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At a Glance

1 Fundamentals and Cell Physiology 2

2 Nerve and Muscle, Physical Work 42

3 Autonomic Nervous System (ANS) 78

4 Blood 88

5 Respiration 106

6 Acid–Base Homeostasis 138

7 Kidneys, Salt, and Water Balance 148

8 Cardiovascular System 186

9 Thermal Balance and Thermoregulation 222

10 Nutrition and Digestion 226

11 Hormones and Reproduction 266

12 Central Nervous System and Senses 310

13 Appendix 372

Further Reading 391

Index 394

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

II

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

Color Atlas

of Physiology

5th edition, completely revised

and expanded

Agamemnon Despopoulos, M.D.

Professor

Formerly: Ciba Geigy

Basel

Stefan Silbernagl, M.D.

Professor

Head of Department

Institute of Physiology

University of Wuerzburg

Wuerzburg, Germany

186 color plates by

Ruediger Gay and

Astried Rothenburger

Thieme

Stuttgart · New York

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

IV

Library of Congress Cataloging-in-Publication

Data

is available from the publisher

This book is an authorized translation of the

5th German edition published and copy￾righted 2001 by Georg Thieme Verlag, Stutt￾gart, Germany.

Title of the German edition:

Taschenatlas der Physiologie

Translated by Suzyon O’Neal Wandrey, Berlin,

Germany

Illustrated by Atelier Gay + Rothenburger, Ster￾nenfels, Germany

! 1981, 2003 Georg Thieme Verlag

Rüdigerstraße 14, D-70469 Stuttgart, Germany

http://www.thieme.de

Thieme New York, 333 Seventh Avenue,

New York, N.Y. 10001, U.S.A.

http://www.thieme.com

Cover design: Cyclus, Stuttgart

Typesetting by: Druckhaus Götz GmbH,

Ludwigsburg, Germany

Printed in Germany by: Appl Druck

GmbH & Co. KG, Wemding, Germany

ISBN 3-13-545005-8 (GTV)

ISBN 1-58890-061-4 (TNY) 1 2 3 4 5

1st German edition 1979

2nd German edition 1983

3rd German edition 1988

4th German edition 1991

5th German edition 2001

1st English edition 1981

2nd English edition 1984

3rd English edition 1986

4th English edition 1991

1st Dutch edition 1981

2nd Dutch edition 2001

1st Italian edition 1981

2nd Italian edition 2001

1st Japanese edition 1982

2nd Japanese edition 1992

1st Spanish edition 1982

2nd Spanish edition 1985

3rd Spanish edition 1994

4th Spanish edition 2001

Important Note: Medicine is an ever-changing

science undergoing continual development.

Research and clinical experience are continu￾ally expanding our knowledge, in particular

our knowledge of proper treatment and drug

therapy. Insofar as this book mentions any do￾sage or application, readers may rest assured

that the authors, editors, and publishers have

made every effort to ensure that such refe￾rences are in accordance with the state of

knowledge at the time of production of the

book.

Nevertheless, this does not involve, imply,

or express any guarantee or responsibility on

the part of the publishers in respect to any do￾sage instructions and forms of applications

stated in the book. Every user is requested to

examine carefully the manufacturers’ leaflets

accompanying each drug and to check, if

necessary in consultation with a physician or

specialist, whether the dosage schedules men￾tioned therein or the contraindications stated

by the manufacturers differ from the state￾ments made in the present book. Such exami￾nation is particularly important with drugs

that are either rarely used or have been newly

released on the market. Every dosage schedule

or every form of application used is entirely at

the user’s own risk and responsibility. The au￾thors and publishers request every user to re￾port to the publishers any discrepancies or

inaccuracies noticed.

Some of the product names, patents, and

registered designs referred to in this book are

in fact registered trademarks or proprietary

names even though specific reference to this

fact is not always made in the text. Therefore,

the appearance of a name without designation

as proprietary is not to be construed as a repre￾sentation by the publisher that it is in the

public domain.

This book, including all parts thereof, is le￾gally protected by copyright. Any use, exploita￾tion, or commercialization outside the narrow

limits set by copyright legislation, without the

publisher’s consent, is illegal and liable to pro￾secution. This applies in particular to photostat

reproduction, copying, mimeographing or

duplication of any kind, translating, prepara￾tion of microfilms, and electronic data pro￾cessing and storage.

