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Exercise Physiology PDF

513 Pages·2001·10.94 MB·English
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PREFACE This book joins the People and Ideas series of the American Physiological Society (APS), and we thank the Society for the opportunity to present various perspectives on the history of exercise physiology. Although APS acknowledgment of the exis tence of exercise physiology as a subdiscipline of physiology is a relatively recent development (1977), it evolved more than a century ago from the classrooms and laboratories of physical education departments whose leaders were usually physi cians (2,3). The People and Ideas series currently consists of five volumes covering the his tory of the circulation of the blood (edited by Fishman and Richards), renal physiol ogy (edited by Gottshalk, Berliner, and Giebisch), endocrine physiology (edited by McCann), membrane physiology (edited by Tosteson), and respiratory physiology (edited by West). Since exercise elicits integrated physiological responses from vir tually all systems of the body, the sixth book of the series was devoted to the history of exercise physiology with an emphasis on how contemporary ideas evolved within the respective systems. Before the responses of major body systems are discussed in the following chap ter, Berryman provides a historian's perspective on ideas about exercise and its phys iological responses from the time of Hippocrates (460-370 B.C.) to that of Edward Smith (1818-1874). Since the central command associated with the initiation of ex ercise elicits motor, circulatory, respiratory, and autonomic responses (1), the early chapters follow this sequence. However, the focus on systems created problems of se lectivity for the authors. Thus, it is not surprising that many were apologetic to their contemporaries for not citing their important findings. Recall that the recent Hand book of Physiology volume on exercise contained references to more than 8200 stud ies in 1183 pages concerned with the physiological responses from five different sys tems (1). The integrative nature of exercise does not elicit equal involvement from all systems. Some are more important than others. This consideration plus a con straint on space led to various emphases within the text: namely, a separate chapter on maximal oxygen consumption, a focus within the autonomic nervous system Vll viii PREFACE chapter on cardiorespiratory responses, an emphasis in the endocrine chapter on metaboUc reactions, and a reference to muscle function in the chapter on aerobic me- tabohsm. Scientific societies have few historians within their membership, and the Amer ican Physiological Society is no exception. Authors invited to present a historical perspective in this book were selected for their demonstrated expertise in the system being discussed and for their experience as physiologists. Every chapter was reviewed by two or more physiologists with equal expertise and experience. The purpose was to retain the individual perspective of the author while verifying the historical accu racy of the material. Time will tell if we were successful. Besides the History Subcommittee of the Publications Committee of APS, thanks are rendered to Jeffrey House and his colleagues at Oxford University Press for their efforts in helping make an idea become a reality. Tucson, Arizona C.M.T. REFERENCES 1. Rowell, L.R. and J.T. Shepherd, Editors, Handbook of Physiology. Section 12, exercise: regulation and integration of multiple systems. New York: Oxford University Press, 1996. 2. Tipton, CM., Exercise physiology, part II: a contemporary historical perspective. In: A History of Exercise and Sport Sciences, edited by R.W. Swanson and J. Massengale. Champaign, IL: Human Kinetics PubHshers, 1997, pp. 396-428, 431-438. 3. Tipton, CM. Contemporary exercise physiology: Fifty years after the closure of the Harvard Fatigue Laboratory. In: Exercise and Sport Sciences Reviews, edited by John O. Holloszy. Baltimore: WiHiams and Wilkins, A Waverly Company, 1998, Vol. 26, pp. 315-340. CONTRIBUTORS R. James Barnard, Ph.D. Carl V. Gisolfi, Ph.D. (deceased) Department of Physiological Science Department of Exercise Science University of California, Los Angeles University of Iowa Los Angeles, California Iowa City, Iowa Jack W. Berryman, Ph.D. L. Bruce Gladden, Ph.D. Department of Medical History and Department of Health and Human Ethics Performance University of Washington Auburn University School of Medicine Auburn, Alabama Seattle, Washington John O. HoUoszy, M.D. George A. Brooks, Ph.D. Division of Geriatrics and Gerontology Exercise Physiology Laboratory Washington University Department of Integrative Biology School of Medicine University of California, Berkeley St. Louis, Missouri Berkeley, California Alan J. McComas, M.B., F.R.C.P(C.) Elsworth R. Buskirk, Ph.D. Professor Emeritus Noll Physiological Research Center Department of Medicine The Pennsylvania State University Division of Neurology University Park, Pennsylvania McMaster University Hamilton, Ontario