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 This International Student Edition is for use outside of the U.S.

Sixteenth Edition

HUMAN
Vander’s

PHYSIOLOGY The Mechanisms of Body Function

Eric P. Widmaier | Hershel Raff | Kevin T. Strang

ISTUDY
 SIXTEENTH EDITION VA N D E R ’ S

Human
Physiology
The Mechanisms of Body Function

ERIC P. WIDMAIER
B O S TO N U N I V E R S I T Y

HERSHEL RAFF
M E D I C A L CO L L E G E O F W I S CO N S I N
AU R O R A S T. LU K E ’ S M E D I C A L C E N T E R /
A DVO C AT E AU R O R A R E S E A R C H I N S T I T U T E

KEVIN T. STRANG
U N I V E R S I T Y O F W I S CO N S I N – M A D I S O N

ISTUDY

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 VANDER’S HUMAN PHYSIOLOGY

Published by McGraw Hill LLC, 1325 Avenue of the Americas, New York, NY 10019. Copyright ©2023 by McGraw Hill LLC.
All rights reserved. Printed in the United States of America. No part of this publication may be reproduced or distributed in
any form or by any means, or stored in a database or retrieval system, without the prior written consent of McGraw Hill LLC,
including, but not limited to, in any network or other electronic storage or transmission, or broadcast for distance learning.

Some ancillaries, including electronic and print components, may not be available to customers outside the United States.

This book is printed on acid-free paper.

1 2 3 4 5 6 7 8 9 LWI 27 26 25 24 23 22

ISBN 978-1-265-13181-4
MHID 1-265-13181-3

Cover Image: Rattanasak Khuentana/Shutterstock

All credits appearing on page are considered to be an extension of the copyright page.

The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website does not indicate an
endorsement by the authors or McGraw Hill LLC, and McGraw Hill LLC does not guarantee the accuracy of the information
presented at these sites.

mheducation.com/highered

ISTUDY
 BRIEF CONTENTS
MEET THE AUTHORS IV ■ FROM THE AUTHORS V ■ INDEX OF EXERCISE PHYSIOLOGY XV ■ GUIDED TOUR THROUGH A CHAPTER XVI
■ UPDATES AND ADDITIONS XX ■ ACKNOWLEDGMENTS XXI ■ CONNECT XXII

■ 1 Homeostasis: A Framework ■ 10 Control of Body ■ 16 Regulation of Organic

for Human Physiology 1 Movement 300 Metabolism and Energy
Balance 574
■ 2 Chemical Composition of ■ 11 The Endocrine System 319 ■ 
Control and Integration of
the Body and Its Relation to Carbohydrate, Protein, and Fat
■ 
General Characteristics of
Physiology 21 Hormones and Hormonal Control
Metabolism 575
Systems 320 ■ 
Regulation of Total-Body Energy
■ 3 Cellular Structure, ■ 
The Hypothalamus and
Balance 589

Proteins, and Metabolic Pituitary Gland 332 ■ 

Regulation of Body
Temperature 594
Pathways 45 ■ The Thyroid Gland 338

■ Cell Structure 46
■ 
T he Endocrine Response to
Stress 343
■ 17 Reproduction 604
■ 
Protein Synthesis, Degradation, ■ 
■ 
Endocrine Control of Growth 347 Overview and Gametogenesis,
and Secretion 57 Sex Determination, and Sex
■  Endocrine Control of Ca2+
■  Differentiation; General
Interactions Between Proteins and
Homeostasis 351 Principles of Reproductive
Ligands 66
■ 
Chemical Reactions and
Enzymes 71
■ 12 Cardiovascular
Endocrinology 605
■ 
Male Reproductive
Physiology 361 Physiology 614
■ Metabolic Pathways 78
■ 
General Features of the ■ 
Female Reproductive
■ 4 Movement of Solutes Circulatory System 362 Physiology 624
and Water Across Cell ■ The Heart 371 ■ 
P regnancy, Contraception,
Infertility, and Hormonal Changes
Membranes 95 ■ The Vascular System 389 Through Life 637

