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Page 1: Comprehensive Human Physiology: From Cellular Mechanisms ...978-3-642-60946-6/1.pdf · Comprehensive Human Physiology: From Cellular Mechanisms to Integration, Vol. 1 . Springer-Verlag

Comprehensive Human Physiology: From Cellular Mechanisms to Integration, Vol. 1

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Springer-Verlag Berlin Heidelberg GmbH

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R. Greger U. Windhorst (Eds.)

Com prehensive Human Physiology From Cellular Mechanisms to Integration

Volume 1

With 700 Figures and 84 Tables

Springer

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Prof. Dr. RAINER GRE GER

Universităt Freiburg

Physiologisches Institut

Hermann-Herder-Str. 7

79104 Freiburg

Germany

Prof. Dr. UWE WINDHORST

University of Calgary

Faculty of Medicine

Department of Clinical Neurosciences

3330 Hospital Drive N.W.

Calgary, Alberta

Canada T2N 4N1

ISBN 978-3-642-64619-5 ISBN 978-3-642-60946-6 (eBook)

DOI 10.1007/978-3-642-60946-6

Library of Congress Cataloging-in-Publication Data. Comprehensive human physiology: from cellular mechanisms to integration/R. Greger, U. Windhorst (eds.). p. cm. Includes bibliographical references and index. ISBN 978-3-642-64619-5 (softcover: alk. paper) 1. Human physiology. 1. Greger, Rainer. II. Windhorst, Uwe, 1946- . QP34.5.C656 1996 612-dc20 96-34793

This work is subject to copyright. All rights are reserved, whether the whole Of part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, 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 1996

Softcover reprint of the hardcover 1 st edition 1996 The use of general descriptive names, registered, 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 for general use.

Product liability: The publishers cannot guarantee the accuracy of any information about dosage and appli­cation contained in this book. In every individual case the user must check such information by consulting the relevant litera ture.

Cover design: Springer-Verlag, Design & Production

Typesetting: Best-set Typesetter Ud., Hong Kong

SPIN: 10634097 27/3111/SPS - 5 4 3 2 1 - Printed on acid-free paper

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Preface

Adding yet another physiology text to the many good textbooks that are available around the world seems like carrying coal to Newcastle. In the physiological sciences, there is no lack of good texts written at different levels and targeting different groups of readers. Our motivation to prepare this new book arose from another, yet very traditional desire, namely to provide a textbook that includes recent developments in the field. These developments confront us not only with an enormous growth of knowledge in quantitative terms, but also with the necessity of revising classical concepts and classifications. Two diverging trends characterize recent developments in our field. One originates from advances in cellular physiology that have provided us deep and fascinating insights into the function of the cell and its organelles. The success of the reductionist approach of identifying ever more refined details is conspicuous and undeniable as evidenced by the flood of data on individual membrane receptors, ion channels, and other membrane proteins which, with ever more sophisticated tools, can now be studied in isolation. Regulatory processes in the individual cell are now understood at several molecular levels, spanning acute modifications of existing matter to selec­tive gene expression. In neurophysiology, methodological progress has made it pos­sible to identify, purify, and localize an ever-increasing number of ion channels, transmitters, cotransmitters, and neuromodulators. Perhaps surprisingly, this suc­cess of reductionism that might have been expected to lead to an ever more diversi­fied parcellation of knowledge, has also provided an unanticipated basis for unification. The reductionist approach continues to reveal that diverse organ func­tions are based on much less diverse basic cellular mechanisms, i.e., many special mechanisms are variations of far fewer basic themes. This makes it much easier to understand the bewildering array of new findings. These developments thus promise to reverse trends towards segregation of specialties, such as cell biology, biochemis­try, molecular biology, and biophysics, which may in future be reunited under the wider umbrella of physiology. In addition, these attractive and fascinating new disci­plines are of importance not only from a theoretical, basic sciences viewpoint, but increasingly for the understanding and treatment of diseases. Each student of physi­ology will thus have to understand such basic mechanisms and their control. And so will physicians. These relations are therefore reflected in sections on pathophysiology throughout this book. While the results of extremely sophisticated and specialized studies continue to accumulate, the trend towards integration is also gaining strength. Indeed, this is a necessary reaction and a countermovement to dissection and division. The ultimate challenge to physiologists is to cope with the new results of reductionist research and at the same time to deal with the fundamental question of how the parts and pieces interact. Our goal remains to understand how cells are coordinated to function as an organ, how organs cooperate in systems, and finally, how system functions are integrated in the soma to motor and neurovegetative behavior of the whole organism when adapting to internal and external needs. Physiology has never been and will not be a mere set of facts, but rather a highly dynamic science based on functional thinking and embedded in the fascinating, ever-continuing process oflearning more about life. Our intention not only to include, but also to combine the molecular and integrative aspects of physiology comes at a time when our science is in a state of transition between classical, partly still valid concepts, and entirely new approaches.

v

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Contents

Volume 1

Physiology Past and Future. R. GREGER and U. WINDHORST ..................................... .

