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Thermoregulation and Human Performance
Medicine and Sport Science
Vol. 53

Series Editors

J. Borms Brussels
M. Hebbelinck Brussels
A.P. Hills Brisbane
T. Noakes Cape Town
Thermoregulation and
Human Performance
Physiological and Biological Aspects

Volume Editor

Frank E. Marino Bathurst

28 figures, and 1 table, 2008

Basel · Freiburg · Paris · London · New York · Bangalore ·

Bangkok · Shanghai · Singapore · Tokyo · Sydney
Medicine and Sport Science
Founder and Editor from 1969 to 1984: E. Jokl†, Lexington, Ky.

Frank E. Marino, PhD

School of Human Movement Studies and
Exercise and Sports Science Laboratories
Charles Sturt University
Bathurst, NSW 2795 (Australia)

Library of Congress Cataloging-in-Publication Data

Thermoregulation and human performance : physiological and biological aspects /

volume editor, Frank E. Marino.
p. ; cm. -- (Medicine and sport science, ISSN 0254-5020 ; v. 53)
Includes bibliographical references and indexes.
ISBN 978-3-8055-8648-1 (hard cover : alk. paper)
1. Body temperature--Regulation. 2. Exercise--Physiological aspects. 3.
Heat--Physiological effect. I. Marino, Frank E. II. Series.
[DNLM: 1. Body Temperature Regulation--physiology. 2. Athletic
Performance--physiology. 3. Exercise--physiology. 4. Heat Stress
Disorders--physiopathology. W1 ME649Q v.53 2008 / QT 165 T4112 2008]
QP135.T482 2008
612⬘.01426--dc22
2008031568

Bibliographic Indices. This publication is listed in bibliographic services, including Current Contents®
Disclaimer. The statements, opinions and data contained in this publication are solely those of the individual authors and
contributors and not of the publisher and the editor(s). The appearance of advertisements in the book is not a warranty,
endorsement, or approval of the products or services advertised or of their effectiveness, quality or safety. The publisher and
the editor(s) disclaim responsibility for any injury to persons or property resulting from any ideas, methods, instructions or
products referred to in the content or advertisements.
Drug Dosage. The authors and the publisher have exerted every effort to ensure that drug selection and dosage set forth in
this text are in accord with current recommendations and practice at the time of publication. However, in view of ongoing
research, changes in government regulations, and the constant flow of information relating to drug therapy and drug
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employed drug.
All rights reserved. No part of this publication may be translated into other languages, reproduced or utilized in any form or
by any means electronic or mechanical, including photocopying, recording, microcopying, or by any information storage
and retrieval system, without permission in writing from the publisher.
© Copyright 2008 by S. Karger AG, P.O. Box, CH–4009 Basel (Switzerland)
www.karger.com
Printed in Switzerland on acid-free and non-aging paper (ISO 9706) by Reinhardt Druck, Basel
ISSN 0254–5020
ISBN 978–3–8055–8648–1
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Section Title

Contents

VII Preface
Marino, F.E. (Bathurst, NSW)

1 The Evolutionary Basis of Thermoregulation and Exercise Performance

Marino, F.E. (Bathurst, NSW)
14 Comparative Thermoregulation and the Quest for Athletic Supremacy
Marino, F.E. (Bathurst, NSW)
26 Thermoregulation, Fatigue and Exercise Modality
Tucker, R. (Cape Town)
39 Neuromuscular Response to Exercise Heat Stress
Cheung, S.S. (St. Catharines, Ont.)
61 Intestinal Barrier Dysfunction, Endotoxemia, and Gastrointestinal
Symptoms: The ‘Canary in the Coal Mine’ during Exercise-Heat Stress?
Lambert, G.P. (Omaha, Nebr.)
74 Effects of Peripheral Cooling on Characteristics of Local Muscle
Drinkwater, E. (Bathurst, NSW)
89 Cooling Interventions for the Protection and Recovery of Exercise
Performance from Exercise-Induced Heat Stress
Duffield, R. (Bathurst, NSW)
104 Ethnicity and Temperature Regulation
Lambert, M.I.; Mann, T. (Newlands); Dugas, J.P. (Chicago, Ill.)
121 Exercise Heat Stress and Metabolism
Mündel, T. (Palmerston North)