■ 5 Cell Signaling in
■ 
Integration of Cardiovascular
Function: Regulation of Systemic ■ 18 The Immune System 659
Physiology 118 Arterial Pressure 409
■ 
Cardiovascular Patterns in Health ■ 19 Medical Physiology:
■ 6 Neuronal Signaling and the
and Disease 417
■ 
Hemostasis: The Prevention of
Integration Using Clinical
Structure of the Nervous Blood Loss 430
Cases 697
System 136
■ Cells of the Nervous System 137
■ 13 Respiratory APPENDIX A A-1
Physiology 445
■ Membrane Potentials 143 APPENDIX B A-42
■ Synapses 158 ■ 14 The Kidneys and APPENDIX C A-46
■ 
Structure of the Nervous Regulation of Water
System 172
and Inorganic Ions 490 GLOSSARY/INDEX GI-1

■ 7 Sensory Physiology 190 ■ 

Basic Principles of Renal
Physiology 491
■ General Principles 191
■ 
Regulation of Ion and Water
■ Specific Sensory Systems 201 Balance 505

■ 8 Consciousness, the Brain, ■ 

Hydrogen Ion Regulation 523
and Behavior 234 ■ 15 The Digestion and
Absorption of Food 534
■ 9 Muscle 257
■ Skeletal Muscle 258
■ Smooth and Cardiac Muscle 286

iii

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 MEET THE AUTHORS
ERIC P. WIDMAIER received his Ph.D. in 1984 in Endocrinology from the University of
California at San Francisco. His postdoctoral training was in molecular endocrinology, neuroscience,
and physiology at the Worcester Foundation for Experimental Biology in Shrewsbury, Massachusetts,
and The Salk Institute in La Jolla, California. His research was focused on the control of body mass and
metabolism in mammals, the mechanisms of hormone action, and molecular mechanisms of intestinal
and hypothalamic adaptation to high-fat diets. He is currently Emeritus Professor of Biology at Boston
University, where he has taught Human Physiology for many years, and he has been recognized with
the Gitner Award for Distinguished Teaching by the College of Arts and Sciences as well as the Metcalf
Prize for Excellence in Teaching by Boston University. He is the author of many scientific and lay
Photo courtesy of: Maria
publications, including books about physiology for the general reader. He has two grown children, Rick
Widmaier
and Carrie; he and his wife Maria divide their time between New Hampshire and Florida.

H ER SH EL R AFF received his Ph.D. in Environmental Physiology from the Johns Hopkins
University in 1981 and did postdoctoral training in Endocrinology at the University of California
at San Francisco. He is now a Professor of Medicine (Endocrinology and Molecular Medicine),
Surgery, and Physiology in the School of Medicine at the Medical College of Wisconsin. He is
Director of the Endocrine Research Laboratory at Aurora St. Luke’s Medical Center/Advocate
Aurora Research Institute. He teaches physiology and pathophysiology to medical, pharmacy,
and graduate students as well as clinical fellows. At the Medical College of Wisconsin, he is
the Endocrinology/Reproduction Course Director for second-year medical students. He was an
inaugural inductee into the Society of Teaching Scholars, elected as a faculty member to Alpha
Omega Alpha (AOA Honor Medical Society), received the Beckman Basic Science Teaching Award
Photo courtesy of: Tonya from the senior MD class five times, and has been one of the MCW’s Outstanding Medical Student
Limberg Teachers in multiple years. He is also an Adjunct Professor of Biomedical Sciences at Marquette
University. Dr. Raff’s basic research focuses on the adaptation to stress. His clinical interest focuses
on pituitary and adrenal diseases, with a special focus on laboratory tests for the diagnosis of
Cushing’s syndrome. He resides outside Milwaukee with his wife Judy and son Jonathan.