Basic Principles and Their Co-ordination

A Homeostasis and Regulation

Regulatory Principles in Physiology U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

2 Physiological Synergetics: A Holistic Concept Concerning Phase Jumps in the Behavior of Driven Nonlinear Systems H. SCHMID-SCHONBEIN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

3 Concepts of Control and Information: A Language for Physiology W.S. YAMAMOTO. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

B Cellular Mechanisms

4 The Cell and Its Membranes R. GREGER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

5 Cellular Transduction Processes R. GREGER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95

6 Cell Communication by Autacoids and Paracrine Hormones R. GREGER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115

7 Cell-to-Cell Communication by Gap Junctions R. GREGER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

8 Cellular Membrane Transport Mechanisms R. GREGER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

9 Connective Tissue: More Than Just a Matrix for Cells G. SIEGEL. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173

10 Nonmuscle Motility and the Actin-Based Cytoskeleton B.M. JOCKUSCH and H. HINSSEN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225

11 Membrane Voltage and Preservation of the Ionic Distribution Across the Cell Membrane R. GREGER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245

VII

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C Nerve and Synaptic Function

12 Ion Channels in Excitable Membranes J.P. RUPPERSBERG . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267

13 Excitation and Nerve Conduction W.F.H.M. MOMMAERTS, D. JUNGE, and M.B. JACKSON. . . . . . . . . . . . . . 283

14 Sensory Transduction and Neural Coding D. JUNGE, W.F.H.M. MOMMAERTS, and U. WINDHORST. . . . . . . . . . . . . 295

15 Innervation of Muscle and Neuromuscular Transmission J.P. RUPPERSBERG and S. HERLITZE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307

16 Synaptic Transmission M. FROTSCHER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321

D Autonomic and Hormonal Control

17 The Autonomic Nervous System: Peripheral and Central Integrative Aspects K.A. SHARKEY and Q.J. PITTMAN. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335

18 The Functions of the Limbic System S.E. Fox. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355

19 The Role of the Hypothalamus in Neuroendocrinology K. KOIZUMI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379

20 Metabolism of CortiCal Steroid Hormones and Their General Mode of Action K. HIERHOLZER and H. BUHLER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403

21 Growth Factors R.A. BRADSHAW................................................ 431

22 The Thyroid Gland. Thyroid Hormones, Their Origin and Their Mechanism of Action R. FINKE, H. SCHLEUSENER, and K. HIERHOLZER. . . . . . . . . . . . . . . . . . 451

23 Hormones Modulate Environmental Control of a Changing Brain B.S. McEwEN. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 473

II The Nervous System

E General Properties of the Central Nervous System

24 Synaptic Circuits and Physiological Operations in the Central Nervous System G.M. SHEPHERD. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 497

25 The Generation of Electric and Magnetic Signals of the Brain by Local Networks F.H. LOPES DA SILVA.......................................... 509

26 Neuron-Glia Interactions in Homeostasis and Degeneration H. KETTENMANN, A. FAISSNER, and J. TROTTER. . . . . . . . . . . . . . . . . . . 533

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27 Blood-Brain Barrier and the Production of Cerebrospinal Fluid W. KUSCHINSKY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 545

28 Circulation in the Central Nervous System L. SOKOLOFF. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 561

29 Cerebral Metabolism and Visualization of Cerebral Activity L. SOKOLOFF....... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 579

30 Establishment of Neural Cell Populations and Interneuronal Connections A. FAISSNER, H. KETTENMANN, and J. TROTTER. . . . . . . . . . . . . . . . . . . 603

F Sensory Systems and Functions

31 Central Projections of Cutaneous and Enteroceptive Senses U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 623

32 Tactile Senses U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 647

33 Temperature Perception and Pain W.D. WILILS, JR . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 677

34 The Vestibular System H.-P. ZENNER and A.W. GUMMER............................... 697

35 Hearing H.-P. ZENNER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 711

36 Central Processing of Auditory Information A.W. GUMMER and H.-P. ZENNER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 729

