minute. (f^c). 1913 Without food. Nov. 29: kUos. meters. meters. cals. cals. gm.'cal. Standing*. . . 75:1^ 1.35 Walking 72.50 5345 3!l5 4.41 3!66 0.562 Dec. 1: Standing 1.34 Walking 72! 60 76!o 6^518 3!26 4.10 2.76 !5o6 76.6 5,661 3.36 4.35 3.01 .641 Dec. 2: Standing 1.37 Walking 73.12 76.2 si 572 3.26 4.03 2.66 .477 76.8 5,616 4.62 4.10 2.73 .486 Dec. 3: Standing . . . . 1.30 Walking. . . . 72.80 76.6 5,569 3.57 4.07 2^77 .497 77.2 5,620 3.62 4.07 2.77 .493 77.5 5,642 3.68 4.19 2.89 .512 Dec. 4: Standing 1.28 Walking 73.61 77!7 5^673 3.41 4.19 2191 isis 77.7 5,673 3.68 4.22 2.94 .618 78.2 6,709 3.67 4.14 2.86 .501 Dec. 5:» Standing .... 1.67 Walking. . . . 72! 51 ■Ts!?!^ 5, 489 3!57 4.43 2.86 .521 78.2 5,670 4.10 4.50 2.93 .517 79.0 5,728 4.10 4.68 3.11 .543 Dec. 8: Standing .... 1.29 Walking 72.33 76.5 5, '533 3!52 4.02 2.73 !493 78.6 5,685 3.78 4.29 3.00 .528 79.3 6,736 4.04 4.27 2.98 .520 79.7 5,766 4.16 4.39 3.10 .538 Dec. 15: Standing .... 1.32 Walking.... 72.10 74.2" 5,'356 3.68 4.73 siii .637 75.6' 6,451 3.83 4.33 3.01 .562 76.3 6,601 3.94 4.24 2.92 .631 76.3 5,501 3.99 4.20 2.88 .624 Dec. 16: Standing .... .... 1.37 Walking 72.05 75.1 5,'4ii 3.89 4.30 2.93 .541 76.7 5,526 3.99 4.18 2.81 .509 77.6 6,591 4.20 3.91 2.54 .454 Dec. 17: Standing 1.31 Walking 72.82 76.6" 5,'462 3. '73 3.99 2.68 .491 76.4 6,563 3.89 4.12 2.81 .605 78.1 5,687 4.16 4.20 2.89 .508 78.6 5,724 4.16 4.33 3.02 .628 ^Computed from the average elevation of the body per step aa obtained from the kvmoeraDh records and the steps per minute. See table 3 for information regarding number of steps 'Usually three periods with the subject standing preceded the walking experiment '"No sleep but much alcohol" on the night preceding the experiment of Dec. 5. DISCUSSION OF RESULTS. 79 Table 13. -Increase in heat output during walking in experiments with subject I — Continued. [Observations made by Dr. Carl Tigerstedt.] Date and condi- tions of experi- (a) Body- weight (6) (c) (d) Distance Horizontal Raising kilogram- of bodv Heat-output (computed). Increase over standing. ments. with per meters. per minute. (e) (.f) to) clothing. minute. (aX6). Total Per horizontal per Total. kilogrammeter. minute. (/-^c). 1913 Without food — Continued. Dec. 18: kilos. meters. meters. cals. cals. gm.-cal. Standing.... 1.29 Walking 74! 76 75.'3^ 5,629 3.52 3.98 2.&9 A78 76.4 5,712 3.68 4.09 2.80 .490 76.7 5,734 3.68 4.19 2.90 .506 76.9 5,749 3.73 4.22 2.93 .510 Dec. 19: Standing 1.36 Walking 74.17 76.1' 5,644 3.83 4.29 2.93 .519 77.9 5,778 3.99 4.23 2.87 .497 78.3 5,808 4.10 4.26 2.90 .499 78.7 5,837 4.20 4.26 2.90 .497 Dec. 20: Standing 1.31 Walking 74.31 76.2 5,662 3.68 4.00 2.69 A75 77.7 5,774 4.04 4.28 2.97 .514 78.6 5,841 4.20 4.43 3.12 .534 78.9 5,863 4.25 4.48 3.17 .541 Deo. 22: Standing 21.34 Walking 73! 77 76!3 . 65.9'^ 5^629 4!67 4.21 2!87 .510 4,861 3.17 3.76 2.42 .498 77.4 5,710 4.28 4.23 2.89 .506 67.51/ 4,979 3.41 3.74 2.40 .482 Deo. 23: Standing n.34 Walking 73! 76 75!9 5^698 3.97 4.03 2.69 .481 66.4 »/ 4,897 3.07 3.55 2.21 .451 76.9 5,671 4.08 4.14 2.80 .494 66.4'' 4,897 3.30 3.66 2.32 .474 Deo. 27: Standing n.34 Walking 72! 96 78 ^4 5,720 i.m 4.07 2.73 .'477 65.2^ 4,757 3.02 3.46 2.12 .446 78.6 5,735 4.13 4.07 2.73 .476 Average. . . With food. 65.1^ 4,750 3.02 3.48 2.14 .451 73.10 75.9 3.78 2.81 .507 Nov. 26: Standing .... 1.61 Walking 73] 30 75] 3 5^519 3.57 5.03 3.42 !626 Deo. 9: Standing 1.61 Walking 72 [33 76!6 6^497 3.68 4.61 3.60 !546 78.3 5,663 3.89 4.39 2.78 .491 79.1 5,721 4.04 4.33 2.72 .475 ■^Computed from the average elevation of the body per step as obtained from the kymograph records and the steps per minute. See table 3 for information regarding number of steps. 'Average of "standing" values obtained in experiments without food, Nov. 29-Dec. 20. 80 ENERGY TKANSFORMATIONS DURING HORIZONTAL WALKING. trained athlete, and although not particularly careful of himself, when out of training, he was nevertheless in reasonably good condition. The fact that alcohoUc excesses preceded the experiment on December 5 shows, of itself, that the subject was not in strict training. Indeed, it was impossible for us to control him outside of the Laboratory. In considering the values without food, it is of interest to note the variations in the standing metabolism in their relation to the total increase due to walking. While we find from the values given in table 6 that the average heat-output for this man without food was 1.34 calories per minute, it will be seen here that the variations ranged from 1.28 calories per minute on December 4 to 1.57 calories on December 5. The latter date was the day following the alcohoUc excesses and it may be noted that this increase in the basal metaboUsm continued through- out the walking periods, as the three highest values for the heat-output during walking were found on this day. On the other hand, when the high basal metabohsm is deducted, the gram-calories per horizontal kUogrammeter are essentially the same as the average value. EXPERIMENTS WITH FOOD. On the two days in which experiments were made with subject I after the ingestion of food, the standing or basal metaboUsm was per- ceptibly higher than in the experiments without food. The total metaboUsm during walking was likewise higher in all of the experi- mental periods. On the other hand, when the standing or basal metab- oUsm is deducted, we find that the gram-calories per horizontal kilo- grammeter were, in the second experiment, quite within the normal Umits. We can find no explanation for the high value obtained on the first day. The fact that the high basal metaboUsm due to food per- sisted through the walking period is of special interest, as it shows that the muscular activity incidental to the amount and rate of walking in these experiments resulted in a metaboUsm which was superimposed upon the increased metaboUsm due to the ingestion of food. In general, we may say that when this subject walked on a level plane, the energy required over and above maintenance to move 1 kilogram over a distance of 1 meter, i. e., 1 horizontal kUogrammeter, was in round numbers 0.5 gram-calorie. The correlation between these results and those obtained with subject II and the earUer observations of Benedict and Cathcart may properly be deferred until the discussion of the experiments with subject II. ENERGY REQUIRED FOR THE ELEVATION OF THE BODY. In walking at the average speed of 75.9 meters per minute, we find, from the records obtained with Dr. Tigerstedt's tracing pointer and by counting the number of steps, that the total distance per minute that the subject raised his body in a vertical direction amounted, on the average, to 3.78 meters per minute. With an average body-weight of DISCUSSION OF BESULTS. 