1st Czech edition 1984

2nd Czech edition 1994

1st French edition 1985

2nd French edition 1992

3rd French edition 2001

1st Turkish edition 1986

2nd Turkish edition 1997

1st Greek edition 1989

1st Chinese edition 1991

1st Polish edition 1994

1st Hungarian edition 1994

2nd Hungarian edition 1996

1st Indonesion edition 2000

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

V

Preface to the Fifth Edition

The base of knowledge in many sectors of phy￾siology has grown considerably in magnitude

and in depth since the last edition of this book

was published. Many advances, especially the

rapid progress in sequencing the human ge￾nome and its gene products, have brought

completely new insight into cell function and

communication. This made it necessary to edit

and, in some cases, enlarge many parts of the

book, especially the chapter on the fundamen￾tals of cell physiology and the sections on

neurotransmission, mechanisms of intracellu￾lar signal transmission, immune defense, and

the processing of sensory stimuli. A list of phy￾siological reference values and important for￾mulas were added to the appendix for quick

reference. The extensive index now also serves

as a key to abbreviations used in the text.

Some of the comments explaining the con￾nections between pathophysiological princi￾ples and clinical dysfunctions had to be slight￾ly truncated and set in smaller print. However,

this base of knowledge has also grown consi￾derably for the reasons mentioned above. To

make allowances for this, a similarly designed

book, the Color Atlas of Pathophysiology

(S. Silbernagl and F. Lang, Thieme), has now

been introduced to supplement the well￾established Color Atlas of Physiology.

I am very grateful for the many helpful com￾ments from attentive readers (including my

son Jakob) and for the welcome feedback from

my peers, especially Prof. H. Antoni, Freiburg,

Prof. C. von Campenhausen, Mainz, Dr. M. Fi￾scher, Mainz, Prof. K.H. Plattig, Erlangen, and

Dr. C. Walther, Marburg, and from my collea￾gues and staff at the Institute in Würzburg. It

was again a great pleasure to work with Rüdi￾ger Gay and Astried Rothenburger, to whom I

am deeply indebted for revising practically all

the illustrations in the book and for designing a

number of new color plates. Their extraordina￾ry enthusiasm and professionalism played a

decisive role in the materialization of this new

edition. To them I extend my sincere thanks. I

would also like to thank Suzyon O’Neal Wan￾drey for her outstanding translation. I greatly

appreciate her capable and careful work. I am

also indebted to the publishing staff, especially

Marianne Mauch, an extremely competent and

motivated editor, and Gert Krüger for invalu￾able production assistance. I would also like to

thank Katharina Völker for her ever observant

and conscientious assistance in preparing the

index.

I hope that the 5th Edition of the Color Atlas

of Physiology will prove to be a valuable tool for

helping students better understand physiolog￾ical correlates, and that it will be a valuable re￾ference for practicing physicians and scien￾tists, to help them recall previously learned in￾formation and gain new insights in physiology.

Würzburg, December 2002

Stefan Silbernagl*

* e-mail: [email protected]

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

VI

A book of this nature is inevitably deriva￾tive, but many of the representations are new

and, we hope, innovative. A number of people

have contributed directly and indirectly to the

completion of this volume, but none more

than Sarah Jones, who gave much more than

editorial assistance. Acknowledgement of

helpful criticism and advice is due also to Drs.

R. Greger, A. Ratner, J. Weiss, and S. Wood, and

Prof. H. Seller. We are grateful to Joy Wieser for

her help in checking the proofs. Wolf-Rüdiger

and Barbara Gay are especially recognized, not

only for their art work, but for their conceptual

contributions as well. The publishers, Georg

Thieme Verlag and Deutscher Taschenbuch

Verlag, contributed valuable assistance based

on extensive experience; an author could wish

for no better relationship. Finally, special

recognition to Dr. Walter Kumpmann for in￾spiring the project and for his unquestioning

confidence in the authors.

Basel and Innsbruck, Summer 1979

Agamemnon Despopoulos

Stefan Silbernagl

Preface to the First Edition

In the modern world, visual pathways have

outdistanced other avenues for informational

input. This book takes advantage of the econo￾my of visual representation to indicate the si￾multaneity and multiplicity of physiological

phenomena. Although some subjects lend

themselves more readily than others to this

treatment, inclusive rather than selective

coverage of the key elements of physiology has

been attempted.

Clearly, this book of little more than 300

pages, only half of which are textual, cannot be

considered as a primary source for the serious

student of physiology. Nevertheless, it does

contain most of the basic principles and facts

taught in a medical school introductory

course. Each unit of text and illustration can

serve initially as an overview for introduction

to the subject and subsequently as a concise

review of the material. The contents are as cur￾rent as the publishing art permits and include

both classical information for the beginning

students as well as recent details and trends

for the advanced student.