37 Introduction to Vision W.F.H.M. MOMMAERTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 757

38 Retinal Processing of Vision J.E. DOWLING. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 773

39 Central Processing of Vision U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 789

40 Eye Movements and the Mechanisms of Accommodation and the Pupil R.H.S. CARPENTER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 829

41 Auditory-Visual Interactions in the Superior Colliculus A.W. GUMMER, P. PLINKERT, and H.-P. ZENNER. . . . . . . . . . . . . . . . . . . 839

42 Olfaction and Taste D. SCHILD. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 847

43 From Neuron to Gestalt: Mechanisms of Visual Perception L. SPILLMANN and W.H. EHRENSTEIN . . . . . . . . . . . . . . . . . . . . . . . . . . . . 861

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G Motor Functions and Their Control

44 Smooth Muscle J.C. RUEGG. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 895

45 Physiology of Skeletal Muscle U. WINDHORST and W.F.H.M. MOMMAERTS . . . . . . . . . . . . . . . . . . . . . . . 911

46 Muscle Contraction: Molecular and Cellular Physiology J.C. RUEGG. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 935

47 Genetic Control of Muscle Function and Molecular Basis of Muscle Diseases H. JOCKUSCH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 959

48 The Peripheral Motor Apparatus and Its Nervous System: General Problems to Be Solved and General Problem Solvers U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 969

49 Spinal Cord and Brainstem: Motor Output, Sensors and Basic Circuits U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 987

50 Spinal Cord and Brainstem: Pattern Generators and Reflexes U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1007

51 Cerebral Cortex, Basal Ganglia, and Cerebellum: Parallel Circuits U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1033

52 Higher Psychomotor Functions U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1059

Appendix. Psychomotor Aspects of the Brain-Mind Problem H.-D. HENATSCH ............................................. 1091

53 The Human Voice J. SUNDBERG. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1095

H Integrated Functions of the Central Nervous System

54 Specific Networks of the Cerebral Cortex: Functional Organization and Plasticity U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1105

55 Bilateral Organization of the Brain U. WINDHORST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1137

56 Memory and Learning R.F. THOMPSON and C.G. LOGAN. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1161

57 Sleep J.M. SIEGEL and R.M. HARPER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1183

58 Biological Rhythms and the Pineal Gland J.C. FLOREZ and J.S. TAKAHASHI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1199

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Volume 2

III Intake and Excretion

Epithelial Transport and Gastrointestinal Physiology

59 Epithelial Transport R. GREGER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1217

60 Mastication and Swallowing J.E. KELLOW ....... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1233

61 Gastric Function J.G. FORTE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1239

62 Function of the Intestine A.D. CONI GRAVE and J.A. YOUNG 1259

63 Gastrointestinal Motility and Defecation J.E. KELLOW ................................................. 1289

64 The Major Salivary Glands J.A. YOUNG and D.L COOK. ... ........... .................. .... 1309

65 Function of the Exocrine Pancreas D.L COOK and J.A. YOUNG ..................................... 1327

66 Endocrine Pancreas G. DREWS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1345

67 Physiological Functions of the Liver D. HXUSSINGER .............................................. 1369

68 Zonal Metabolism in the Liver D. HXUSSINGER .............................................. 1393

69 Circulation of the Alimentary Canal J.S. DAVISON....... ...... ... ........ ... ......... ....... ...... 1403

70 The Enteric Nervous System J.S. DAVISON................................................. 1415

71 Energy Metabolism and Nutrition K. JUNGERMANN and C.A. BARTH. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1425

72 Internal Control of Food Intake: Hunger and Satiety H.S. KOOPMANS ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1459

Functions of the Kidney, Fluid- and Electrolyte-Balance

73 Introduction to Renal Function, Renal Blood Flow and the Formation of Filtrate R. GREGER .................................................. 1469

74 Principles of Renal Transport; Concentration and Dilution of Urine R. GREGER .................................................. 1489

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75 Renal Handling of the Individual Solutes of Glomerular Filtrate R. GREGER .................................................. 1517

76 The Body Compartments and Dynamics of Water and Electrolytes F. LANG. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1545

77 Na+ Cl- and Water Metabolism F. LANG. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1557

78 Acid-Base Metabolism F. LANG. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1571

79 K+ Metabolism F. LANG. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1585

80 Caz+, Mgz+ and Phosphate Metabolism F. LANG. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1595

81 Regulation of the Lower Urinary Tract W. JANIG . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1611

82 Central Control of Water and Salt Metabolism R. GREGER .................................................. 1625

IV Acute Vital Functions; Heart; Circulation and Respiration

K Blood and Immune Defense

83 Blood, Plasma Proteins, Coagulation, Fibrinolysis, and Thrombocyte Function C. BAUER and W. WUILLEMIN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1651