81 73.1 kilograms, this would correspond to a work equivalent of 276.32 kilogrammeters or 0.65 large calorie per minute. Since the total increase over the standing metabolism due to walking averaged 2.81 large calories per minute, it is seen that approximately 23 per cent of the energy required for the work of forward progression was employed in raising the body in a vertical direction. These experiments give no data regarding the amount of energy required to lower the body after being raised at each step. Various assumptions are found for this in the Uterature, ranging from one- quarter to three-quarters of the work done in the elevation of the body, but it is obviously out of place to attempt such gross interpretations of the results. In conclusion, then, we may say that when walking in a horizontal direction at the rate of 76 meters per minute, a man with a body-weight of 73 kilograms produces 2.81 calories per minute above his standing basal metabohsm, of which 0.65 calorie or 23 per cent is required to raise the body through a distance of approximately 4 meters per minute. It is thus apparent at the outset that a very important factor in the energy consumption while walking in a horizontal direction may be the type of step employed. Unfortunately our observations were not made upon a sufficient number of persons to permit a complete discussion of this point. WALKING EXPERIMENTS WITH SUBJECT II. Subject II, who was a professional athlete, though not in continuous training, had exercised excessively in bicycle riding and had been the subject in Cathcart's research on the muscular work of bicycle riding. He had done a considerable amount of walking when in training, but was not an especially weU-trained walker and certainly could not be classed as a professional pedestrian. On the other hand, he performed all of the tests on the treadmill at even the highest speeds with perfect ease and was at no time unduly distressed. The walking experiments with this subject covered a period of approamately two months, i. e., from March 16 to May 15, 1914. About 40 per cent of the experiments were made without food; the remainder of the experiments followed either the breakfast or the mid- day meal, the diet in some cases being controlled. The speed of rota- tion of the treadmill varied during the experiments without food from 56.0 meters per minute to 146.8 meters per minute and in a few experi- ments, with the subject running, it increased to 148.7 meters per minute. The greater part of the experiments were made with a speed between 56.0 and 93.3 meters per minute. In computing the energy per horizontal kilogrammeter, we have the same problem to consider that was met with in the case of subject I, namely, the selection of a suitable base-Une. As the tests with sub- ject II were more varied than those with the first subject, at least 82 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. three base-lines are possible, if not indeed permissible. These are, (1) the average resting value with the subject standing in a relaxed posi- tion, without su'pport, which was obtained in 25 experimental days from March 20 to May 14, 1914, this value being 1.25 calories per minute; (2) the average resting, lying value of 51 experiments in 1911-12 by Benedict and Cathcart, this being 1.17 calories per minute, referred to here as lying value I; and (3) the resting lying basal value found as an average of four experimental periods on April 18, 1914, namely, 1.11 calories per minute, referred to here as lying value II. The subject in all these experiments was without food. EXPERIMENTS WITHOUT FOOD. All of the experiments without food for subject II have been brought together in table 14 and are arranged chronologically, the only dis- turbance in this order being for the experiment on May 13. The change in position for the experiment of May 13 was made to bring the results more nearly in the order of the increasing velocity of walking. The heat-output per horizontal kilogrammeter for each experiment has been computed in gram calories on the three base-Unes previously cited. The results using a basal value obtained with the subject standing in a relaxed position are given in column g; those with an average basal value obtained by Benedict and Cathcart in their experiments between December 7, 1911, and April 16, 1912 (lying value I), are given in col- umn h; and those with a basal value obtained from the special experi- ment made by Mr. L. E. Emmes on April 18, 1914 (lying value II), are given in column i. Inasmuch as the values for the subject standing relaxed were determined specifically for this research, and hence in this particular, at least, are more appropriate than the other basal values used, we shaU lay great stress in our discussion upon the incre- ments computed from this base-line. A general inspection of the values in column g shows that there is a tendency towards constancy until May 4, at which time the rate of walking first exceeded 100 meters per minute. Considering the experiments when the speed was below 100 meters per minute, we find 57 periods with speeds ranging from 56 to 93.3 meters inclusive, the average speed being 71.5 meters per minute and the energy require- ment for one horizontal kilogrammeter averaging 0.493 gram-calorie. An inspection of the table shows considerable variation above and below these average figures in isolated cases as, for instance, on April 15, when in one period the heat output per horizontal kilogrammeter was 0.603 gram-calorie. On the other hand the value of 0.555 gram- calorie is exceeded but six times and the individual values fall below 0,45 gram-calorie likewise only six times out of the 57 periods. It is thus clear that the value 0.493 gram-calorie is distinctly representative of the heat requirements for moving 1 kilogram 1 meter at speeds ranging from 56 to 93.3 meters per minute. DISCUSSION OF BESULTS. 83 Table 14. — Increase in heat-output during walking in experiments mth subject II ivithmUfood. to) (b) to) to) Heat-output toomputed). to) Total per minute. (/) Increase above standing relaxed. Increase per kilogram- Date and condi- tions of experi- ments. Body- weight with cloth- ing. Dis- tance per minute. Horizon- tal kilo- gram- meters toXft). Raising of body per minute. meter (gram-calorie.) to) Above standing relaxed to) Above lying. (i) Above lying, (/-i-c). I. II. 1914 Standing, re- kUos. meters. meters. cals. cals. gm.-cal. gm.-cal. gm.-cal. laxed» 1.25 Lying, P 1.17 Lying, IP 1.11 Walking: Mar. 20.. 72.4 (*) 2.61 3.48 2.23 24.. 72.0 60.6 4,363 1.09 3.74 2.49 0.571 o!689 o!663 25.. 72.6 60.6 4,363 1.77 3.40 2.16 .493 .611 .626 26.. 72.0 62.7 4,514 3.09 3.42 2.17 .481 .498 .612 27.. 72.0 57.6 4,147 2.09 3.16 1.90 .468 .477 .492 31.. 72.0 57.2 4,118 2.01 3.27 2.02 .491 .510 .626 Apr. 1.. 72.0 59.3 4,269 2.22 3.36 2.11 .494 .513 .527 3.. 72.0 76.2 5,486 2.38 4.39 3.14 .572 .587 .598 6.. 72.0 57.8 4,161 1.78 3.32 2.07 .498 .517 .631 56.8 4,090 1.83 3.26 2.00 .489 .609 .623 56.0 4,032 1.91 3.20 1.95 .484 .603 .618 7.. 72.0 59.6 4,291 1.98 3.63 2.28 .631 .650 .664 58.2 4,190 1.95 3.19 1.94 .463 .482 .496 8.. 72.0 76.1 5,479 3.01 4.45 3.20 .584 .699 .610 77.6 5,587 2.81 3.97 2.72 .487 .501 .612 78.3 6,637 2.85 3.89 2.64 .468 .483 .493 9.. 72.0 59.3 4,269 1.92 3.40 2.15 .604 .622 .536 58.8 4,233 2.05 3.20 1.95 .461 .480 .494 59.0 4,248 1.98 3.36 2.11 .497 .516 .530 10.. 72.0 77.3 5,566 2.77 4.09 2.84 .610 .525 .636 78.9 5,681 3.21 4.09 2.84 .500 .614 .525 14.. 71.9 61.7 4,436 2.45 3.43 2.18 .491 .609 .523 62.1 4,465 2.34 3.28 2.03 .456 .473 .486 62.4 4,487 2.37 3.26 2.01 .448 .466 .479 15.. 71.5 77.2 5,520 3.33 4.58 3.33 .603 .618 .629 79.4 5,677 3.52 4.02 2.77 .488 .502 .613 80.2 5,734 3.49 4.04 2.79 .487 .501 .611 16.. 