Jerome A. Dempsey, Ph.D. Canada Department of Preventive Medicine University of Wisconsin Medical Jere H. Mitchell, M.D. School Harry S. Moss Heart Center Madison, Wisconsin University of Texas Southwestern Medical Center Dallas, Texas XI Xll CONTRIBUTORS Jacques R. Poortmans, Ph.D. Charles M. Tipton, Ph.D. Professor Emeritus Department of Physiology Department of Physiological Chemistry University of Arizona Institut Superieur d'Education Physique Tucson, Arizona et de Kinesitherapie Universite Libre de Bruxelles Evangelia Tsiani, Ph.D. Brussels, Belgium Department of Community Health Sciences Michael C. Riddell, Ph.D. Brock University Department of Kinesiology and Health Faculty of Applied Health Science Science Brock University York University St. Catharines, Ontario Faculty of Pure and Applied Science Canada Toronto, Ontario Canada Mladen Vranic, M.D., D.Sc, F.R.S.C. F.R.C.P(C.) Loring B. Rowell, Ph.D. Department of Physiology Department of Physiology and University of Toronto Biophysics Toronto, Ontario University of Washington Canada School of Medicine Seattle, Washington Brian J. Whipp, Ph.D., D.Sc. Division of Respiratory and Critical Neil B. Ruderman, M.D., D. Phil. Care Physiology and Medicine Department of Medicine Harbor-UCLA Medical Center Boston University School of Medicine Torrance, California Boston, Massachusetts Edward J. Zambraski, Ph.D. Bengt Saltin, M.D., Ph.D. Department of Cell Biology and The Copenhagen Muscle Research Neuroscience Centre Rutgers University Rigshospitalet and University of Piscataway, New Jersey Copenhagen Copenhagen, Denmark chapter 1 ANCIENT AND EARLY INFLUENCES Jack W. Berryman I DENTIFYING and analyzing the ancient and early influences on what would become the discipline known today as exercise physiology are formidable tasks. The range of disciplines involved and the time period of potential influences are both expansive, and the fact that no identifiable field of exercise physiology existed before the mid-nineteenth century is particularly troublesome. Accordingly questions con cerning what to read, where to look, and what years to examine loom large. Whereas the other authors in this volume could focus on a particular organ system, I did not have that luxury. This chapter begins with the contributions of Hippocrates (460-370 B.C.), the most influential physician from Greek antiquity, and then moves chronologically through relevant geography. Specific sections are devoted to the ancient period, the medieval period, the sixteenth century, the seventeenth century, the eighteenth cen tury, and to part of the nineteenth century. Most of the general thought and study concerning physiology during these years took place within the larger context of medicine and medical schools. For example, it was not until the sixteenth century that the French physician Jean Fernel (1497-1558) became the first to apply the term ''physiology'' to the science of the functions of the body. Concerning the physiolog ical aspects of exercise, most of the relevant information published appeared in liter ature dealing with the value of exercise in promoting health and preventing disease. Some of it also appeared in the early literature relating to athletic performance. Gen erally, then, those engaged in the physiological study of exercise were physicians by training. This continued to be the case in the seventeenth, eighteenth, and nineteenth centuries in France, Germany, Italy, England, and the United States. Those who did 2 EXERCISE PHYSIOLOGY most of the early research related to the effects of exercise on the body were physi cians. Some also held appointments at the most prestigious medical schools. The chapter concludes in the mid- to late nineteenth century when the term "physiology of exercise'' began to appear. The American physician William Byford (1817-1890) first used the term in 1855 in an article published in The American journal of Medical Sciences. He was followed in the 1880s by another American physician, Edward Hartwell (1850-1922), who pubUshed two articles on the physiol ogy of exercise in the Boston Medical and Surgical Journal. Also, Fernand Lagrange (1846-1909), a French physician, authored a book entitled the Physiology of Bodily Exercise in 1888. Because of the unique nature of much of the early material, several passages in the actual words of the writer are reproduced below. In these writings illustra tions were not very plentiful, so only paintings of individual scientists and title pages from the more prominent books are available to embellish the material within this chapter. ANCIENT PERIOD To grasp the underlying conceptions of physiological processes upon which the an cient Greek and Roman physicians based their medical theory, and consequently their practice, is ultimately a frustrating endeavor. The most prominent and influen tial physician from Greek antiquity was Hippocrates (460-370 B.C.). Unfortunately, the collection of medical writings attributed to his influence, the Corpus Hippo- craticum, is filled with physiological explanations that are mostly inconsistent and, from a