84 Hematopoiesis and the Red Blood Cell M.J. KOURY and C. BAUER ..................................... 1679

85 Nonerythroid and Immune Competent Cells of the Blood J. LINDEN MANN and C. BAUER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1695

86 Neuroendocrine Modulation of the Immune System F. CHIAPPELLI, C. FRANCESCHI, E. OTTAVIANI, G.F. SOLOMON, and A.N. TAYLOR. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1707

87 Psychoneuroimmunology M. JERRY. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1731

L Heart and Circulation

88 Hemorheology H. SCHMID-SCHONBEIN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1747

89 The Cardiac Function Cycle H. ANTONI .................................................. 1793

90 Functional Properties of the Heart H. ANTONI .................................................. 1801

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91 Electrophysiology of the Heart at the Single Cell Level and Cardiac Rhythmogenesis H. Antoni ................................................... 1825

92 Electrocardiography H. ANTONI .................................................. 1843

93 Calcium-Mediated Control of Cardiac Contractility at the Cellular Level G.A. LANGER ................................................ 1857

94 Peripheral Circulation: Fundamental Concepts, Comparative Aspects of Control in Specific Sections and Lymph Flow J. HOLTZ .................................................... 1865

95 Hemodynamics in Regional Circulatory Beds and Local Vascular Reactivity J. HOLTZ.................................................... 1917

96 Vascular Smooth Muscle G. SIEGEL ................................................... 1941

97 Microcirculation and Capillary Exchange E.M. RENKIN and C. CRONE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1965

98 Biology of the Vascular Wall and Its Interaction with Migratory and Blood Cells B. NILIUS and R. CASTEELS .................................... 1981

99 Control of the Circulation: An Integrated View H.E.D.J. TER KEURS and J.V. TYBERG ............................ 1995

M Respiration

100 Pulmonary Ventilation B.J. WHIPP. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2015

101 Pulmonary Gas Exchange J. PIIPER .................................................... 2037

102 Respiratory Gas Transport and Acid-Base Equilibrium in Blood J. PIIPER .................................................... 2051

103 Oxygen Supply and Energy Metabolism J. PIIPER .................................................... 2063

104 Pulmonary and Bronchial Circulation N.C. STAUB and C.A. DAWSON.................................. 2071

105 Neural Regulation of Respiration: Rhythmogenesis and Afferent Control D.W. RICHTER ....................... . . . . . . . . . . . . . . . . . . . . . . . . 2079

106 A Systems View of Respiratory Regulation M.E. SCHLAFKE and H.P. KOEPCHEN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2097

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107 Central Nervous Integration of Cardiorespiratory Control K.M. SPYER . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2129

108 Coordination of Circulation and Respiration During Exercise S.A. WARD and B.J. WHIPP. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2145

109 Gravitational and Hyper- and Hypobaric Stress S.A. WARD ............... :.................................. 2175

N Regulation of Body Temperature

110 Basic Thermoregulation K.E. COOPER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2199

III Body Temperature Control and Its Disorders K.E. COOPER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2207

112 The Formation of Sweat R. GREGER .................................................. 2219

V The Life Cycle

o Reproduction

113 Hormonal Regulation of Reproductive Organs G.F. WEINBAUER and E. NIESCHLAG . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2231

114 Behavioral and Neurovegetative Components of Reproductive Functions W. JANIG . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2253

115 Fertilization B. DALE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2265

116 Endocrinology of Pregnancy M. BRECKWOLDT, J. NEULEN, and C. KECK........................ 2277

117 Functions of the Placenta U. KARCK and M. BRECKWOLDT 2291

118 Cardiovascular and Respiratory Systems of the Fetus A. JENSEN. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2307

119 Fetal Endocrinology D.A. FISHER ................................................. 2339

120 The Birth Process H.P. ZAHRADNIK 2347

121 Lactation M. BRECKWOLDT, J. NEULEN, and C. KECK . . . . . . . . . . . . . . . . . . . . . . . . 2365

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P Development, Maturation, Aging, and Death

122 Normal Development and Growth M. BRANDIS ................................................. 2373

123 Physiology of Aging: Standards for Age-Related Functional Competence P.S. TIMIRAS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2391

124 Death R. GREGER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2407

VI Appendix

125 Units Used in Physiology and Their Definitions R. GREGER .................................................. 2421

126 Normal Values for Physiological Parameters R. GREGER and M. BLEICH ......... . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2427

Subject Index ............................................. 2451

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List of Contributors

ANTONI, H.