71.5 61.9 4,426 2.38 3.16 1.91 .432 .460 .463 62.3 4,454 2.37 3.22 1.97 .442 .460 .474 61.8 4,419 2.20 3.21 1.96 .444 .462 .475 17.. 71.5 76.6 6,477 3.36 4.36 3.11 .568 .582 .593 78.1 5,584 3.45 3.94 2.69 .482 .496 .507 79.3 6,670 3.80 4.14 -2.89 .510 .624 .634 21.. 70.5 60.8 4,287 2.32 3.13 1.88 .439 .467 .471 22.. 70.5 76.8 5,415 3.33 3.91 2.66 .491 .506 .617 78.6 6,642 3.44 3.97 2.72 .491 .505 .616 79.2 6,684 3.42 3.94 2.69 .482 .496 .607 23.. — _ ^ — 70.6 61.1 {') 4,308 2.53 2.46 3.27 3.17 2.02 1.92 .469 .487 .501 3ee table 91, Bene- 'Average of periods "standing, relaxed" as shown in table 10, p. 71. 'Average of results with this subject from Dec. 7, 1911 to Apr. 16, 1912. diet and Cathcart, Carnegie Inst. Wash. Pub. 187, 1913, p. 78.) 'Determined during four periods of an experiment on Apr. 18, 1914, the subject being without food. ^Probably at the rate of about 67 meters per minute. 'Probably at the rate of about 61 meters per minute. 84 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. Table 14 — Increase in heat-output during walking in experiments with subject II without /ood—Continued. (a) (6) (c) (d) Heat-output (computed). (e) Total per minute (/) Increase above standing relaxed. Increase per kUogram- Date and condi- tions of experi- ments. Body- weight with cloth- ing. Dis- tance per minute Horizon- tal kilo- gram- meters (aXb). Raising of body per minute meter (gram-calorie). (ff) Above standing relaxed (A) Above lying, (i) Above lying, C/+C). I. II. 1914 Walking — Continued ; kilos. meters. meters. cats. cats. gm.-cal. gm.-cal. gm.-cal. Apr. 24.. 70.5 76.4 5,386 3.60 3.79 2.54 .472 .486 .498 78.4 5,527 3.69 3.94 2.69 .487 .501 .512 25.. 69.9 63.0 4,404 2.65 3.21 1.96 .445 .463 • .477 64.0 4,474 2.69 3.30 2.05 .458 .476 .489 27.. 70.5 73.5 5,182 3.25 3.77 2.52 .486 .502 .513 78.1 5,506 3.64 3.97 2.72 .494 .509 .519 . 28.. 70.5 78.2 5,513 3.41 4.04 2.79 .506 .521 .531 79.8 5,626 3.60 3.99 2.74 .487 .501 .512 80.0 5,640 3.53 4.06 2.81 .498 .512 .523 80.7 5,689 3.80 4.12 2.87 .504 .519 .529 80.4 5,668 3.33 4.40 3.15 .556 .570 .580 80.2 5,654 3.67 4.07 2.82 .499 .513 .524 80.2 5,654 3.80 4.14 2.89 .511 .525 .536 80.4 5,668 3.46 4.09 2.84 .501 .515 .526 May 13.. 71.5 82.6 5,906 3.76 4.13 2.88 .488 .501 .511 89.7 6,414 4.51 4.50 3.25 .507 .519 .529 80.1 5,727 3.29 3.94 2.69 .470 .484 .494 76.6 5,477 2.84 3.75 2.50 .456 .471 .482 93.3 6,671 5.25 4.72 3.47 .520 .532 .541 91.9 6,571 5.15 4.67 3.42 .520 .533 .542 4.. 71.7 114.5 8,210 6.43 6.71 5.46 .665 .675 .682 109.1 7,823 6.16 5.79 4.54 .580 .591 .598 102.0 7,313 5.88 5.42 4.17 .570 .581 .589 5.. 71.7 106.0 7,601 4,86 5.54 4.29 .564 .575 .583 102.6 7,356 5.82 5.33 4.08 .555 .566 .574 103.6 7,428 6.06 5.51 4.26 .574 .584 .592 6.. 71.7 140.7 10,088 9.76 10.23 8.98 .890 .898 .904 139.6 10,009 8.90 10.38 9.13 .912 .920 .926 142.9 10,246 8.79 10.91 9.66 .943 .951 .956 10.. 72.3 145.5 10,519 5.37 10.99 9.74 .926 .934 .939 146.8 10,613 5.81 11.68 10.43 .983 .990 .996 146.6 10,598 8.02 11.00 9.75 .920 .928 .933 Running: May 11.. 72.0 146.6 10,554 13.45 9.47 8.22 .779 .786 .792 146.7 10,562 13.89 9.57 8.32 .788 .795 .801 148.3 10,677 14.80 9.69 8.44 .790 .798 .804 12.. 72.0 147.1 10,591 15.40 10.80 9.55 .902 .909 .915 148.4 10,684 14.09 9.19 7.94 .743 .751 .756 148.1 10,663 15.40 9.80 8.55 .802 .809 .815 148.1 10,663 14.74 9.97 8.72 .818 .825 .831 148.7 10,706 15.37 10.06 8.81 .823 .830 .836 Walking: May 14.. 71.5 146.4 10,467 7.59 11.23 9.98 .953 .961 .967 Running: May 14.. 146.5 10,474 15.44 11.13 9.88 .943 .951 .957 147.2 10,524 12.47 9.15 7.90 .751 .758 .764 147.6 10,553 12.34 9.26 8.01 .759 .767 .772 148.1 10,588 11.09 9.66 8.41 .794 .802 .808 15.. 71.0 144.7 10,273 12.48 9.64 8.39 .817 .824 .830 148.4 10,536 12.14 9.67 8.42 .799 .807 .812 147.7 10,486 13.13 9.47 8.22 .784 .792 .797 DISCUSSION OF RESULTS. 85 In two of the walking experiments without food, those of April 6 and 7, we were able to obtain electrocardiograms of the pulse of this subject and the results are recordedintable4 (pp. 54 and 55). In previous researches a striking uniformity has been found in the changes of the metaboUsm and the pulse-rate, amounting at times to a distinct percentage relationship between them. Such a uniformity was noted by Benedict and Cathcart in their study of a bicycle rider when the metaboUsm and pulse-rate during riding were compared with the values obtained in the lying or sitting position. In this research, on the contrary, the pulse-rate in the experiments of April 6 and 7 showed a distinct lowering when the subject changed from the standing posi- tion to walking, particularly in the experiment on April 7, notwith- standing the fact that the total metaboUsm during walking increased from 100 to 200 per cent above the basal metaboUsm of standing. This lowering of the pulse-rate was so positive that it is difl&cult to beUeve that any error was made in the measurements. Furthermore, in the fourth period of the experiment on April 7, we find that the pulse- rate with the subject standing after walking increased materially over that during walking. The evidence thus impUes that when standing still upon the treadmill, this subject had a much higher pulse-rate than when he was walking at a slow speed. These records are entirely contrary to our previous experience but the lowered pulse-rate during walking has been confirmed by Professor H. Monmouth Smith, of the Laboratory staff, in similar experiments with three other subjects, thus estabUshing the fact. Further experimenting is now in progress and the results will be incorporated in a subsequent report. Influence of Velocity. On all days subsequent to May 4, with the single exception of May 13, the speed of walking exceeded 100 meters per minute. At this higher speed there was a distinctly higher energy requirement per horizontal kilogrammeter. Thus, we have six periods when the rate of walking was from 102 to 114.5 meters per minute and averaged 106.3 meters per minute. At this speed the average heat-output per hori- zontal kilogrammeter was 0.585 gram-calorie, a material increase over 0.493 gram-calorie, the average heat-output for the lower speed. A further grouping of the experiments is permissible when the speed ranged from 139.6 to 148.7 meters per minute, as with this higher speed a considerable increase is shown in the heat requirement per unit of work. Some of the experiments in this group show that when the subject ran instead of walked, the heat-output per horizontal kilo- grammeter was lowered, although the rate of progression was prac- tically the same; accordingly we may not advantageously draw average values. In the two walking experiments on May 6 and 10 the speed averaged 143.7 meters per minute and the heat per horizontal kilo- grammeter 0.929 gram-calorie. In the first period of the experiment 86 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. of May 14 the subject walked at the rate of 146.4 meters per minute, with a heat-output of 0.953 gram-calorie per horizontal kilogrammeter. In the four following periods the subject ran, although the speed per minute was approximately the same as that in the walking period. Nevertheless, the average heat-output per unit of work was