K EVI N T. STR ANG received both his Master’s Degree in Zoology (1988) and his Ph.D.
in Physiology (1994) from the University of Wisconsin–Madison, where he is now an emeritus
Distinguished Faculty Associate in the Departments of Neuroscience and Kinesiology. His thesis
research focused on cellular mechanisms of contractility modulation in cardiac muscle. For over
30 years he taught a large undergraduate systems physiology course as well as the first-year medical
physiology course in the UW–Madison School of Medicine and Public Health. He was elected to
UW–Madison’s Teaching Academy and as a Fellow of the Wisconsin Initiative for Science Literacy.
Photo courtesy of: Kevin Strang He has been a frequent guest speaker at colleges and high schools on the physiology of alcohol
consumption. Twice awarded the UW Medical Alumni Association’s Distinguished Teaching
Award for Basic Sciences, he also received the University of Wisconsin System’s Underkofler/
Alliant Energy Excellence in Teaching Award. In 2012 he was featured in The Princeton Review
publication The Best 300 Professors. Interested in teaching technology, Dr. Strang has produced
numerous physiology animations, some of which were adopted for use with Vander’s Human
Physiology. He has two adult children, Jake and Amy, and lives in Madison with his wife Sheryl.

T O O U R FA M I L I E S : M A R I A , C A R O L I N E , A N D R I C H A R D ; J U DY A N D J O N A T H A N ;
S H E RY L , J A K E , A N D A M Y

iv

ISTUDY
 FROM THE AUTHORS
Lifeline to success in physiology

We are pleased to offer an integrated package of textual and

digital material to deliver basic and clinical content, real-
life applications, and educational technologies to students of
physiology. With the sixteenth edition of Vander’s Human
Physiology, all these pieces come together to facilitate learn-
ing and enthusiasm for understanding the mechanisms of
body function.
The cover of this edition reflects several areas of focus
of the book, including homeostasis, exercise, and human
health. These and other areas of interest are elaborated
upon, beginning with Chapter 1, where the key “General
Principles of Physiology,” an underlying theme in the book,
is first introduced. Unifying themes, such as homeostasis,
are explored throughout the book at all levels of system,
organ, tissue, and cellular function. As in previous editions,
these themes are always related to pathophysiology through
the use of compelling clinical case studies in all chapters,
and a final chapter with several cases that integrate material
across the entire book.
We are certain that you will find the sixteenth edition
of this textbook to be the most up-to-date and comprehen-
sive book available for students of physiology. Thank you
and happy reading!

Cover Image: Jacob Lund/Shutterstock

ISTUDY

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 CONTENTS
MEET THE AUTHORS IV ■ FROM THE AUTHORS V ■ INDEX OF EXERCISE PHYSIOLOGY XV ■ GUIDED TOUR THROUGH A CHAPTER XVI
■ UPDATES AND ADDITIONS XX ■ ACKNOWLEDGMENTS XXI ■ CONNECT XXII

Hydrogen Bonds 27

1
Molecular Shape 27
Homeostasis: A Framework Ionic Molecules 27
for Human Physiology 1 2.3 Solutions 28
Water 28
1.1 The Scope of Human Physiology 2 Molecular Solubility 29
Concentration 30
1.2 How Is the Body Organized? 2
Hydrogen Ions and Acidity 30
Muscle Cells and Tissue 3
2.4 Classes of Organic Molecules 31
Neurons and Nervous Tissue 3
Epithelial Cells and Epithelial Tissue 3 Carbohydrates 31
Connective-Tissue Cells and Connective Tissue 4 Lipids 33
Organs and Organ Systems 4 Proteins 35
Nucleic Acids 39
1.3 Body Fluid Compartments 5
Chapter 2 Clinical Case Study 42
1.4 Homeostasis: A Defining Feature of Physiology 7
1.5 General Characteristics of Homeostatic Control ASSESSMENT QUESTIONS 43
Systems 8
Feedback Systems 9

3
Resetting of Set Points 9
Feedforward Regulation 10 Cellular Structure, Proteins, and
1.6 Components of Homeostatic Control Systems 11 Metabolic Pathways 45
Reflexes 11
Local Homeostatic Responses 13
1.7 The Role of Intercellular Chemical Messengers in Cell Structure 46
Homeostasis 13 3.1 Microscopic Observations of Cells 46
1.8 Processes Related to Homeostasis 14 3.2 Membranes 48
Adaptation and Acclimatization 14 Membrane Structure 48
Biological Rhythms 14 Membrane Junctions 50
Balance of Chemical Substances in the Body 15 3.3 Cell Organelles 52
1.9 General Principles of Physiology 16 Nucleus 52
Chapter 1 Clinical Case Study 17 Ribosomes 52
Endoplasmic Reticulum 53
ASSESSMENT QUESTIONS 20
Golgi Apparatus 53
Endosomes 53
Mitochondria 53