Physiologisches lnstitut der Universitat Freiburg, Hermann-Herder-Str. 7, 79104 Freiburg, Germany

BARTH, C.A.

Deutsches lnstitut fUr Ernahrungsforschung, Arthur-Scheunert-Allee 114-116, 14558 Bergholz-Rehbrucke, Germany

BAUER, C.

Physiologisches lnstitut der Universitat Zurich, Winterthurer Str. 190,8057 Zurich, Switzerland

BLEICH, M.

Physiologisches Institut der Universitat Freiburg, Hermann-Herder-Str. 7, 79104 Freiburg, Germany

BRADSHAW, R .A.

University of California at Irvine, Department of Biological Chemistry, College of Medicine, Irvine, CA 92717, USA

BRANDIS, M. Universitats-Kinderklinik, Abteilung Allgemeine Kinderheilkunde, Mathildenstr. 1, 79106 Freiburg, Germany

BRECK WOLDT, M. Universitats-Frauenklinik, Abteilung Frauenheilkunde und Geburtshilfe II, Hugstetter Str. 55, 79106 Freiburg, Germany

BOHLER, H.

Freie Universitat Berlin, Universitatsklinikum Benjamin Franklin, Institut fUr Klinische Physiologie, Hindenburgdamm 30, 12200 Berlin, Germany

CARPENTER, R.H.S.

University of Cambridge, The Physiological Laboratory, Downing Street, Cambridge CB2 3EG, UK

CASTEELS, R.

Laboratorium voor Fysiologie, Campus Gasthuisberg K.U.L., Herestraat, 3000 Leuven, Belgium

CHIAPPELLl, F. University of California at Los Angeles, Department of Anatomy and Cell Biology,

UCLA Medical Center, 10833 Le Conte Avenue, Los Angeles, CA 90024-1763, USA

CONIGRAVE, A.D.

University of Sydney, Faculty of Medicine, Edward Ford Building (A2?), Sydney, NSW 2006, Australia

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COOK, D.l.

University of Sydney, Department of Physiology (F13), Sydney, NSW 2006, Australia

COOPER, K.E.

The University of Calgary, Faculty of Medicine, Health Sciences Centre, Department of Medical Physiology, 3330 Hospital Drive N.W., Calgary, Alberta, Canada T2N 4N1

CRONE, e. (t) University of California at Davis, School of Medicine, Department of Human Physiology, Davis, CA 95616, USA

DALE, B. Stazione Zoologic a "Anton Dohrn", Laboratory of Cell and Developmental Biology, Villa Communale 1, 80121, Naples, Italy

DAVISON, J.S.

University of Calgary, Faculty of Medicine, Department of Medical Physiology, 3330 Hospital Drive N.W., Calgary, Alberta, Canada T2N 4N1

DAWSON, C.A.

Medical College of Wisconsin, Department of Clinical Physiology, VA Medical Center, Research Service 151,5000 West National Avenue, Milwaukee, WI 53295-1000, USA

DOWLING, J.E.

Harvard University, The Biological Laboratories, Department of Cellular and Developmental Biology, 16 Divinity Avenue, Cambridge, MA 02138, USA

DREWS, G. Universitat Tiibingen, Pharmazeutisches Institut, Auf der Morgenstelle 8, 72076 Tiibingen, Germany

EHRENSTEIN, W.H. Institut fUr Arbeitsphysiologie an der Universitat Dortmund, Abteilung Sinnes­und Neurophysiologie, Ardeystr. 67, 44139 Dortmund, Germany

FAISSNER, M. Universitat Heidelberg, Institut fUr Neurobiologie, 1m Neuenheimer Feld 364, 69120 Heidelberg, Germany

FINKE, R. Freie Universitat Berlin, Universitatsklinikum Benjamin Franklin, Institut fUr Klinische Physiologie, Hindenburgdamm 30, 12200 Berlin, Germany

FISHER, D.A.

Coining Nichols Institute, 33608 Ortega Highway, San Juan, Capistrano, CA 92690, USA

FLOREZ, J.e. Northwestern University, Department of Neurobiology and Physiology, 2153 N Campus Drive, Evanston, IL 60208-3520, USA

FORTE, J.G. University of California at Berkeley, Department of Molecular and Cell Biology, Division of Cell and Developmental Biology, Berkeley, CA 94720, USA

Fox, S.E.