materially less, i. e., 0.812 gram-calorie. It is thus apparent that the velocity had a very considerable influ- ence upon the heat per unit of work when the subject was walking. This confirms the earlier contention of Durig and his associates that the heat required to move 1 kilogram 1 meter increases with the speed. Furthermore, the method of progression, i. e., walking or running, had likewise a very considerable influence on the heat per unit of work. Both of these factors require subsequent elaboration. Comparison of the Heat peh Unit of Work as Computed fbom Difpehent Base-linhb. The last three columns of table 14 permit the comparison of the heat per unit of work, i. e., per horizontal kilogranuneter, as computed from the three base-hnes. Instead of using the individual values in table 14, however, this comparison can better be made by using only the average figures for the several groups presented in the foregoing discussion on the influence of velocity. To find exactly the differ- ences due to the selection of the base-line, we have therefore brought together in table 15 a general average for all of the experiments without food, both walking and running. The walking experiments are divided into three groups, according to the speed at which the subject walked. Thus we have 57 periods when the subject walked at a slow rate ranging from 56 to 93.3 meters and averaging 71.5 meters, 6 periods with a medium speed averaging 106.3 meters per minute, and 7 periods with a high speed averaging 144.1 meters per minute. During the running experiments, the speed averaged 147.5 meters per minute. The values showing the heat-output per unit of work done, using the Benedict and Cathcart basal value (lying I), are given in column 6, those for which the basal value of April 18, 1914, was employed (lying II) in column c, and those calculated from the basal value found with the subject standing in the relaxed position are given in column d. With the large amounts of heat involved in the walking experiments the relatively sUght differences appearing in these three base-Unes should not have a very great effect. It is obvious, moreover, that the greater the amount of work done in walking, the less will be the per- centage change due to the selection of the base-line. Thus we note that the heat per unit of work done at the slow speed is lower when the standing basal value is employed than it is when the lying value II (that of April 18, 1914) is used, the difference between the two averages being approximately 6 per cent. With the increase in the work involved in walking at the moderate speed, the difference due to the selection of the basal value becomes considerably less, i. e., 3 per cent, DISCUSSION OF RESULTS. 87 and finally with the large heat production incidental to walking at the rate of 144.1 meters per minute, the difference between the calculations with the two base-Unes becomes approximately only 1 .5 per cent. This small difference likewise holds in the running experiments. The values computed from the Benedict and Cathcart base-line show the same general characteristics, although the difference between the two results is smaller than with the basal value of April 18, 1914. It thus appears that as the total amount of work increases it becomes less important which of the three base-lines is used. For slow and moderate speeds a possible difference of 3 to 6 per cent in the average value of the heat per unit of work may be expected, while with the highest speed this difference decreases to from 1 to 1.5 per cent. The Table 16. — Comparison of heat-output during walking and running as referred to basal values for the lying and standing positions. [Experiments with subject II without food.] Number of periods. Average distance per minute (meters) (a) Heat (computed) per hori- zontal kilogrammeter (gram-calorie). Above lying I' Above lying 11' (0 Above standing relaxed' (d) Walking: 57 6 7 Running : 15 71.5 106.3 144.1 147.5 0.509 .595 .940 .814 0.521 .603 .946 .819 0.493 .585 .932 .806 'See table 14, p. 83, for explanation of basal values used. importance of the selection of a basal value is therefore greatest when the amount of walking to be performed is smallest, i. e., when the sub- ject is walking at moderate speed. This is somewhat unfortunate, for a large proportion of walking for exercise, for the carrying of burdens, and in marching is usually done at a moderate speed. Since, however, the basal values obtained with the subject standing in the relaxed position were daily and accurately determined, and since there is such a high degree of constancy in the standing relaxed values, we consider that they may be properly used as the basal values for these compu- tations. The figures obtained per unit of work on this basis may there- fore be considered to represent the increased metaboUsm necessary to move 1 kilogram 1 meter in a horizontal direction. EXPERIMENTS WITH FOOD. A large number of experiments following the ingestion of food were made with subject II. The results of all these experiments are pre- sented in table 16, the data being grouped arbitrarily according to the preceding meal and especially according to the character of the diet. 88 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. Table 16. — Increase in heat-oiUpvt during walking in experiments with, subject II with food. (a) (6) (c) (d) Heat-output (computed). Increase over Body weigh with Horizon- tal kilo- gram- standing. Diet and date. Condition. t ^^^ tance Raising of body (e) (/) (0) Per cloth- per meters per Total horizon- ing. minute. (0X6). minute. per minute. Total. tal kilo- gram- meter 1914 Breakfast. kUos meters. meters. cats. caXs. gm.-cal. Not controlled: Standing'. . '1.43 Mar. 16 . . Walking... iiA I Vi.Z 5^379 2^88 3.92 2.49 0.463 18.. ....do 72. < I 76.0 78.0 5,502 5,647 3.34 3.62 4.06 4.09 2.63 2.66 .478 .471 19.. ....do 72.-: 72.^ 72.^ t 74.9 y 57.9 1 57.9 57.5 5,423 4,192 4,192 4,163 3.16 3.11 2.74 S.61 4.21 3.74 3.40 3.74 2.78 2.31 1.97 2.31 .513 .551 .470 .555 21. . do 23.. ....do 28.. ....do 72. ( ) 59.4 4,277 2.52 t 3.48 2.05 .479 30. . ....do 72. ( 72. ( ) 58.5 ) 55.1 4,212 3,967 2.26 1.91 3.65 3.23 2.22 1.80 .527 .454- Apr. 4.. ....do 56.8 4,090 1.98 3.07 1.64 .401 56.2 4,046 1.87 3.24 1.81 .447 Controlled : Carbohydrate Apr. 29. . Standing^. . n.59 Walking... 76.5 ) 76.9 5^452 sies 4.31 2!72 .499 78.2 5,544 3.56 4.26 2.67 .482 79.5 5,637 3.52 4.20 2.61 .463 80.9 5,736 3.74 4.07 2.48 .432 81.1 5,750 3.76 4.09 2.50 .435 80.9 5,736 3.29 3.87 2.28 .397 81.0 5,743 3.73 4.10 2.51 .437 81.1 5,750 3.69 3.98 2.39 .416 Lunch. Not controlled: Standing'. . n.55 Mar. 16. . Walkine... 72^ t 75!3 5,452 3.17 3.99 2.44 !448 18.. ....do 72.' t 76.3 5,524 3.75 4.02 2.47 .447 19.. ....do 72.' t 74.9 5,423 3.55 4.09 2.54 .468 20.. ....do 72.' t 52.7 3,815 2.39 3.16 1.61 .422 21.. ....do 72.' 72. ( 72. ( 72. ( t 52.8 ) 62.7 ) 61.1 ) 60.7 3,823 4,514 4,399 4,370 4,219 2.59 .81 1.50 2.30 3.40 3.99 3.64 3.67 1.85 2.44 2.09 2.12 .484 .541 .475 .485 24. . do 25.. do 26.. do 27.. ....do 72. ( ) 58.6 2.15 3.62 2.07 .491 28.. ....do 72. ( ) 59.3 4,270 2.12 3.81 2.26 .529 30.. ....do 72. ( ) 58.5 4,212 2.40 3.59 2.04 .484 57.3 4,126 2.24 3.53 1.98 .480 31.. ....do 72!