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

VII

From the Preface to the Third Edition

The first German edition of this book was al￾ready in press when, on November 2nd, 1979,

Agamennon Despopoulos and his wife, Sarah

Jones-Despopoulos put to sea from Bizerta, Tu￾nisia. Their intention was to cross the Atlantic

in their sailing boat. This was the last that was

ever heard of them and we have had to aban￾don all hope of seeing them again.

Without the creative enthusiasm of Aga￾mennon Despopoulos, it is doubtful whether

this book would have been possible; without

his personal support it has not been easy to

continue with the project. Whilst keeping in

mind our original aims, I have completely re￾vised the book, incorporating the latest advan￾ces in the field of physiology as well as the wel￾come suggestions provided by readers of the

earlier edition,to whom I extend my thanks for

their active interest.

Würzburg, Fall 1985

Stefan Silbernagl

Dr. Agamemnon Despopoulos

Born 1924 in New York; Professor of Physiology at the

University of New Mexico. Albuquerque, USA, until 1971;

thereafter scientific adviser to CIBA-GEIGY, Basel.

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

VIII

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

IX

Table of Contents

1

Fundamentals and Cell Physiology 2

The Body: an Open System with an Internal Environment · · · 2

Control and Regulation · · · 4

The Cell · · · 8

Transport In, Through, and Between Cells · · · 16

Passive Transport by Means of Diffusion · · · 20

Osmosis, Filtration, and Convection · · · 24

Active Transport · · · 26

Cell Migration · · · 30

Electrical Membrane Potentials and Ion Channels · · · 32

Role of Ca2+ in Cell Regulation · · · 36

Energy Production and Metabolism · · · 38

2

Nerve and Muscle, Physical Work 42

Neuron Structure and Function · · · 42

Resting Membrane Potential · · · 44

Action Potential · · · 46

Propagation of Action Potentials in Nerve Fiber · · · 48

Artificial Stimulation of Nerve Cells · · · 50

Synaptic Transmission · · · 50

Motor End-plate · · · 56

Motility and Muscle Types · · · 58

Motor Unit of Skeletal Muscle · · · 58

Contractile Apparatus of Striated Muscle · · · 60

Contraction of Striated Muscle · · · 62

Mechanical Features of Skeletal Muscle · · · 66

Smooth Muscle · · · 70

Energy Supply for Muscle Contraction · · · 72

Physical Work · · · 74

Physical Fitness and Training · · · 76

3

Autonomic Nervous System (ANS) 78

Organization of the Autonomic Nervous System · · · 78

Acetylcholine and Cholinergic Transmission · · · 82

Catecholamine, Adrenergic Transmission and Adrenoceptors · · · 84

Adrenal Medulla · · · 86

Non-cholinergic, Non-adrenergic Transmitters · · · 86

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

X

4

Blood 88

Composition and Function of Blood · · · 88

Iron Metabolism and Erythropoiesis · · · 90

Flow Properties of Blood · · · 92

Plasma, Ion Distribution · · · 92

Immune System · · · 94

Hypersensitivity Reactions (Allergies) · · · 100

Blood Groups · · · 100

Hemostasis · · · 102

Fibrinolysis and Thromboprotection · · · 104

5

Respiration 106

Lung Function, Respiration · · · 106

Mechanics of Breathing · · · 108

Purification of Respiratory Air · · · 110

Artificial Respiration · · · 110

Pneumothorax · · · 110

Lung Volumes and their Measurement · · · 112

Dead Space, Residual Volume, and Airway Resistance · · · 114

Lung–Chest Pressure—Volume Curve, Respiratory Work · · · 116

Surface Tension, Surfactant · · · 118

Dynamic Lung Function Tests · · · 118

Pulmonary Gas Exchange · · · 120

Pulmonary Blood Flow, Ventilation–Perfusion Ratio · · · 122

CO2 Transport in Blood · · · 124

CO2 Binding in Blood · · · 126

CO2 in Cerebrospinal Fluid · · · 126

Binding and Transport of O2 in Blood · · · 128

Internal (Tissue) Respiration, Hypoxia · · · 130

Respiratory Control and Stimulation · · · 132

Effects of Diving on Respiration · · · 134

Effects of High Altitude on Respiration · · · 136

Oxygen Toxicity · · · 136

6

Acid–Base Homeostasis 138

pH, pH Buffers, Acid–Base Balance · · · 138

Bicarbonate/Carbon Dioxide Buffer · · · 140

Acidosis and Alkalosis · · · 142

Assessment of Acid–Base Status · · · 146

7

Kidneys, Salt, and Water Balance 148

Kidney Structure and Function · · · 148

Renal Circulation · · · 150

Glomerular Filtration and Clearance · · · 152

Transport Processes at the Nephron · · · 154

Reabsorption of Organic Substances · · · 158

Despopoulos, Color Atlas of Physiology © 2003 Thieme