SUNY Health Sciences Center, Department of Physiology, Box 31, 450 Clarkson Avenue, Brooklyn, NY 11203, USA

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FRANCESCHI, C. University of Modena, Faculty of Pathology, Department of Immunology, 41100 Modena, Italy

FROTSCHER, M. Anatomisches Institut der Universitat Freiburg, Abteilung Anatomie I, Albertstr. 17, 79104 Freiburg, Germany

GREGER, R. Physiologisches Institut der Universitat Freiburg, Hermann-Herder-Str. 7, 79104 Freiburg, Germany

GUMMER, A.W. U niversitats-Hals-N asen-Ohren-Klinik, Horforschungslaboratorien, Silcherstr. 5, 72076 Tiibingen, Germany

HARPER, R.M. UCLA School of Medicine, Department of Anatomy, Los Angeles, CA 90024, USA

HAUSSINGER, D.

Klinik flir Gastroenterologie und Infektiologie, Heinrich -Heine-U niversitat, Moorenstr. 5, 40225 Diisseldorf, Germany

HENATSCH, H.D.

Georg-August-Universitat Gottingen, Zentrum Physiologie und Pathophysiologie, Abteilung Neuro- und Sinnesphysiologie, Humboldtallee 23, 37073 Gottingen, Germany

HERLITZE, S. HNO-Universitatsklinik Tiibingen, Sektion Sensorische Biophysik, Rontgenweg 11, 72076 Tiibingen, Germany

HIERHOLZER, K. Freie Universitat Berlin, Universitatsklinikum Benjamin Franklin, Institut flir Klinische Physiologie, Hindenburgdamm 30, 12200 Berlin, Germany

HINSSEN, H.

Universitat Bielefeld, Fakultat Biologie, Biochemische Zellbiologie, 33501 Bielefeld, Germany

HOLTZ, J.

Martin-Luther-Universitat Halle, Institut fiir Pathophysiologie, Magdeburger Str. 6, 06112 Halle, Germany

JACKSON, M.B. University of Wisconsin School of Medicine, Department of Physiology, 1300 University Avenue, Madison, WI 53706, USA

Jj\NIG, W. Universitat Kiel, Physiologisches Institut, Olshausenstr. 40, 24118 Kiel, Germany

JENSEN, A. Universitats-Frauenklinik, Knappschaftskrankenhaus, In der Schornau 23-25, 44892 Bochum, Germany

JERRY, M. Tom Baker Cancer Center, 1331, 29th Street N.W., Calgary, Alberta, Canada T2N 4N2

JOCKUSCH, B.M. Technische Universitat Braunschweig Carolo Wilhelmina, Zoologisches Institut, Zellbiologie, Spielmannstr. 7, 38106 Braunschweig, Germany

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JOCKUSCH, H. Universitat Bielefeld, Entwicklungsbiologie, W7, 33501 Bielefeld, Germany

JUNGE, D. UCLA School of Medicine, Department of Physiology, 10833 Le Conte Avenue, Los Angeles, CA 90024-1751, USA

JUNGERMANN, K. Universitat G6ttingen, Biochemisches Institut, Humboldtallee 23, 37073 G6ttingen, Germany

KARCK, U. Klinikum der Universitat Freiburg, Universitats-Frauenklinik, Abteilung Frauenheilkunde und Geburtshilfe II, Hugstetter Str. 55, 79106 Freiburg, Germany

KECK, C. Klinikum der Universitat Freiburg, Universitats-Frauenklinik, Abteilung Frauenheilkunde und Geburtshilfe II, Hugstetter Str. 55, 79106 Freiburg, Germany

KELLOW, J.E. University of Sydney, Department of Medicine, The Royal North Shore Hospital, Pacific Highway, St. Leonards, NSW 2065, Australia

KETTENMANN, H. Max-De1bruck-Zentrum fur Molekulare Medizin, Zellulare Neurowissenschaften, Robert-R6ssle-Str. 10, 13122 Berlin, Germany

KOEPCHEN, H.P. (t) Freie Universitat Berlin, Institut flir Physiologie, Arnimallee 22, 14195 Berlin, Germany

KorZUMI, K. SUNY Downstate Medical Center, Department of Physiology, 450 Clarkson Avenue, Box 31, Brooklyn, NY 11203-2098, USA

KOOPMANS, H.S. The University of Calgary, Department of Medical Physiology, Gastrointestinal Research Group, 3330 Hospital Drive N.W., Calgary, Alberta, Canada T2N 4Nl

KOURY, M.J.