modern perspective, completely nonsensical. Translator W. H. S. Jones pointed out that "We must not expect of them [Hippocratic works] too much con sistency, too much conformity with experience, too much scientific method. We must realise that they are in part works of imagination, often figurative, allusive and metaphorical. They portray truth, or what the writers consider to be truth, in an allegorical disguise'' (43; p. xxiv). He also noted that "the arts were distinguished from the sciences only when Greek thought was past its zenith" and that the Greeks were known for their "luxuriant" imagination (43; p. xxiii). However, it is fascinat ing to consider that although physicians from this era relied heavily upon a luxuri ant imagination for their view of the systems of the body, the therapies they pre scribed and the advice they provided for the preservation and restoration of health were based almost solely on experience and observation—clear, empirical knowledge. So, although their explanations of exact physiological processes were often quite fan tastical, their careful observations and their reliance on experience made their prac tice better than their theory. Hippocrates is considered "the founder of Western scientific medicine," [65; p. 4). Though the term "physiology" was coined only in the sixteenth century, the early Hippocratic physicians were intent on understanding the functioning of the human body and its specific organic processes. But they pursued knowledge of the Ancient and Early Influences 3 functioning of the human body in order to improve their abihty to preserve and re store the health of their patients. And they examined exercise in relation to training for athletic competition in light of what were deemed excesses or deviations from normalcy. The ancient Greek and Roman physicians' application of close observation dis tinguished them as the first practitioners of the scientific method, embryonic though it may have been. According to Rothschuh, because of their ''purely empirical proce dures and practices . . . medicine was the first discipline to come into its own/' and consequently served as a starting point for other scientific disciplines, including physiology {6S; pp. 2-5). It appears that Hippocrates authored two separate works on healthful living— Regimen in Health, with nine very short chapters, and Regimen, composed of four long sections or books. The first seven chapters of Regimen in Health offered advice on the preservation of health and were directed to the layman. Advice was given on what to drink and eat at certain times of the year. Rapid walking was suggested in winter and slow in summer, emetics and clysters were recommended for the bowels, and a chapter was devoted to athletic training. Regimen's four books expressed many of the same physiological principles and considerations as Regimen in Health. In Book I, Hippocrates emphasized that when establishing a regimen, one must exert careful attention "to proportion exercise to bulk of food, to the constitution of the patient, to the age of the individual, to the sea son of the year, to the changes in the winds, to the situation of the region in which the patient resides, and to the constitution of the year.'' Hippocrates clearly had some notion of what we call metabolic processes in relation to exercise: eating alone will not keep a man well; he must also take exercise. For food and exercise, while possessing opposite qualities, yet work together to produce health. For it is the na ture of exercise to use up material, but of food and drink to make good deficiencies. And it is necessary, as it appears, to discern the power of the various exercises, both natural exercises and artificial, to know which of them tends to increase flesh and which to lessen it. (43; p. 229) In their article "Hippocratic and Galenic Concepts of Metabolism," C. E. A. Winslow and R. R. Bellinger concluded from this statement that Hippocrates "postulated [the] relation between the need for fuel foods and muscular activity, bringing us almost to the threshold of quantitative metabolism" (84; p. 130). In Book II, Hippocrates also explained the excretory effects of exercise: Early morning walks too reduce [the body], and render the parts about the head light, bright and of good hearing, while they relax the bowels. They reduce because the body as it moves grows hot, and moisture is thinned and purged, partly by the breath, partly when the nose is blown and the throat cleared, partly being consumed by the heat of the soul for the nourishment thereof. (43; pp. 351-353) According to Winslow and Bellinger, this statement suggests a connection between respiration and exercise (84; p. 132). The statement that "running in a circle dissolves flesh least, but reduces and contracts the belly most, because, as it causes the most 4 EXERCISE PHYSIOLOGY rapid respiration, it is the quickest to draw moisture to itself/' more explicitly sup ports this connection (43; p. 355). Hippocrates emphasized the reduction of "flesh'' through exercise in Book II. He observed that athletes with "physiques of a less firm flesh and inclined to be hairy are more capable of forcible feeding and of fatigue/' and that "their good con dition is of longer duration" (43; pp. 55-57). With regard to hydration and excessive exercise, Hippocrates advised: When people are attacked by thirst, diminish food and fatigue, and let them drink their wine well diluted and as cold as possible. Those who feel pains in the abdomen after ex ercise or after other fatigue are benefited by resting without food; they ought also to drink that of which the smallest quantity will cause the maximum of urine to be passed, in order that the veins across the abdomen may not be strained by repletion. (43; p. 57) In Regimen, Hippocrates also discussed "trained bodies" and "untrained people," es pecially in relation to "fatigue pains," or the potential for muscle soreness: The fatigue pains that arise in the body are as follow Men out of training suffer these pains after the slightest exercise, as no part of their body has been inured to any exer cise; but trained bodies feel fatigue pains after unusual exercises, some even after usual exercises if they be excessive. These are the various kinds of fatigue pains; their prop erties are as follow. Untrained people, whose flesh is moist, after exercise undergo a con siderable melting, as the body grows warm. Now whatever of this melted substance passes out as sweat, or is purged away with the breath, causes pain only to the part of the body that has been emptied contrary to custom; but such part of it as remains be hind causes pain not only to the part of the body emptied contrary to custom, but also to the part that has received the moisture, as it is not congenial to the body but hostile to it. It tends to gather, not at the fleshless, but at the fleshy parts of the body, in such a way as to cause them pain until it has passed out. (43; pp. 359-361). His explanation for those in training suffering "fatigue pains from unaccustomed ex ercises" was that any unexercised part of the body must of necessity have its flesh moist, just as persons out of training are moist generally throughout. So the flesh must of necessity melt, se crete itself and collect itself, as in the former case. ... Accustomed exercises should be practised, so that the collected humour may grow warm, become thin, and purge itself away, while the body generally may become neither moist nor yet unexercised. (43; pp. 361-363) For accustomed exercises causing pain, Hippocrates said that "moderate toil is not followed by pain; but when immoderate it dries the flesh overmuch, and this flesh, being emptied of its moisture, grows hot, painful and shivery, and falls into a longish fever" (43; p. 363). Clearly, the most important figure in medicine and physiology since Hip pocrates was Claudius Galenus—or Galen (ca. 129-200 A.D.). His theories and writ ings dominated medicine and were unchallenged at least through the later Middle Ages, and were still widely accepted during the Renaissance. He was born in Perga- mum—a city in the Roman empire on the western coast of Asia Minor, now Ancient and Early Influences 5 Turkey—and went on to acquire an extensive education in philosophy and medicine in the cities of Smyrna, Corinth, and Alexandria. During this time, he did extensive study in anatomy based largely on animal structures (71). Around 158-159 A.D., Galen returned to Pergamum to serve as the physician for the gladiators, perform ing his duties in the Asclepeion. Here he not only learned anatomy, physiology, and surgery, but also experimented with various training regimens to produce overall health, strength, and endurance for the combatants {67). Galen was a prominent author as well as a physician, and wrote about four hun dred treatises on mostly medical and philosophical subjects. Some of his works were commentaries on the Hippocratic writings—which he respected and used—but the majority were unique and innovative. Galen's extensive systematic medical theory also had its basis in the Hippocratic wisdom, but he moved far beyond it because he based his beliefs on anatomical and physiological facts acquired through observation and dissection. While his anatomy had some shortcomings, he far exceeded his pre decessors and went on to establish anatomy as the basis for medicine as well as phys iology. In fact, Galen's physiological concepts dealt with the physiology related to specific organs as well as to a general theory of vitality. Central to what became known as ''Galenic medical theory" or in some cases "humoral theory" was Galen's identification of the "naturals" (referring to physiol ogy), the "non-naturals" (things not innate—hygiene), and the "contra-naturals" (against nature—pathology). The non-naturals were six in number and included those things which humans were exposed to in daily life: (1) air and environment, (2) food and drink, (3) motion and rest, (4) sleep and wake, (5) excretions and reten tions, and, (6) passions of the mind (emotions). The non-naturals needed to be uti lized in moderation as to quantity, quality, time, and order; for