2
Chemical Composition of Lysosomes 54
the Body and Its Relation to Peroxisomes 55
Physiology 21 Cytoskeleton 55

Protein Synthesis, Degradation, and Secretion 57

2.1 Atoms 22
3.4 Genetic Code 57
Components of Atoms 22
Atomic Number 23 3.5 Protein Synthesis 58
Atomic Mass 23 Transcription: mRNA Synthesis 59
Ions 24 Translation: Polypeptide Synthesis 60
Atomic Composition of the Body 24 Regulation of Protein Synthesis 62
2.2 Molecules 25 Mutation 64
Covalent Chemical Bonds 25 3.6 Protein Degradation 65
Ionic Bonds 26

vi

ISTUDY
 3.7 Protein Secretion 65 4.4 Endocytosis and Exocytosis 110
Endocytosis 111
Interactions Between Proteins and Ligands 66 Exocytosis 112
4.5 Epithelial Transport 113
3.8 Binding Site Characteristics 66
Chemical Specificity 66 Chapter 4 Clinical Case Study 115
Affinity 67 ASSESSMENT QUESTIONS 116
Saturation 68
Competition 69
3.9 Regulation of Protein-Binding Activity 69
Allosteric Modulation 70
Covalent Modulation 71
5 Cell Signaling in Physiology 118

Chemical Reactions and Enzymes 71

3.10 Chemical Reactions 71 5.1 Receptors 119
Determinants of Reaction Rates 72 Types of Receptors 119
Reversible and Irreversible Reactions 72 Interactions Between Receptors and Ligands 119
Law of Mass Action 73 Regulation of Receptors 122
3.11 Enzymes 74 5.2 Signal Transduction Pathways 122
Cofactors 74 Pathways Initiated by Lipid-Soluble Messengers 123
3.12 Regulation of Enzyme-Mediated Pathways Initiated by Water-Soluble Messengers 124
Reactions 75 Major Second Messengers 126
Other Messengers 130
Substrate Concentration 75
Cessation of Activity in Signal Transduction Pathways 132
Enzyme Concentration 75
Enzyme Activity 76 Chapter 5 Clinical Case Study 133
3.13 Multienzyme Reactions 76 ASSESSMENT QUESTIONS 134

Metabolic Pathways 78

6
3.14 Cellular Energy Transfer 78 Neuronal Signaling and
Glycolysis 78 the Structure of the Nervous
Krebs Cycle 80 System 136
Oxidative Phosphorylation 82
3.15 Carbohydrate, Fat, and Protein Metabolism 84
Cells of the Nervous System 137
Carbohydrate Metabolism 84
Fat Metabolism 86 6.1 Structure and Maintenance of Neurons 137
Protein and Amino Acid Metabolism 88 6.2 Functional Classes of Neurons 138
Metabolism Summary 89 6.3 Glial Cells 141
3.16 Essential Nutrients 90
6.4 Neural Growth and Regeneration 142
Vitamins 91
Growth and Development of Neurons 142
Chapter 3 Clinical Case Study 91
Regeneration of Axons 142
ASSESSMENT QUESTIONS 93
Membrane Potentials 143
6.5 Basic Principles of Electricity 143

4
6.6 The Resting Membrane Potential 144
Movement of Solutes and Water
Nature and Magnitude of the Resting Membrane Potential 144
Across Cell Membranes 95
Contribution of Ion Concentration Differences 145
Contribution of Different Ion Permeabilities 147
Contribution of Ion Pumps 148
4.1 Diffusion 96 Summary of the Development of a Resting Membrane
Magnitude and Direction of Diffusion 96 Potential 148
Diffusion Rate Versus Distance 97 6.7 Graded Potentials and Action Potentials 149
Diffusion Through Membranes 97 Graded Potentials 149
4.2 Mediated-Transport Systems 100 Action Potentials 151
Facilitated Diffusion 102
Active Transport 102 Synapses 158
4.3 Osmosis 106
6.8 Functional Anatomy of Synapses 158
Extracellular Osmolarity and Cell Volume 108
Contents vii