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XI

Excretion of Organic Substances · · · 160

Reabsorption of Na+ and Cl– · · · 162

Reabsorption of Water, Formation of Concentrated Urine · · · 164

Body Fluid Homeostasis · · · 168

Salt and Water Regulation · · · 170

Diuresis and Diuretics · · · 172

Disturbances of Salt and Water Homeostasis · · · 172

The Kidney and Acid–Base Balance · · · 174

Reabsorption and Excretion of Phosphate, Ca2+ and Mg2+ · · · 178

Potassium Balance · · · 180

Tubuloglomerular Feedback, Renin–Angiotensin System · · · 184

8

Cardiovascular System 186

Overview · · · 186

Blood Vessels and Blood Flow · · · 188

Cardiac Cycle · · · 190

Cardiac Impulse Generation and Conduction · · · 192

Electrocardiogram (ECG) · · · 196

Excitation in Electrolyte Disturbances · · · 198

Cardiac Arrhythmias · · · 200

Ventricular Pressure–Volume Relationships · · · 202

Cardiac Work and Cardiac Power · · · 202

Regulation of Stroke Volume · · · 204

Venous Return · · · 204

Arterial Blood Pressure · · · 206

Endothelial Exchange Processes · · · 208

Myocardial Oxygen Supply · · · 210

Regulation of the Circulation · · · 212

Circulatory Shock · · · 218

Fetal and Neonatal Circulation · · · 220

9

Thermal Balance and Thermoregulation 222

Thermal Balance · · · 222

Thermoregulation · · · 224

10

Nutrition and Digestion 226

Nutrition · · · 226

Energy Metabolism and Calorimetry · · · 228

Energy Homeostasis and Body Weight · · · 230

Gastrointestinal (GI) Tract: Overview, Immune Defense and Blood Flow · · · 232

Neural and Hormonal Integration · · · 234

Saliva · · · 236

Deglutition · · · 238

Vomiting · · · 238

Stomach Structure and Motility · · · 240

Gastric Juice · · · 242

Small Intestinal Function · · · 244

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

XII

Pancreas · · · 246

Bile · · · 248

Excretory Liver Function—Bilirubin · · · 250

Lipid Digestion · · · 252

Lipid Distribution and Storage · · · 254

Digestion and Absorption of Carbohydrates and Protein · · · 258

Vitamin Absorption · · · 260

Water and Mineral Absorption · · · 262

Large Intestine, Defecation, Feces · · · 264

11

Hormones and Reproduction 266

Integrative Systems of the Body · · · 266

Hormones · · · 268

Humoral Signals: Control and Effects · · · 272

Cellular Transmission of Signals from Extracellular Messengers · · · 274

Hypothalamic–Pituitary System · · · 280

Carbohydrate Metabolism and Pancreatic Hormones · · · 282

Thyroid Hormones · · · 286

Calcium and Phosphate Metabolism · · · 290

Biosynthesis of Steroid Hormones · · · 294

Adrenal Cortex and Glucocorticoid Synthesis · · · 296

Oogenesis and the Menstrual Cycle · · · 298

Hormonal Control of the Menstrual Cycle · · · 300

Estrogens · · · 302

Progesterone · · · 302

Prolactin and Oxytocin · · · 303

Hormonal Control of Pregnancy and Birth · · · 304

Androgens and Testicular Function · · · 306

Sexual Response, Intercourse and Fertilization · · · 308

12

Central Nervous System and Senses 310

Central Nervous System · · · 310

Cerebrospinal Fluid · · · 310

Stimulus Reception and Processing · · · 312

Sensory Functions of the Skin · · · 314

Proprioception, Stretch Reflex · · · 316

Nociception and Pain · · · 318

Polysynaptic Reflexes · · · 320

Synaptic Inhibition · · · 320

Central Conduction of Sensory Input · · · 322

Motor System · · · 324

Hypothalamus, Limbic System · · · 330

Cerebral Cortex, Electroencephalogram (EEG) · · · 332

Sleep–Wake Cycle, Circadian Rhythms · · · 334

Consciousness, Memory, Language · · · 336

Glia · · · 338

Sense of Taste · · · 338

Sense of Smell · · · 340

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

XIII

Sense of Balance · · · 342

Eye Structure, Tear Fluid, Aqueous Humor · · · 344

Optical Apparatus of the Eye · · · 346

Visual Acuity, Photosensors · · · 348

Adaptation of the Eye to Different Light Intensities · · · 352

Retinal Processing of Visual Stimuli · · · 354

Color Vision · · · 356

Visual Field, Visual Pathway, Central Processing of Visual Stimuli · · · 358

Eye Movements, Stereoscopic Vision, Depth Perception · · · 360

Physical Principles of Sound—Sound Stimulus and Perception · · · 362

Conduction of Sound, Sound Sensors · · · 364

Central Processing of Acoustic Information · · · 368

Voice and Speech · · · 370

13 Appendix 372

Dimensions and Units · · · 372

Powers and Logarithms · · · 380

Graphic Representation of Data · · · 381

The Greek Alphabet · · · 384

Reference Values in Physiology · · · 384

Important Equations in Physiology · · · 388