if they were taken in excess or put into imbalance, Galen believed disease would result. In particular, the role of motion (exercise) and rest in his theory was related to their respective effects on the qualities and humors. Galen believed that excessive rest would increase cold and moisture, whereas excessive exercise would at first heat the body and then pro duce cold and dryness. Moderate exercise would maintain warmth. Since causes for disease were thought to be due to heat, cold, dryness, or moisture, exercise and the other five non-naturals played important roles in therapy. Therefore, regulating the non-naturals along with drug prescription and surgery were the key components of the physician's activities. However, the non-naturals played a more significant role in hygiene. At the time, hygiene referred to the science of health and its preserva tion. Consequently, those liable to illness or those wishing to maintain their health- fulness were given a regimen by their physician whereby the non-naturals were reg ulated to provide a proper balance (4). Most of Galen's material on exercise was in the first three books of On Hygiene. Book 1, "The Art of Preserving Health," was composed of 15 chapters. Chapter 8 was entitled, "The Use and Value of Exercise" and dealt mainly with the need for motion at all ages. Whether by sailing, riding on horseback, driving, or via cradles, swings, and arms, everyone, even infants, Galen said, needed exercise (32). Other chapters pertained to bathing and massage, fresh air, beverages, and evacuations. "Exercise and 6 EXERCISE PHYSIOLOGY Massage" was the title of Book 2 and it comprised 12 chapters. Chapter 2, "Purposes, Time, and Methods of Exercise and Massage,'' included some very important mate rial on the role exercise played in Galen's conception of hygiene. For example, in ref erence to the type and definition of exercise, Galen said: To me it does not seem that all movement is exercise, but only when it is vigorous. But since vigor is relative, the same movement might be exercise for one and not for an other The criterion of vigorousness is change of respiration; those movements which do not alter the respiration are not called exercise. But if anyone is compelled by any movement to breathe more or less faster, that movement becomes exercise for him. This therefore is what is commonly called exercise or gymnastics. (32; pp. 53-54) He saw the uses and values of exercise as follows: The uses of exercise, I think, are twofold, one for the evacuation of the excrements, the other for the production of good condition of the firm parts of the body. For since vig orous motion is exercise, it must needs be that only these three things result from it in the exercising body—hardness of the organs from mutual attrition, increase of the in trinsic warmth, and accelerated movement of respiration. These are followed by all the other individual benefits which accrue to the body from exercise; from hardness of the organs, both insensitivity and strength for function; from warmth, both strong attrac tion for things to be eliminated, readier metabolism, and better nutrition and diffusion of all substances, whereby it results that solids are softened, liquids diluted, and ducts dilated. And from the vigorous movement of respiration the ducts must be purged and the excrements evacuated. (32; p. 54) Space does not permit elaboration, but Galen's ideas about the proper time for exer cise, factors to consider before exercise, the varieties of exercise, the different quali ties of exercises, and the places for exercise are very perceptive. In The Pulse for Beginners, Galen discussed the impact of exercise on the pulse in a section on "the result of non-natural causes": Exercise to begin with—and so long as it is practised in moderation—renders the pulse vigorous, large, quick, and frequent. Large amounts of exercise, which exceed the capac ity of the individual, make it small, faint, quick, and extremely frequent. In cases of great excess, whereby the subject is scarcely still able to move, and only at great inter vals if at all, and there is a considerable loss of power, the pulse becomes very small, faint, slow, and sparse. If a state of dissolution of the faculty is reached, then the pulse will be that specific to this state; the nature of this pulse will be discussed in due course. (30; p. 332) In Good Condition, Galen identified "good condition" as "a kind of excellence of health, and is thus found in those bodies which have the best constitutions." Ath letes, as Galen saw in the writings of Hippocrates, were viewed as being purveyors of "the extremes of good condition" and were therefore seen as being "dangerous." Galen himself argued that "athletes' health, for example, is so far from being highly desirable that it was rightly criticized" and explained that athlete's strive for "the ac quisition not only of good mixture, but also of physical mass—which cannot take place without an ill-balanced type of fiUing. And thus the state is rendered both dan-

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