Vanderbilt University, Department of Medicine, Hematology Division, Nashville, TN 37232-2287, USA

KUSCHINSKY, W. 1. Physiologisches Institut der Universitat Heidelberg, 1m Neuenheimer Feld 326, 69120 Heidelberg, Germany

LANG, F. Physiologisches Institut der Universitat Tubingen, Gmelinstr. 5, 72076 Tubingen, Germany

LANGER, G.A. UCLA School of Medicine, Cardiovascular Research Laboratories, Department of Physiology, Macdonald Research Laboratory Building, 675 Circle Drive South, Los Angeles, CA 90095, USA

LINDENMANN, J.

Institut fur Immunologie und Virologie, Gloriastr. 30, 8028 Zurich, Switzerland

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LOGAN, e.G. The University of Iowa College of Medicine, Department of Neurology,

200 Hawkins Dr., Iowa City, IA 52242-1053, USA

LOPEs DA SILVA, F.H. University of Amsterdam, Faculty of Biology, Graduate School for the Neurosciences, Institute of Neurobiology, Kruislaan 320, 1098 SM Amsterdam, The Netherlands

McEwEN, B.S. Rockefeller University, Laboratory of Neuroendocrinology, 1230 York Avenue, New York, NY 10021, USA

MOMMAERTS, W.F.H.M. (t) UCLA School of Medicine, Center for the Health Sciences, Department of Physiology, 10833 LeConte Avenue, Los Angeles, CA 90024-1751, USA

NEULEN, J. Universitats-Frauenklinik, Abteilung Frauenheilkunde und Geburtshilfe II, Hugstetter Str. 55, 79106 Freiburg, Germany

NIESCHLAG, E. Westfalische Wilhelms-Universitat, Klinische Forschergruppe fUr Reproduktionsmedizin der Max-Planck-Gesellschaft, Dogmagkstr. 11, 48149 Munster, Germany

NILIUS, B. Laboratorium voor Fysiologie, Campus Gasthuisberg K.U.L., Herestraat, 3000 Leuven, Belgium

OTTAVIANI, E. University of Modena, Department of Animal Biology, 41100 Modena, Italy

PIIPER, J. Max-Planck-Institut fUr experimentelle Medizin, Hermann-Rein-Str. 3, 37075 G6ttingen, Germany

PITTMAN, Q.J. University of Calgary, Health Sciences Center, Department of Medical Physiology, 3330 Hospital Drive N.W., Calgary, Alberta, Canada T2N 4Nl

PLINKERT, P. Universitats-Hals-Nasen-Ohren-Klinik, Silcherstr. 5, 72076 Tiibingen, Germany

RENKIN, E.M. University of California at Davis, School of Medicine, Department of Human Physiology, Davis, CA 95616, USA

RICHTER, D.W. Zentrum Physiologie und Pathophysiologie, Abteilung Neuro- und Sinnesphysiologie, Humboldtallee 23, 37073 G6ttingen, Germany

RUEGG, J.e. II. Physiologisches Institut der Universitat Heidelberg, 1m Neuenheimer Feld 326, 69120 Heidelberg, Germany

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RUPPERSBERG, J.P. HNO-Universitatsklinik Ttibingen, Sektion Sensorische Biophysik, Rontgenweg 11, 72076 Ttibingen, Germany

SCHILD, D. Universitat GoUingen, Zentrum Physiologie und Pathophysiologie, Abteilung Neuro- und Sinnesphysiologie, Humboldtallee 23, 37073 GoUingen, Germany

SCHLAFKE, M.E. Ruhr-Universitat Bochum, Abteilung flir Angewandte Physiologie, Universitatsstr. 150,44780 Bochum, Germany

SCHLEUSENER, H. Freie Universitat Berlin, Universitiitsklinikum Benjamin Franklin, Institut ftir Klinische Physiologie, Hindenburgdamm 30, 12200 Berlin, Germany

SCHMID-SCHONBEIN, H. Rheinisch-Westfalische Technische Hochschule Aachen, Medizinische Fakultat, Institut flir Physiologie, Pauwelsstra~e, 52074 Aachen, Germany

SHARKEY, K.A. University of Calgary, Health Sciences Center, Department of Medical Physiology, 3330 Hospital Drive N.W., Calgary, Alberta, Canada T2N 4N1