( ) 57.7 4,154 1.72 3.53 1.98 .477 Apr. 3 . . ....do 72. ( ) 57.0 4,104 2.02 3.44 1.89 .461 77.0 5,544 2.84 4.68 3.13 .565 78.9 5,681 3.38 4.47 2.92 .514 4.. ....do 72!( ) 54.4 3,917 4,183 2.05 3.51 1.96 .500 58.1 2.21 3.74 2.19 !524 'Average of all periods with subject standing after a light meal (breakfast). See table 11, p. 72. 'Standing, relaxed. Average of values obtained with similar diet on Apr. 25 and Apr. 27. See table 12, p. 74. ^Average of results obtained for all positions standing after lunch, diet not controlled. See table 12, p. 74. DISCUSSION OF RESULTS. 89 Table 16. — Increase in heat^ouljmt during walking in experiments with subject II with food — Continued . (a) (b) (c) (d) Heat-output (computed). Increase over Body- weight with Horizon- tal kilo- gram- standing. Diet and date. Condition. Dis- tance Raising of body (e) (/) (ff) Per doth- ing. per minute. meters (aXh). per minute. Total per minute. Total. horizon- tal kilo- gram- meter (/-^c). 1914 Lunch — Cont'd. Gontrolled: Protein: kilos. meters. meters. cats. cats. gm.-cal. Apr. 6., Standing. . 1.43 Walking... 72.6' 57.5 4,140 2^21 3.54 2;ii isib 57.2 4,118 1.94 3.51 2.08 .505 56.3 4,054 1.81 3.46 2.03 .501 7.. Standing. . 1.37 Walking... 72^0 58.8 4^234 i!78 3.59 2^22 .524 58.7 4,226 1.83 3.77 2.40 .568 56.9 4,097 1.82 3.41 2.04 .498 8.. Standing . . 1.33 Walking... 72.0 76^9 5^536 2.50 4.28 2.95 .533 78.0 5,616 3.01 4.38 3.05 .543 79.2 5,702 2.70 4.46 3.13 .549 23.. Standing. . 1.24 Walking... 70!5 6i.2 4^315 2.74 3.55 2.31 .535 61.6 4,343 2.57 3.44 2.20 .507 61.6 4,343 2.48 3.45 2.21 .509 61.7 4,350 2.63 3.45 2.21 .508 24.. Standing . . 1.41 Walking... 70.5 77!o 5,428 3.74 4.05 2.64 .486 78.8 5,555 3.88 4.11 2.70 .486 80.0 5,640 3.84 4.26 2,85 .505 80.0 5,640 3.88 4.24 2.83 .502 May 5 . . Standing^. . 11,36 Walking... 71 !7 112.6 8^073 7'69 6.34 4.98 .617 113.7 8,152 7.83 6.43 5.07 .622 106.0 7,600 6.40 5.89 4.53 .596 111.3 7,980 7.76 6.31 4.95 .620 May 10.. Standing'. . n.36 Walking... 72!3 146.5 10,592 7.83 11.05 9.69 .915 146.9 10,621 8.28 11.07 9.71 .914 Carbohydrate: 1.43 Apr. 9 . . Standing . . Walking . . . 72.0 59.4 4i277 i!95 3.48 2.05 .479 59.0 4,248 1.94 3.43 2.00 .471 58.8 4,234 1.94 3.53 2.10 .496 Apr. 10.. Standing. . 1.47 Walking.. 72!o 78.0 5,616 3.01 4.70 3.23 .575 79.1 5,695 3.17 4.59 3.12 ,548 79.7 5,738 3.35 4.37 2.90 .505 14. Standing . 1.42 Walking.. 7i!9 6i!2 4,400 2!38 3.40 i.98 .450 62.0 4,458 2.20 3.54 2.12 .476 62.2 4,472 2.27 3.47 2.05 .458 15. Standing . 1.38 Walking . . . 71.6 78!2 5^591 3!88 4.52 3.14 .562 80.1 5,727 3.72 4.41 3.03 .529 80.6 5,763 3.88 4.63 3.25 .564 I ^ . iStandingi relaxed; average obtained on 6 days with protein diet (table 12). 90 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. Table 16. — Increase in heat-output during walking in experiments mth subject II vnth food — Continued. (a) (fc) (c) (d) Heat-output (computed). Increase over standing. Diet and date. Condition. Body- weight with Dis- tance Horizon- tal kilo- gram- Raising of body (e) Total (/) (g) Per cloth- ing. per niinute meters (oXfc). per minute per minute Total. horizon- tal kUo- gram- meter (f^c). 1914 Lunch — Cont'd. Carbohydrate — cont'd. kUoa. meters. meters. cats. coils. gm.-cal. Apr, 25.. Standing. . 1.58 Walkine.. 69.9 63.1 4^411 2.88 3.52 1.94 .'446 63.8 4,460 2.70 3.53 1.95 .437 63.7 4,453 2.85 3.49 1.91 .429 63.3 4,425 2.64 3.34 1.76 .398 27.. Standing. . 1.61 Walking... Tois 77.5 5,464 3.60 4.30 2.69 .492 ' .... 78.9 5,562 3.56 4.02 2.41 .433 .... 79.8 5,626 3.88 4.24 2.63 .467 80.2 5,654 3.74 4.04 2.43 .430 May 4 . . Standing^ . '1.59 Walking... 7i;7 lis 14 sjisi 6!84 6.59 5.66 .615 112.3 8,052 7.88 6.46 4.87 .605 110.3 7,909 7.41 6.10 4.51 .570 6.. Standing^ . '1.59 Walking... 7i!7 143.6 10,253 7!82 10.73 9!i4 !89i 11.. Standing!. '1.59 Running. . . 72.0 148.7 10,706 13^95 9.50 7.91 !739 Fat; Apr. 16.. Standing . . 1.41 Walking... 71.5 61.6 4^405 2.49 3.35 1.94 .440 62.3 4,455 2.48 3.15 1.74 .391 62.2 4,447 2.24 3.43 2.02 .454 61.9 4,426 2.21 3.42 2.01 .454 17.. Standing. . 1.36 Walking... n.'s 77.6 5^548 3.96 4.06 2.76 .487 79.5 5,684 3.78 4.27 2.91 .512 80.0 5,720 3.85 4.56 3.20 .559 80.0 5,720 3.67 4.15 2.79 .488 21.. Standing. . 1.42 Walking... 70.5 60 !4 4; 258 2.45 3.33 1.91 .449 60.6 4,272 2.56 3.43 2.01 .471 60.5 4,265 2.34 3.44 2.02 .474 60.5 4,265 2.23 3.43 2.01 .471 22.. Standing. . 1.42 Walking... 70.5 77.3 5, '456 3.86 4.07 2.65 .486 78.7 5,548 3.86 4.24 2.82 .508 79.5 5,604 3.92 4.25 2.83 .505 79.7 5,619 3.84 4.18 2.76 .491 'Standing, relaxed. Average of values obtained with .sim lar diet on Apr. 25 and Apr. 27. See table 12, p. 74. DISCUSSION OF RESULTS. 91 As we have already seen from the discussion in an earlier section/ the metabolism in the standing relaxed position after the ingestion of food was considerably greater than that obtained when the subject was without food, this being due to the katabolic stimuh of the foodstuffs. Of most importance, however, is the question whether the increment due to walking is superimposed upon the increased katabohsm due to food or whether this increment is lessened by the fact that the body is previously stimulated to a greater katabolic activity. With a view to studying the influence of variations in the intensity of the pre-walking stimulation, certain of the diets were so controlled as to consist in large part of one of the three principal food constituents, i. e., protein, fat, or carbohydrate. With the experimental conditions and the impos- sibility of controlling the subject's diet while he was outside of the laboratory, it was impracticable to make a thorough study of this par- ticular phase of the problem. Our data do not therefore present a conclusive statement as to the influence of the special protein, fat, or carbohydrate diets, but merely contribute to the interesting question as to whether or not the increase in the katabohsm following the inges- tion of food persists during increased muscular activity or if there be a summation effect under these conditions. Since we have seen not only from the work of previous investigators, but from tables 14 and 15, that there is a marked influence upon the heat per unit of work as the result of an increase in velocity, particu- larly when the velocity is above 95 meters per minute, it seems prefer- able to consider the data presented in table 16 from this point of view. As in the experiments with this subject when no food was taken, here again we find that in the greater number of the experiments the speed varied from 52.7 to 81.1 meters per minute, there being but 7 periods at speeds ranging from 106 to 113.7 meters per minute, and but 4 periods with a speed over 140 meters per minute. Considering first the experiments at the lower speed, we find that 91 periods are available for study. The average velocity diu-ing these periods was 68.2 meters per minute and the heat per unit of work done, i. e., per horizontal kilogrammeter, was 0.486 gram-calorie. It will be seen from the table that the standing basal value for the total heat- output per minute in the experiments with food ranged from 1.24 to 1.61 calories per minute, averaging considerably above those obtained when the subject was standing in the post-absorptive condition. On the other hand, since the average value per unit of work done in 57 periods without food was 0.493 gram-calorie and the average of 91 periods with food was 0.486 gram-calorie, it is clear that the increase over the basal metabolism was the same in both instances; in other words, the increment due to the work of forward progression was 'See p. 75. 