ISTUDY

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 Electrical Synapses 158 7.4 Association Cortex and Perceptual Processing 200
Chemical Synapses 159 Factors That Affect Perception 200
6.9 Mechanisms of Neurotransmitter Release 159
6.10 Activation of the Postsynaptic Cell 160 Specific Sensory Systems 201
Binding of Neurotransmitters to Receptors 160 7.5 Somatic Sensation 201
Removal of Neurotransmitter from the Synapse 160
Touch and Pressure 201
Excitatory Chemical Synapses 160
Posture and Movement 202
Inhibitory Chemical Synapses 161
Temperature 202
6.11 Synaptic Integration 162 Pain and Itch 202
6.12 Synaptic Strength 163 Neural Pathways of the Somatosensory System 206
Presynaptic Mechanisms 163 7.6 Vision 207
Postsynaptic Mechanisms 164 Light 207
Modification of Synaptic Transmission by Drugs and Overview of Eye Anatomy 208
Disease 164 The Optics of Vision 208
6.13 Neurotransmitters and Neuromodulators 166 Photoreceptor Cells and Phototransduction 210
Acetylcholine 166 Neural Pathways of Vision 213
Biogenic Amines 167 Color Vision 214
Amino Acid Neurotransmitters 168 Color Blindness 216
Neuropeptides 170 Eye Movement 216
Gases 171 Common Diseases of the Eye 217
Purines 171 7.7 Audition 218
Lipids 171 Sound 218
6.14 Neuroeffector Communication 171 Sound Transmission in the Ear 219
Hair Cells of the Organ of Corti 222
Structure of the Nervous System 172 Neural Pathways in Hearing 223
7.8 Vestibular System 224
6.15 Central Nervous System: Brain 172
The Semicircular Canals 224
Forebrain: The Cerebrum 172
The Utricle and Saccule 225
Forebrain: The Diencephalon 175
Vestibular Information and Pathways 225
Hindbrain: The Cerebellum 175
Brainstem: The Midbrain, Pons, and Medulla Oblongata 175 7.9 Chemical Senses 226
6.16 Central Nervous System: Spinal Cord 176 Gustation 226
Olfaction 228
6.17 Peripheral Nervous System 176
Chapter 7 Clinical Case Study 229
6.18 Autonomic Nervous System 179
ASSESSMENT QUESTIONS 232
6.19 Protective Elements Associated with the Brain 183
Meninges and Cerebrospinal Fluid 183
The Blood–Brain Barrier 184

8
Chapter 6 Clinical Case Study 185
Consciousness, the Brain,
ASSESSMENT QUESTIONS 188 and Behavior 234

8.1 States of Consciousness 235

7 Sensory Physiology 190 Electroencephalogram 235

The Waking State 236
Sleep 236
Neural Substrates of States of Consciousness 238
General Principles 191 Coma and Brain Death 240
8.2 Conscious Experiences 241
7.1 Sensory Systems and Receptors 191
Selective Attention 242
The Receptor Potential 192
Neural Mechanisms of Conscious Experiences 242
7.2 Primary Sensory Coding 193
8.3 Motivation and Emotion 244
Stimulus Type 194
Motivation 244
Stimulus Intensity 194
Emotion 245
Stimulus Location 195
Central Control of Afferent Information 197 8.4 Altered States of Consciousness 246
7.3 Ascending Neural Pathways in Sensory Systems 198 Schizophrenia 246
The Mood Disorders: Depression and Bipolar Disorders 247

viii Contents

ISTUDY
 Psychoactive Substances, Tolerance, and Substance Membrane Activation 289
Use Disorders 248 Types of Smooth Muscle 291
8.5 Learning and Memory 249 9.10 Cardiac Muscle 292
Memory 249 Cellular Structure of Cardiac Muscle 292
The Neural Basis of Learning and Memory 250 Excitation–Contraction Coupling
8.6 Cerebral Dominance and Language 251 in Cardiac Muscle 293
Chapter 8 Clinical Case Study 253 Chapter 9 Clinical Case Study 295