SHEPHERD, G.M. Yale University School of Medicine, Section of Neurobiology, 333 Cedar Street, PMB-236, New Haven, CT 06510-8001, USA

SIEGEL, G. Institut flir Physiologie der Freien Universitat Berlin, Fachbereich Natur und sozialwissenschaftliche Grundlagenmedizin und medizinische Okologie, Arnimallee 22, 14195 Berlin, Germany

SIEGEL, J.M. University of California at Los Angeles, School of Medicine, Department of Psychiatry, Neurobiology Research 151A3, Sepulveda V AMC, North Hills, CA 91343, USA

SOKOLOFF, L. National Institute of Mental Health, Laboratory of Cerebral Metabolism, Building 36, Bethesda, MD 20892, USA

SOLOMON, G.F. University of California at Los Angeles, Department of Psychiatry and Biobehavioral Sciences, Los Angeles, CA 90024-1759, USA

SPILLMANN, L. Universitat Freiburg, Institut flir Biophysik und Strahlenbiologie, Hansastr. 9, 79104 Freiburg, Germany

SPYER, K.M. Royal Free Hospital, School of Medicine, Department of Physiology, Rowland Hill Street, London NW3 2PF, UK

STAUB, N.C. University of California, Cardiovascular Research Institute, Department of Physiology, Box 0130, San Francisco, CA 94143, USA

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SUNDBERG, ). Kung Tekniska H6gskolan, Department of Speech Communication and Music Acoustic, Drotning Kristinas vag 31, Box 700 14, 10044 Stockholm, Sweden

TAKAHASHI, J.S. Northwestern University, Department of Neurobiology and Physiology, 2153 N Campus Drive, Evanston, IL 60208-3520, USA

TAYLOR, A.N. University of California at Los Angeles, Department of Neurobiology, Los Angeles, CA 90024-1763, USA

TER KEURS, H.E.D.).

The University of Calgary, Department of Medicine, 3330 Hospital Drive N.W., Calgary, Alberta, Canada T2N 4Nl

THOMPSON, R.F. University of Southern California, Program for Neural, Informational and Behavioral Sciences, HDCO Neurosciences Building, HNB 122, Los Angeles, CA 90089-2520, USA

TIMIRAS, P.S.

University of California at Berkeley, Department of Molecular and Cell Biology, Division of Cell and Developmental Biology, 221 Life Sciences Addition, Berkeley, CA 94720, USA

TROTTER, ).

Universitat Heidelberg, Institute flir Neurobiologie, 1m Neuenheimer Feld 364, 69120 Heidelberg, Germany

TYBERG, J.V. The University of Calgary, Department of Medicine, 3330 Hospital Drive N.W., Calgary, Alberta, Canada T2N 4Nl

WARD, S.A.

South Bank University, Division of Human and Exercise Science, School of Applied Science, 103 Borough Rd., London SEI OAA, UK

WEINBAUER, G.F. Westfalische Wilhelms-Universitat, Klinische Forschergruppe fiir Reproduktionsmedizin der Max-Planck-Gesellschaft, Dogmagkstr. 11, 48149 Munster, Germany

WHIPP, B.). St. George's Hospital Medical School, Department of Physiology, Cranmer Terrace (Tooting), London SW17 ORE, UK

WILLIS, JR., W.D. The University of Texas Medical Branch at Galveston, School of Medicine, Department of Anatomy and Neurosciences, Marine Biomedical Institute, 200 University Boulevard, Galveston, TX 77550-2772, USA

WINDHORST, U.

University of Calgary, Faculty of Medicine, Departments of Clinical Neurosciences and Medical Physiology, 3330 Hospital Drive N.W., Calgary, Alberta, Canada T2N 4Nl

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WUILLEMIN, W.

Hamatologisches Zentrallabor, Universitatsspital Bern, Inselspital, 3010 Bern, Switzerland

YAMAMOTO, W.S.

The George Washington University School of Medicine, Department of Computer Medicine, 2300 K St. N.W., Washington, DC 20037, USA

YOUNG, J.A. University of Sydney, Faculty of Medicine, Edward Ford Building (A27), Sydney, NSW 2006, Australia

ZAHRADNIK, H.P. Universitats-Frauenklinik II, Hugstetter Str. 55, 79106 Freiburg, Germany

ZENNER, H.-P. U niversitats-Hals-Nasen -Ohren-Klinik, Horforschungslaboratorien, Silcherstr. 5, 72076 Tiibingen, Germany

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