130 Author Index

131 Subject Index
Section Title

Preface

There are many excellent books which examine the relationship between ther-
moregulation and human performance. The collection of papers in this present
book is in no way meant to replace those texts but rather add to the stock of knowl-
edge in this area of human physiology. Over the past century and in particular the
last decade, our understanding of the relationship between thermoregulation, per-
formance and fatigue has changed dramatically. For example, early texts on the
general subject of muscular exercise gave only a cursory account of the effect of ris-
ing body temperature on exercise [1, pp. 28–31]. Nevertheless, the common thread
in many seminal texts could be summarized by the statement ‘The ill-effects
become more marked as the temperature of the body rises, until they culminate in
the disorganization of the bodily functions, which constitute heat-stroke’ [1, p. 31].
However, the majority of research since the seminal work of Pembrey and Nicol [2]
shifted to understanding the cardiovascular responses with increasing body tem-
perature because it was noted that cardiovascular strain was indeed increased dur-
ing times of thermal stress. The development of muscle biopsy techniques and
radioactive tracer studies permitted the study of rising and decreasing body tem-
perature on metabolism. Further development of electromyography and electrical
stimulation techniques have provided insights into the role the nervous system
plays in determining human performance under thermally stressful situations.
Even with the technological and methodological advances, we still do not com-
pletely understand why thermal strain induces premature fatigue other than to
presume that this is to protect the organism from irreversible cellular damage.
This fact alone has necessitated further analysis of previous research in addition to
considering theories derived from complimentary areas of research such as evolu-
tionary biology, anthropology and cellular and molecular biology. This book is an
attempt to propose different interpretations of the research which might explain
the behavioral and physiological processes which could increase our understand-
ing of the limitations of thermoregulation and in so doing also explain the limits
of human performance.
The Earth provides extreme environments with temperature differences rang-
ing from about –80⬚C at the South Pole to well over 50⬚C in the Sahara desert. It is
not surprising then that temperature has a universal effect on life and that all life
processes depend very much on temperature so that the necessary chemical reactions
and subsequent biological processes take place when just the right thermal envi-
ronment is present. However, exercise physiologists have not, as a general rule,
considered the evolutionary history of humans as a possible avenue for answers
regarding the relationship between thermoregulation and human performance.
An attempt has been made by Marino in the first two chapters of this book to turn
back the clock and re-discover the foundations of human evolution and the forces
which may have shaped our physiology together with a comparison of the ther-
moregulatory strategies employed by other mammals.
A novel feature of this book is the inclusion of a ‘new understanding’ of exercise
performance which claims that humans, like other mammals, are able to predict
the requirements for successful performance in order to make use of the limited
physiological capacity. Perhaps the attractiveness of this new school of thought is
related to its ability to explain many observations under many different condi-
tions. Tucker, in his paper, provides evidence for an anticipatory exercise response
particularly when there is impending thermal strain. The centerpiece of this chap-
ter is the consideration of the self-paced exercise modality which, unlike the fixed-
intensity exercise modality, does not mask the individual’s physiological response.
The neuromuscular system provides an exciting avenue to explore the effects of
thermal strain on human performance. The last decade of research has shown that
the central nervous system is deeply affected by increasing body temperature and
that there is a distinct reduction in muscle recruitment when this occurs.
However, the reasons for this remain elusive. Cheung provides a detailed account
of the research in this area and the methodologies by which these observations
have been made. Another novel area of research is that of intestinal barrier dys-
function and endotoxemia during times of increasing thermal strain. Lambert
proposes that the symptoms associated with this response could possibly provide
the organism with a warning signal of impending cellular disruption.
A performance enhancement method which has become increasingly popular
amongst athletes is that of body cooling before exercise in order to reduce thermal
strain but more recently, the application of cooling for recovery from strenuous
physical exercise has been utilized. However, the effects of cooling on local muscle
characteristics are seldom considered when these cooling methods are employed.
Drinkwater reviews the physiological responses and the various methods utilized