ASSESSMENT QUESTIONS 255 ASSESSMENT QUESTIONS 297

9 Muscle 257
10 Control of Body
Movement 300

Skeletal Muscle 258 10.1 Motor Control Hierarchy 301

Voluntary and Involuntary Actions 303
9.1 Structure 258
10.2 Local Control of Motor Neurons 303
Cellular Structure 258
Connective Tissue Structure 259 Interneurons 303
Filament Structure 259 Local Afferent Input 304
Sarcomere Structure 260 10.3 The Brain Motor Centers and the Descending Pathways
Other Myofibril Structures 261 They Control 308
9.2 Molecular Mechanisms of Cerebral Cortex 308
Skeletal Muscle Contraction 262 Subcortical and Brainstem Nuclei 310
Membrane Excitation: Cerebellum 310
The Neuromuscular Junction 262 Descending Pathways 311
Excitation–Contraction Coupling 265 10.4 Muscle Tone 313
Sliding-Filament Mechanism 267 Abnormal Muscle Tone 313
9.3 Mechanics of Single-Fiber Contraction 270 10.5 Maintenance of Upright Posture and Balance 313
Twitch Contractions 271 10.6 Walking 315
Load–Velocity Relation 272 Chapter 10 Clinical Case Study 315
Frequency–Tension Relation 272
Length–Tension Relation 274 ASSESSMENT QUESTIONS 317
9.4 Skeletal Muscle Energy Metabolism 275
Creatine Phosphate 276

11
Oxidative Phosphorylation 276
Glycolysis 276 The Endocrine System 319
Muscle Fatigue 277
9.5 Types of Skeletal Muscle Fibers 278
9.6 Whole-Muscle Contraction 279
Control of Muscle Tension 280 General Characteristics of Hormones and Hormonal Control
Control of Shortening Velocity 281 Systems 320
Muscle Adaptation to Exercise 281 11.1 Hormones and Endocrine
Lever Action of Muscles and Bones 282 Glands 320
9.7 Skeletal Muscle Disorders 285 11.2 Hormone Structures and Synthesis 322
Muscle Cramps 285 Amine Hormones 322
Hypocalcemic Tetany 285 Peptide and Protein Hormones 322
Muscular Dystrophy 285 Steroid Hormones 323
Myasthenia Gravis 286 11.3 Hormone Transport in the Blood 326
11.4 Hormone Metabolism and Excretion 327
Smooth and Cardiac Muscle 286
11.5 Mechanisms of Hormone Action 327
9.8 Structure of Smooth Muscle 286
Hormone Receptors 327
9.9 Smooth Muscle Contraction and Its Control 287 Events Elicited by Hormone–Receptor Binding 328
Cross-Bridge Activation 287 Pharmacological Effects of Hormones 328
Sources of Cytosolic Ca2+ 289 11.6 Inputs That Control Hormone Secretion 329

Contents ix

ISTUDY

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 Control by Plasma Concentrations of Mineral Ions or Organic

12
Nutrients 329
Control by Neurons 329
Cardiovascular
Control by Other Hormones 330 Physiology 361
11.7  Types of Endocrine Disorders 330
Hyposecretion 331 General Features of the Circulatory System 362
Hypersecretion 331
Hyporesponsiveness and Hyperresponsiveness 331 12.1  Components of the Circulatory System 362
Blood 362
The Hypothalamus and Pituitary Gland 332 Plasma 363
The Blood Cells 363
11.8  Control Systems Involving the Hypothalamus and Blood Flow 367
Pituitary Gland 332 Circulation 367
Posterior Pituitary Hormones 332 12.2  Pressure, Flow, and Resistance 369
Anterior Pituitary Gland Hormones and the Hypothalamus 334