92 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. constant, irrespective of whether the subject was with or without food, this being true for subject II with the rate of speed varying from 52.7 to 93.3 meters per minute. At moderate speeds above 100, we have seven periods with food in which the velocity varied from 106 to 113.7 meters per minute, the average speed being 111.4 meters per minute, and the average output of heat per unit of work done 0.606 gram-calorie. Certain difficulties appear when an attempt is made to compare the results of these periods with food with periods without food. First, the speed per minute without food was 106.3 meters and that with food 111.4 meters, this being a perceptible increase. Second, but few periods are available for comparison either with or without food, there being but 7 for the former and 6 for the latter. If we make such a comparison, however, we find that with food the heat per unit of work is 0.606 gram-calorie, while that without food was only 0.585 gram-calorie. It is probably not possible to attribute this increase in the heat-output per unit of work solely to the increase of 5 per cent in the speed, and we can only state that with speeds averaging 111.4 meters per minute the increase in the heat per horizontal kilogrammeter with food is perceptibly greater than that with food at a speed of 68.2 meters per minute. The comparison becomes even more difficult when we consider the experiments with the highest speeds, namely, those of 140 meters per minute and over. Here we have but 4 periods with food, with the speed ranging from 143 to 148.7 meters per minute and an average speed of 146.3 meters per minute. The comparison is still further comphcated by the fact that in one of these four periods the subject was running. The average heat-output per unit of work done for the three walking periods was 0.907 gram-calorie and the value found for the one period with the subject running was 0.739 gram-calorie, a value much less than that obtained with the subject walking. We may thus say that all of the observations made after the taking of food are completely in accord with those made when no food was taken, i. e., an increase in the energy per unit of work done as the speed increased and a considerably less energy per unit of work done when the subject was running as compared with that when he was walking. An analysis of the processes of running and walking and the character of the steps is necessary to interpret these differences intelhgently.^ Electrocardiograms of the pulse-rate were obtained in a few of the experiments following the ingestion of food, namely, those of April 4, 6, and 7.^ These records show the same discrepancy between the pulse- rate and the metabolism as was found in the experiments without food.* 'See p. 98. *See table 4, pp. 54 and 55. 'See p. 85. DISCUSSION OF RESULTS. 93 INFLUENCE OF THE CHARACTER OF DIET ON THE HEAT-OUTPUT PER UNIT OF WORK. While from the general observations of the metabolism with and without food the inference was properly drawn that there was no material increase in the heat-output per unit of work in the food periods, nevertheless certain of the experiments permit a more detailed examination of this point since a reasonably satisfactory grouping of the results may be made upon the basis of the character of the diet. The experimental plan as originally outlined called for a series of walking experiments at each of the three standard speeds with diets containing a preponderance of each of the three principal nutrients. The average figures representing fhe heat per unit of work with the three diets and the three speeds are collected in table 17. In this table the average speed for the different groups is given, as the speed per minute, especially with the low speed, was not exactly the same for the three diets. Only the walking experiments are considered in this connection, none of the running experiments being included. Table 17 — Influen:: jj charader of diet on the fieat-output per horizontal kilogranvmeter during walking experiments with subject II. Character of diet. Low speed. Moderate speed. High speed. Meters. Gm.-eal. Meters. Gm.-oal. Meters. Gm.-cal. Carbohydrate Protein 73.2 67.1 70.1 0.471 .516 .478 112.0 110.9 0.597 .614 143.0 146.7 0.891 .915 Fat . .■ Aside from the generally increasing value of the heat per unit of work as the speed increases, which is clearly shown in table 17, we note, in comparing the values obtained with the different diets, a dis- tinctly higher, although not striking, increase in the heat per unit of work with the protein diet as compared with the carbohydrate diet, this increase being approximately 9 per cent at the low speed and 3 per cent at the moderate and high speeds. With the fat diet the com- parison was made only with the low speed, and we find that the heat- output per unit of work was essentially the same as that with the car- bohydrate diet and measurably less than was obtained with the protein diet. The general impression derived from the comparison of all the experiments with subjects I stnd II was that the heat per unit of work was practically independent of the taking of food. It appears from the foregoing discussion, however, that this conclusion should be slightly modified when diets containing large quantities of protein are con- sidered, for the data in table 17 indicate that with such a diet a slightly higher heat-output per unit of work may be expected. 94 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. INFLUENCE OF FATIGUE UPON THE HEAT-OUTPUT PEE UNIT OF WORK. Many of our experiments were made with a sufficient number of con- secutive walking periods to permit a study of the heat per unit of work done under conditions of accumulated work. The average length of the walking period was 14 minutes. The distance traveled in a period in the prolonged walking experiments varied from 800 to 1,200 meters, and the subject walked not only diu-ing the experimental periods, when the gaseous metaboUsm was being measured, but also between these periods. When, therefore, the experiment consisted of four or more experimental periods, it will be seen that the continuous walking would amount frequently to several kilometers, thus providing conditions for studying the effect of fatigue. In relatively few of these experiments, however, was there substantial evidence of extreme fatigue. The results of the observations in which there were four or more periods of continuous walking are brought together in table 18. The data given show the total distance walked by the subject prior to the beginning of each experimental period^ and both the distance walked per minute and the heat-output per unit of work performed during the period. On two days, April 28 and 29, 1914, there were eight walking periods; the total distance traversed each day in