VIII Preface
for cooling which could be detrimental to certain modes of exercise. Duffield, in a
complimentary paper, provides an analysis of the possible benefits of cooling for
thermally stressful conditions in addition to the use of body cooling for protection
and recovery from physical exercise.
The morphology of humans is intimately tied to the climate in which various
ethnic groups live. There is a notable difference between the athletic performances
of certain ethnic groups in given events and therefore the assumption that there is
a genetic advantage. Lambert and colleagues provide an in-depth analysis of the
various physical characteristics of ethnic groups and the advantages that these
characteristics bring in certain environmental conditions. The debate between
phenotypic versus genotypic variation is explored.
Finally, the effects of thermal strain on metabolism are considered in the chap-
ter by Mündel. However, this paper considers the consequences of metabolism
and the ability for the brain to deal with these consequences rather than concen-
trate solely on the skeletal muscle requirements during exercise and heat stress.
I would like to acknowledge the very enthusiastic assistance of S. Karger
Publishers in the preparation of this edited volume. The merit of this book lies in
the work of the authors and I am indebted to them for their efforts and coopera-
tion. Last but not least, my sincere gratitude goes to Professor Tim Noakes for his
initial ideas and faith in the Editor!
Frank E. Marino

References
1 Bainbridge FA: The Physiology of Muscular 2 Pembrey MS, Nicol BA: Observations upon the
Exercise, ed 3. New York, Longmans, Green and Co., deep and surface temperature of the human body. J
1931. Physiol (Lond) 1898;23:386–406.

Preface IX
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Section Title
Marino FE (ed): Thermoregulation and Human Performance. Physiological and Biological Aspects.
Med Sport Sci. Basel, Karger, 2008, vol 53, pp 1–13

The Evolutionary Basis of

Thermoregulation and
Exercise Performance
Frank E. Marino
School of Human Movement Studies and Exercise and Sports Science Laboratories,
Charles Sturt University, Bathurst, NSW, Australia

Abstract
The evolutionary history of mammals, but more specifically humans, indicates that heat stress was a
decisive and powerful selection pressure. There is good evidence that early hominids had to adapt to
a changing environment by assuming an upright posture and consequently bipedalism. Because of
further distances between food sources and the need for prolonged hunting, bipedal locomotion
over longer distances required higher aerobic capacities and as a consequence an increase in
endogenous heat production. A cooling mechanism to balance heat loads was essential for survival
and adaptations by other bodily systems such as the brain must have developed to deal with the
increased heat stress. This chapter discusses the evolutionary forces which are thought to have pro-
duced the thermoregulatory system used by modern day humans in exercise performance. A partic-
ular feature that has been overlooked by thermal physiologists is the way in which mammals use the
thermoregulatory system to anticipate thermal limits during physical activity and thus avoid physio-
logical catastrophe. Copyright © 2008 S. Karger AG, Basel

The past century has witnessed extensive research into thermoregulation during
exercise performance. The ability to thermoregulate effectively to avoid lethal dis-
ruption to homeostasis has been an important aspect of the research conducted, in
doing so there is a greater understanding of the limits to performance under
extreme conditions [1, 2]. However, this research highlights the susceptibility of
living organisms to extreme environments and the practical limitations for living
and performing in such conditions.
Perhaps the pinnacle in the understanding of thermoregulation was the inven-
tion of the thermometer by Galileo Galilei c1593 and its subsequent improve-
ments culminating in the publication of Ars de Medicina Statica (1624) by Santorio
Santorio [3]. In this publication, descriptions of temperature measurements in
man were available for comparison to ill health, for at this time it was not yet
known that humans regulated their body temperature around a ‘set point’. What
was clear, however, was that body temperature was a critical part of human physi-
ology especially when fever was present.
The relationship between thermoregulation and human performance is inti-
mately connected to the evolutionary forces which are seldom considered in the
exercise sciences but could provide pivotal insights into the way in which modern
humans deal with exercise heat stress. This chapter will discuss and outline the key
evolutionary forces which are thought to have fashioned the development of the
thermoregulatory system in humans with a view to providing an understanding of
the limitations within which this system functions and perhaps help explain more
recent findings in exercise and human performance.