The Heart 371

The Thyroid Gland 338
12.3  Anatomy 371
11.9  Synthesis of Thyroid Hormone 338
Cardiac Muscle 372
11.10 Control of Thyroid Function 340
12.4  Heartbeat Coordination 373
11.11 Actions of Thyroid Hormone 341 Sequence of Excitation 374
Metabolic Actions 341 Cardiac Action Potentials and Excitation of the SA Node 375
Permissive Actions 341 The Electrocardiogram 377
Growth and Development 341 Excitation–Contraction Coupling 377
11.12 Hypothyroidism and Hyperthyroidism 342 Refractory Period of the Heart 378
12.5  
Mechanical Events
The Endocrine Response to Stress 343 of the Cardiac Cycle 380
11.13 Physiological Functions of Cortisol 343 Mid-Diastole to Late Diastole 382
Systole 382
11.14 Functions of Cortisol in Stress 344 Early Diastole 383
11.15 Adrenal Insufficiency and Cushing’s Syndrome 345 Pulmonary Circulation Pressures 383
11.16 Other Hormones Released During Stress 346 Heart Sounds 384
12.6   The Cardiac Output 384
Endocrine Control of Growth 347 Control of Heart Rate 385
Control of Stroke Volume 385
11.17 Bone Growth 347
12.7  Measurement of Cardiac Function 389
11.18 Environmental Factors Influencing Growth 348
11.19 Hormonal Influences on Growth 348 The Vascular System 389
Growth Hormone and Insulin-Like Growth Factors 348
12.8  Overview of the Vascular System 389
Thyroid Hormone 350
Insulin 350 12.9  Arteries 391
Sex Steroids 350 Arterial Blood Pressure 391
Cortisol 350 Measurement of Systemic Arterial Pressure 392
12.10 Arterioles 394
Endocrine Control of Ca2+ Homeostasis 351 Local Controls 395
11.20 Effector Sites for Ca 2+
Homeostasis 351 Extrinsic Controls 396
Endothelial Cells and Vascular Smooth Muscle 397
Bone 351
Arteriolar Control in Specific Organs 398
Kidneys 352
Gastrointestinal Tract 352
12.11 Capillaries 399
11.21 Hormonal Controls 352 Anatomy of the Capillary Network 400
Velocity of Capillary Blood Flow 400
Parathyroid Hormone 353
Diffusion Across the Capillary Wall: Exchanges of Nutrients and
1,25-Dihydroxyvitamin D 353
Metabolic End Products 401
Calcitonin 354
Bulk Flow Across the Capillary Wall: Distribution of the
11.22 Metabolic Bone Diseases 354 Extracellular Fluid 402
Hypercalcemia 355 12.12 Venules and Veins 405
Hypocalcemia 355 Determinants of Venous Pressure 406
Chapter 11 Clinical Case Study 356 12.13 The Lymphatic System 407
ASSESSMENT QUESTIONS 359 Mechanism of Lymph Flow 409

x Contents

ISTUDY
 Integration of Cardiovascular Function: Regulation of Inspiration 453
Systemic Arterial Pressure 409 Expiration 455
12.14 Overview of Regulation of Systemic Arterial 13.3  Lung Mechanics 456
Pressure 409 Lung Compliance 456
Airway Resistance 458
12.15 Baroreceptor Reflexes 413
Lung Volumes and Capacities 459
Arterial Baroreceptors 413
13.4  Alveolar Ventilation 460
The Medullary Cardiovascular Center 414
Operation of the Arterial Baroreceptor Reflex 415 Dead Space 461
Other Baroreceptors 415 13.5  Exchange of Gases in Alveoli and Tissues 462
12.16 Blood Volume and Long-Term Regulation of Arterial Partial Pressures of Gases 462
Pressure 416 Alveolar Gas Pressures 464
Gas Exchange Between Alveoli and Blood 465
12.17 Other Cardiovascular Reflexes and Responses 417
Matching of Ventilation and Blood Flow in Alveoli 466
Cardiovascular Patterns in Health and Disease 417 Gas Exchange Between Tissues and Blood 467
13.6  Transport of Oxygen in Blood 468
12.18 Hemorrhage and Other Causes of Hypotension 417
What Is the Effect of PO2 on Hemoglobin Saturation? 468
Shock 419 Effects of Other Factors on Hemoglobin Saturation and Oxygen-
12.19 The Upright Posture 419 Carrying Capacity 470
12.20 Exercise 420 13.7  Transport of Carbon Dioxide in Blood 473
Maximal Oxygen Consumption and Training 422 13.8  Transport of Hydrogen Ion Between Tissues and
12.21 Hypertension 424 Lungs 474
12.22 Heart Failure 425 13.9   Control of Respiration 475
12.23 Hypertrophic Cardiomyopathy 427 Neural Generation of Rhythmic Breathing 475
Control of Ventilation by PO2 , PCO2, and H+ Concentration 476
12.24 Coronary Artery Disease and Heart Attacks 428
Control of Ventilation During Exercise 480
Causes and Prevention 428
Other Ventilatory Responses 482
Drug Therapy 430
13.10 Hypoxia 483
Interventions 430
Stroke and TIA 430 Why Do Ventilation–Perfusion Abnormalities Affect O2 More
Than CO2? 483
Hemostasis: The Prevention of Blood Loss 430 Emphysema 484
Acclimatization to High Altitude 484
12.25 Overview of Hemostasis 430 13.11 Nonrespiratory Functions of the Lungs 485
12.26 Formation of a Platelet Plug 431 Chapter 13 Clinical Case Study 486
12.27 Blood Coagulation: Clot Formation 432
ASSESSMENT QUESTIONS 488
12.28 Anticlotting Systems 436
Factors That Oppose Clot Formation 436
The Fibrinolytic System 436