continuous walking was approximately 23 kilometers. Throughout both experiments the distance per minute remained essentially constant at about 80 meters; we thus have ideal conditions for studying the influence of fatigue resulting from continuous walking. On April 28, when the subject was without food, there was no essential difference in the unit of work done as the experiment progressed, although there were variations from period to period. On the next day, a similar experiment was made after food had been taken. The average heat per unit of work done on this day was considerably less than that on April 28 and a general tendency was also shown for the heat per unit of work done to decrease as the work progressed. Inspection of the values found for another experiment of 6 periods on May 13, when the subject was without food, shows more or less fluctuation in the values obtained for the heat per unit of work done, but there is no uniformity in the changes and, indeed, the values are somewhat greater in the last periods than in the earlier periods. With the experiments having only four periods, in which the total distance walked prior to the last period ranged from 8 to 12 kilometers, we find that on certain days there was a tendency for the heat per unit of work to decrease as the walking progressed and on other days to increase, but in the majority of the experiments there was no regularity in the variations. From the data given in table 18, we may conclude that there may be noticeable differences in the absolute heat per unit of work done on different days, a point particularly brought out in the experiments of 'For total distance walked, aee table 4, pp. 56 to 60. DISCUSSION OF RESULTS. 95 April 28 and April 29. Nevertheless, there is no definitely uniform influence of prolonged walking upon the efficiency for horizontal movement shown by this subject and the heat per unit of work after he had walked 20 kilometers is as liable to be lower than the initial value as it is to be higher. In other words, in this series of experiments, the measured metabolism agrees perfectly with the subjective impression that there was not a sufficient degree of fatigue to affect the comfort Table 18. — Heat-outpiU per unit of work during prolonged walking in experiments with subject II . Distance walked. Heat Distance walked. Heat (com- puted) (com- puted) Date. Total to Per per hori- zontal Datel Total to Per per hori- zontal beginning minute during period. kilo- beginning minute during period. kilo- of period.' gram- meter.' of period.' gram- meter.* 1914 1914 Without food: meters. meters. gm.-cal. With food: meters. meters. gm.-cal. Apr. 28. . . 382 78.2 0.506 Apr. 29.. 650 76.9 0.499 3,322 79.8 .487 3,519 78.2 .482 6,316 80.0 .498 6,585 79.5 .463 9,538 80.7 .504 9,699 80.9 .432 13,062 80.4 .556 12,615 81.1 .435 16,416 80.2 .499 15,547 80.9 .397 19,033 80.2 .511 18,662 81.0 .437 22,479 80.4 .501 21,676 81.1 .416 May 13... 521 82.6 .488 Apr. 22.. 368 77.3 .486 3,993 89.7 .507 3,452 78.7 .508 7,737 80.1 .470 7,067 79.5 .505 11,391 76.6 .456 10,345 79.7 .491 16,413 93.3 .520 Apr. 23.. 348 61.2 .535 20,527 91.9 .520 2,723 61.6 .507 With food: 5,512 61.6 .509 Apr. 16... 303 61.6 .440 7,976 61.7 .508 2,557 62.3 .391 Apr. 24.. 354 77.0 .486 5,901 62.2 .454 2,879 78.8 .486 9,424 61.9 .454 5,541 80.0 .505 Apr. 17..-. 507 77.6 .487 8,295 80.0 .502 3,945 79.5 .512 Apr. 25.. 323 63.1 .440 7,572 80.0 .559 2,330 63.8 .437 12,266 80.0 .488 5,122 63.7 .429 Apr. 21... 307 60.4 .449 8,125 63.3 .398 2,658 60.6 .471 Apr. 27. . 372 77.5 .492 5,374 60.5 .474 2,905 78.9 .433 8,615 60.5 .471 5,693 9,366 79.8 80.2 .467 .430 'For total distance walked, see table 4, pp. 56 to 60. 'Calculated for each period from the increment above values obtained with the subject standing relaxed. See tables 14 and 16, pp. 83 and 88. of the subject or his metaboUsm and that at the speed used throughout these tests, namely, 80 meters per minute, walking may be done for many kilometers without affecting the efficiency of the body for loco- motion in a forward direction. The stamina of the subject and his capacity for excessive work was severely tested in the earlier research on prolonged bicycle riding, in 96 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. which it was demonstrated that this subject had ridden the equivalent of 100 miles over average roads on an empty stomach; but in this research it was impracticable to carry out prolonged endurance tests at high rates of speed. Indeed, at the highest speed, i. e., 140 kilometers per minute, it is very certain that the subject would have run instead of walked, as distances as great as 20 or more kilometers are rarely covered rapidly by walking, but usually by running. On the other hand, a speed of 80 meters per minute represents an average rapidity of motion when long-distance walking is to be done and hence the values obtained at this speed have a much greater practical use than would values secured with the subject walking at a higher speed. COMPARISON OF THE HEAT-OUTPUT PER UNIT OF WORK DURING RUNNING WITH THAT OBTAINED DURING WALKING. The striking difference between the heat per unit of work observed when the subject was running and that obtained when he was walking has already been noted in table 15 (see p. 87), the figures showing that considerably less energy per unit of work was required for moving the body forward in running than in walking under the experimental conditions. The values obtained when the subject was moving forward at the rate of approximately 144 meters per minute give the only data for a direct comparison, no running experiments being made at the lower rates of walking. At this speed, the value computed for walking (using the standing basal value) is 0.932 gram-calorie and for running 0.806 gram-calorie. That these are not accidental figures is shown by the fact that the first value is the average of 7 reasonably agreeing periods and the second value is the average of 15 well agreeing periods; in other words, we deal here with a positive difference of approximately 15 per cent. It thus becomes important to analyze, if possible, the mechanical movements incidental to walking, and par- ticularly rapid walking, to find the reason for this discrepancy. TraveUng at the speed of 144 meters per minute, the ordinary individual without load generally runs rather than walks. On the other hand, in transporting heavy material as, for example, nuhtary accouterments and trappings, running would be wholly impracticable and walking must be resorted to. It has been noted, especially in observations of professional walkers, that in rapid walking there is commonly very great extraneous arm movement, the arms being violently swung back and forth with each step. In the belief that this unusual extraneous muscular movement results in a large energy transformation which does not directly contribute to the forward pro- gression, the study previously referred to was made in which the energy requirement of subject II was observed while he stood and swung his arms violently as in rapid walking. These values have already been discussed in connection with the standing resting values,^ when it was 'See table 10 and p. 71. DISCUSSION OF RESULTS. 