The Primordial Soup and the Thermal Environment

Although there is no absolute determination of how life on Earth began, there is

consensus that early life was subjected to a hot environment. The hypothesis that life
began in a primordial soup was first postulated in the 1920s independently by both
Oparin and Haldane [4]. In this model, the Earth’s atmosphere was composed of
nitrogen, ammonia, methane and hydrogen whereby heat produced chemical reac-
tions giving rise to molecules that eventually made their way into water forming a
primordial soup of amino acids and thus the building blocks of life [5]. Although the
primordial soup hypothesis is not regarded as the only process by which life evolved,
the point is that heat must have been an essential ingredient in this process.
Due to the sensitivity to fluctuations in environmental temperature, it would be
reasonable to conclude that behavioural motivation was related to the inherent
need to control the internal temperature. From this point of view it was likely that
primitive organisms were able to perform metabolic functions, be excitable, repro-
duce and find ways of moving from one point to another. A graduation from
unicellular to multicellular organisms was most likely a fundamental point in
evolution and thus cellular reactions must have occurred [6]. There is very little
doubt that a pivotal feature in evolution must have been the ability of organisms to
respond to environmental challenges, which could only occur if structures for cel-
lular responsiveness were available. In addition to these molecular and cellular
adaptations, a major advancement in the evolution of hominids was the appear-
ance of mammals [7]. The sequence of events leading to this is beyond the scope
of this chapter and, therefore, the reader is referred to a more in-depth discussion
of the events in this timeline [8].
Suffice it to say that the appearance of mammals is arguably the most important
step in the evolutionary history of homo sapiens, clearly the development of

2 Marino
upright posture and bipedal locomotion was a significant shaping force for our
species [9, 10]. The relationship between upright posture and the ability to effec-
tively regulate body temperature is not readily apparent. However, because bipedal
locomotion permitted hominids to pursue prey over long distances and possibly
for days [11], the production of endogenous heat must have been a significant
impediment. More modern studies show that the cost of running for humans is
relatively high leading to endogenous heat production for a given body mass and
thus a critical factor to deal with during exercise of long duration [12, 13]. To deal
with this inefficiency the development of a heat dissipating mechanism would be
required for continued activity.

Evidence of Preferred Temperature of Organisms

The relationship between the origins of life and temperature were examined in a
classical experiment by Mendelssohn [14] who showed that paramecia reacted to
varying environmental temperatures by either dispersing or congregating
according to their seemingly preferred temperature medium. When the tempera-
ture of their medium was 19⬚C the paramecia randomly dispersed, whereas,
when the temperature was increased to 38⬚C they would congregate at the cooler
end of 25–26⬚C as depicted in figure 1. What is even more remarkable is that the
paramecia had a ‘preferred’ temperature of 24–28⬚C where they would congre-
gate and avoid the extremes of either 12 or 36⬚C (fig. 2). Notably, the paramecia
were attracted toward the moderate rather than the warmer medium suggesting
that optimal function was either linked to temperature or that these unicellular
organisms were capable of controlling their internal temperature by behavioural
means.
Although, the temperature on much of the Earth’s surface ranges from 0–50⬚C,
some animals such as polar fish and invertebrates can live in temperatures below
0⬚C, whilst some algae can survive in temperatures above 70⬚C [15]. There is no
clear evidence as to the ambient temperature in which early hominids evolved
albeit that technology has permitted the ambient temperature range in which
humans can live to be extended considerably. The temperature range in which
humans might have evolved is an important consideration as it could explain why
most animals (mammals and birds) have a set core temperature of about 37⬚C.
The major theories about the significance of 37⬚C have been constructed
around the observation that death ensues in some animals following high thermal
strain. A treatise on each of these is not within the scope of this review so the
reader is referred to other more pertinent texts [15]. Briefly, however, these theo-
ries include (1) denaturation of proteins and thermal coagulation, (2) thermal
inactivation of enzymes, (3) inadequate oxygen supply, (4) different temperature

Evolution and Thermoregulation 3

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