Temperature Regulation in Humans and Other Mammals

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1 Temperature Regulation in Humans and Other Mammals

2 Springer Berlin Heidelberg New York Barcelona Hong Kong London Milan Paris Singapore Tokyo

3 Claus Jessen Temperature Regulation in Humans and Other Mammals With 100 Figures Springer

4 Professor Dr. Claus Jessen Justus-Liebig-UniversiHit Giegen Physiologisches Institut Aulweg Giegen Germany and Redwitzstr. 5A Viereth-Trunstadt Germany ISBN-I3 : Springer-Verlag Berlin Heidelberg New York Library of Congress Cataloging-in-Publication Data Jessen, Claus, Temperature regulation in humans and other mammals I Claus Jessen p. cm. Includes bibliographical references and index. ISBN- I 3 : DOl: / e-isbn-13: Body temperature--regulation. I. Title. QP135.J '619--dc This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitations, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. Springer-Verlag Berlin Heidelberg New York a member of BertelsmannSpringer Science+Business Media GmbH Springer-Verlag Berlin Heidelberg 2001 Softcover reprint of the hard cover 1 st edittion The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free general use. Cover Design: design & production, Heidelberg Typesetting: Camera-ready by the author SPIN: /3130XZ Printed on acid free paper

5 Preface In 1996, environmental physiology was the subject of two volumes published in the Handbook series of the American Physiological Society. The largest section deals with temperature regulation. It covers the research ofthe preceding 30 years and is an excellent reference book for advanced scientists working in this or closely related fields. However, handbooks are rarely read from cover to cover. This text is intended to fill the gap between detailed information for specialists and the more cursory treatment of temperature regulation in general textbooks of physiology. It is a brief comprehensive presentation, melding the results of recent research into the background knowledge collected in earlier periods. The intended audience for this book includes students, teachers, physicians and scientists interested in various aspects of thermal physiology, such as exercise, adaptation, circulation, fluid balance or physiological regulation in general. Any other reader is welcome to share my interest in this fascinating field of integrative physiology. Viereth-Trunstadt, September 2000

6 Contents 1 Introduction 1.1 Poikilotherms and Homeotherms 1.2 The Capacity of the System Homeothermy in Terms of Control Theory A First Approach to the Controller 6 2 The Skin as a Source of Temperature Signals The Inner Body as a Source of Temperature Signals Identifying Single Feedback Loops 3.2 Hypothalamus Spinal Cord Other Thermosensitive Sites of the Body Core The Neuronal Basis of Temperature Reception 4.1 Skin Thermoreceptors Afferent Processing of Skin Temperature Signals Core Temperature Sensors Outside the Central Nervous System Thermosensitive Neurons in the Central Nervous System Neuronal Temperature Transduction Sensitivity to Local and Remote Temperatures Heat Production and Heat Balance of the Body The Heat Balance Equation Body Mass, Heat Exchange and Metabolic Rate Metabolic Heat Production vs. Metabolic Rate Basal Heat Production vs. Resting Heat Production Shivering Non-Shivering Thermogenesis in Brown Adipose Tissue Cold-Induced Non-Shivering Thermogenesis in Other Tissues 36

7 VIII Contents 6 Physics of Heat Exchange with the Environment 6.1 Conduction Convection Natural Convection Forced Convection The Heat-Transferring Medium 6.3 Radiation Long-Wave Infrared Solar Radiation 6.4 Evaporation Sweating Panting External and Internal Insulation 7.1 External Insulation: Fur Fur and Solar Radiation Fur and Exercise 7.2 Internal Insulation: Fat and the Principle of Core and Shell Core Temperature vs. Mean Body Temperature Internal Conductance 7.3 Total Conductance The Temperature Field of the Body Core Passive Temperature Deviations Where to Measure a "Representative" Body Core Temperature Regulated Temperature Deviations Selective Brain Cooling Testes Cooling Behavioural Control of Heat Exchange with the Environment 9.1 Locomotion Orientation and Posture 9.3 Wallowing and Saliva Spreading 9.4 Social Behaviour 9.5 Operant Behaviour Autonomic Control of Dry Heat Loss from the Skin 10.1 Pilomotion Peripheral Blood Flow

8 Contents IX Skin Blood Flow and Skin Temperature Anatomy of Peripheral Circulation The Central Drive Local Effects of Skin Temperature Specialized Surface Regions Autonomic Control of Evaporative Heat Loss 77 1l.l Sweating Types of Sweat Glands and Modes of Transmission Sweat Rate: Central Drive and Local Effects Panting 11.3 Sweating vs. Panting Interaction of Various Body Temperatures in Control of Thermoregulatory Responses Core Temperature as a Compound Input Selective Brain Cooling: a Special Pattern of Core Temperatures Natural Spinal Cord Warming Skin and Core Temperatures Autonomic Effector Mechanisms Behavioural Responses 12.3 The Regulated Variable The Central Interface Between Afferent Temperature Signals and Efferent Drives The Controller: Concepts and Facts 13.2 Neuronal Models 13.3 The Set-Point Problem Short-Term Temperature Regulation in Various Environments: Inputs and Responses 14.1 The Medium Range l.l The Thermoneutral Zone 14.2 Short-Term Exposure to Cold 14.3 Short-Term Exposure to Heat

9 X Contents 15 Exercise in the Heat: the Ultimate Challenge 15.1 Cardiovascular System 15.2 Body Fluid Balance Changes of Set-Point 16.1 Ovarian Cycle and Circadian Rhythm, But Not Sleep 16.2 Fever Pyrogens and Fever Adaptation to Cold 17.1 Smaller Animals Hibernation Daily Torpor Everybody Starts Small 17.2 Larger Animals 17.3 Humans Adaptation to Heat 18.1 Smaller Animals 18.2 Larger Animals 18.3 Humans Pathophysiology of Temperature Regulation 19.1 Hyperthermia and Heat Stroke 19.2 Hypothermia References 169 Subject Index 191

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