97 found that in one experiment the heat-output per minute was 2.53 calories and in the other 3.13 calories per minute, or an average heat- output of 2.83 calories per minute as compared with 1.25 calories per minute, the average of the basal values obtained with the subject standing relaxed. It will therefore be seen that when the arms were swung violently, the metaboUsm was increased 126 per cent. It is accordingly important for us to eUminate, if possible, the influ- ence of this extraneous arm motion. To this end we have computed in table 19 the heat-output per unit of work during rapid walking, using Table 19. — Increase in heat-output during fast walking in experiments with subject II. [Basal value, standing swinging arms.'] Date. Body- weight with clothing. Distance per minute. Heat-output (computed). Total per minute. Increase due to walking. Total. Per horizontal kilogrammeter. 1914 Without food: May 6 10.... 14.... Average With food: Protein — May 10 Carbohydrate — May 6 Average kilos. 71.7 72.3 7i]5 meters. 140.7 139.6 142.9 145.5 146.8 146.6 146.4 cais. 10.23 10.38 10.91 10.99 11.68 11.00 11.23 cals. 7.40 7.55 8.08 8.16 8.85 8.17 8.40 gm.-cal. 0.734 .754 .789 .776 .834 .771 .803 .780 72,3 71.7 146.5 146.9 143.0 11.05 11.07 10.73 8.11 8.13 7.56 .766 .765 .737 .756 'The subject stood without food swinging his arms violently as in the most rapid walking. (See table 10, p. 71.) In computing the results for walking without food, the average value (2.83 cals.) is used; for walking with food the increment (2.83 — 1.25 = 1.58 cals.) due to swinging the arms is added to the value obtained for "standing, relaxed" with the respective diets. 2.83 calories as the base-line, i. e., the average value found with the subject standing and swinging his arms violently. Under these condi- tions we find a marked depression in the heat-output per unit of work as compared with that computed with a base-Une obtained with the sub- ject standing in a relaxed position. In the experiments without food, this averaged with the high speeds, 0.932 gram-calorie,^ but with the new basal value, it averages 0.780 gram-calorie. This value compares favorably with the value of 0.806 gram-calorie^ found in the 15 periods with the subject running, being some 3 per cent less. >See table 15, p. 87. DISCUSSION OF RESULTS. 99 At the low speed the subject walked 69.3 meters per minute, and took 108.2 steps per minute, the average length of the step being 64.0 cm. With an increase in speed we note an increase not only in the number of steps per minute, but likewise in the length of the step, the number of steps per minute for the fast walking being 152.4 and the length of the step 94.8 cm. When we consider the height to which the body was raised, we find that the average height per minute was greatly increased as the speed increased, that obtained at the medium rate being more than twice the value secured with the low speed, i. e., an increase from 2.88 meters to 6.69 meters. When the average values obtained at the medium and fast speeds are compared, we find that although the average distance walked per minute increased from 109.0 meters to 144.5 meters and the number of steps taken per minute from 130.9 to 152.4, the increment in the distance over which the body was raised was but 1.21 meters. Of special interest in this connection is the extraordinary influence of the change in locomotion from fast walking to running. The increase in the speed was inconsiderable, being but 3.1 meters per minute. We find, however, that in running the average height to which the body was lifted increased from 7.90 meters to 13.76 meters per minute and the average number of steps from 152.4 _to 181.9 per minute, the latter increment not being at all in proportion to the former. On the other hand, the length of the step was decreased in running from 94.8 cm. to 81.1 cm. It is thus apparent that in running the steps were taken much more rapidly and considerably shortened and that the body was raised to a much higher point at each step. In any analysis of the mechanics of forward progression, therefore, we should bear in mind the fact that in running the body is actually lifted to nearly twice the height that it is raised during walking. This would of itself involve mechanical work not directly contributory to the work of forward progression and we should therefore expect to find, on this basis alone, that the work of walking would be much more economi- cally done than the work of running. On the other hand, it has been pointed out in connection with the fast walking experiments that the subjects involuntarily, or possibly as the result of previous training, did an excessive amount of muscular work with the arms which likewise was not contributory to the motion of forward progression. If, how- ever in comparing the values for the heat output per unit of work done in walking and running, we use as a base-Une the value obtained with the subject standing and swinging his arms vigorously, we find that the advantage stiU lies with the walking rather than with the running. Thus, while the heat-output per unit of work with the subject running without food is 0.806 gram-calorie with the standing relaxed base-line, when we use the base-line obtained with the subject swinging his arms the value becomes 0.780 gram-calorie for the 7 periods without food and 100 ENERGY TRANSFORMATIONS DURING HORIZONTAL WALKING. 0.756 gram-calorie for the three periods with food, with an average value of approximately 0.768 gram-calorie. While these experiments are relatively few in number, it is quite possible that the apparent lower value obtained for walking as compared with running, after deducting the standing active base-line, may be explained by the fact that the body is elevated nearly twice as' much when running as when walking and that to cover the same distance about 20 per cent more steps were needed. In the motion of forward progression we have therefore to take into account, first, the extraneous muscular activities not directly contributory to the work of forward progression, chief among these being the extraneous muscular movements of the arms in rapid walking which require the expenditure of a considerable amount of energy. Second, the type of step or the gait plays an important r61e, since in raising the body work is performed, requiring an expenditure of energy. When the body is raised approximately 14 meters per minute during running, it may be readily computed that with a man weighing 70 kilograms, this would be equivalent to raising 980 kilograms one meter in one minute corresponding to over 2.2 large calories. Any tjrpe of locomotion, therefore*, which minimizes the raising of the body is the most economical. As a natural outcome of the study it will be seen that it would be desirable for athletes and others interested in the work of forward progression to develop a gait which will eliminate these two apparently unnecessary and extraneous factors, each of which requires the expenditure of a considerable amount of energy which is not directly contributory to the motion of forward progression. Nutrition Laboratory of the Carnegie Institution of Washington. Boston, Massachusetts, June 19, 1915.