Further Reading 391

Index 394

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

2

!

1 Fundamentals and Cell Physiology

“. . . If we break up a living organism by isolating its different parts, it is only for the sake of ease in

analysis and by no means in order to conceive them separately. Indeed, when we wish to ascribe to a

physiological quality its value and true significance, we must always refer itto the whole and draw our

final conclusions only in relation to its effects on the whole.”

Claude Bernard (1865)

The Body: an Open System with an

Internal Environment

The existence of unicellular organisms is the

epitome of life in its simplest form. Even

simple protists must meettwo basic but essen￾tially conflicting demands in order to survive.

A unicellular organism must, on the one hand,

isolate itself from the seeming disorder of its

inanimate surroundings, yet, as an “open sys￾tem” (! p. 40), it is dependent on its environ￾ment for the exchange of heat, oxygen,

nutrients, waste materials, and information.

“Isolation” is mainly ensured by the cell

membrane, the hydrophobic properties of

which prevent the potentially fatal mixing of

hydrophilic components in watery solutions

inside and outside the cell. Protein molecules

within the cell membrane ensure the perme￾ability of the membrane barrier. They may

exist in the form of pores (channels) or as more

complex transport proteins known as carriers

(! p. 26 ff.). Both types are selective for cer￾tain substances, and their activity is usually

regulated. The cell membrane is relatively well

permeable to hydrophobic molecules such as

gases. This is useful for the exchange of O2 and

CO2 and for the uptake of lipophilic signal sub￾stances, yet exposes the cellto poisonous gases

such as carbon monoxide (CO) and lipophilic

noxae such as organic solvents. The cell mem￾brane also contains other proteins—namely,

receptors and enzymes. Receptors receive sig￾nals from the external environment and con￾vey the information to the interior of the cell

(signal transduction), and enzymes enable the

cell to metabolize extracellular substrates.

Let us imagine the primordial sea as the ex￾ternal environment of the unicellular or￾ganism (! A). This milieu remains more or less

constant, although the organism absorbs

nutrients from it and excretes waste into it. In

spite of its simple structure, the unicellular or￾ganism is capable of eliciting motor responses

to signals from the environment. This is

achieved by moving its pseudopodia or

flagella, for example, in response to changes in

the food concentration.

The evolution from unicellular organisms to

multicellular organisms, the transition from

specialized cell groups to organs, the emer￾gence of the two sexes, the coexistence of in￾dividuals in social groups, and the transition

from water to land have tremendously in￾creased the efficiency, survival, radius of ac￾tion, and independence of living organisms.

This process required the simultaneous devel￾opment of a complex infrastructure within the

organism. Nonetheless, the individual cells of

the body still need a milieu like that of the

primordial sea for life and survival. Today, the

extracellular fluid is responsible for providing

constant environmental conditions (! B), but

the volume of the fluid is no longer infinite. In

fact, it is even smaller than the intracellular

volume (! p. 168). Because of their metabolic

activity, the cells would quickly deplete the

oxygen and nutrient stores within the fluids

and flood their surroundings with waste prod￾ucts if organs capable of maintaining a stable

internal environment had not developed. This

is achieved through homeostasis, a process by

which physiologic self-regulatory mecha￾nisms (see below) maintain steady states in

the body through coordinated physiological

activity. Specialized organs ensure the con￾tinuous absorption of nutrients, electrolytes

and water and the excretion of waste products

via the urine and feces. The circulating blood

connects the organs to every inch of the body,

and the exchange of materials between the

blood and the intercellular spaces (interstices)

creates a stable environment for the cells. Or￾gans such as the digestive tract and liver ab￾sorb nutrients and make them available by

processing, metabolizing and distributing

Despopoulos, Color Atlas of Physiology © 2003 Thieme

All rights reserved. Usage subject to terms and conditions of license.

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