14
12.29 Anticlotting Drugs 437 The Kidneys and
Chapter 12 Clinical Case Study 438 Regulation of Water
and Inorganic Ions 490
ASSESSMENT QUESTIONS 442

Basic Principles of Renal Physiology 491

13
14.1  Renal Functions 491
Respiratory Physiology 445
14.2  Structure of the Kidneys and Urinary System 492
14.3   Basic Renal Processes 496
Glomerular Filtration 497
13.1  Organization of the Respiratory System 446 Tubular Reabsorption 499
The Airways and Blood Vessels 446 Tubular Secretion 501
Site of Gas Exchange: The Alveoli 448 Metabolism by the Tubules 501
Relation of the Lungs to the Thoracic (Chest) Wall 449 Regulation of Membrane Channels and Transporters 501
“Division of Labor” in the Tubules 502
13.2   Principles of Ventilation 450
14.4  The Concept of Renal Clearance 502
Ventilation 450
Boyle’s Law 451 14.5  Micturition 504
Transmural Pressures 451 Involuntary (Spinal) Control 504
How Is a Stable Balance of Transmural Pressures Achieved Voluntary Control 504
Between Breaths? 452 Incontinence 505
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 Regulation of Ion and Water Balance 505 15.6 The Small Intestine 551
14.6  Total-Body Balance of Sodium and Water 505 Anatomy 551
Secretions 552
14.7  Basic Renal Processes for Sodium and Water 506
Digestion and Absorption in the Small Intestine 557
Primary Active Na+ Reabsorption 506 Motility of the Small Intestine 563
Coupling of Water Reabsorption to Na+ Reabsorption 507 15.7 The Large Intestine 564
Urine Concentration: The Countercurrent Multiplier
System 509 Anatomy 564
Secretion, Digestion, and Absorption in the Large Intestine 565
14.8   Renal Sodium Regulation 513
Motility of the Large Intestine and Defecation 565
Control of GFR 513
15.8 Pathology of the Digestive System 567
Control of Na+ Reabsorption 514
Ulcers 567
14.9   Renal Water Regulation 517
Vomiting 567
Osmoreceptor Control of Vasopressin Secretion 517 Gallstones 569
Baroreceptor Control of Vasopressin Secretion 518 Lactose Intolerance 569
14.10 A Summary Example: The Response to Sweating 519 Constipation and Diarrhea 569
14.11 Thirst and Salt Appetite 519 Chapter 15 Clinical Case Study 570
14.12 Potassium Regulation 520 ASSESSMENT QUESTIONS 573
Renal Regulation of K+ 520
14.13 Renal Regulation of Calcium and Phosphate Ions 522
14.14 Summary—Division of Labor 522

16
Regulation of Organic
14.15 Diuretics 523 Metabolism and Energy
Balance 574
Hydrogen Ion Regulation 523
14.16 Sources of Hydrogen Ion Gain or Loss 523 Control and Integration of Carbohydrate, Protein,
14.17 Buffering of Hydrogen Ion in the Body 524 and Fat Metabolism 575
14.18 Integration of Homeostatic Controls 525 16.1 Events of the Absorptive and Postabsorptive States 575
14.19 Renal Mechanisms 525 Absorptive State 575
HCO3− Handling 526 Postabsorptive State 579
Addition of New HCO3− to the Plasma 526 16.2 Endocrine and Neural Control of the Absorptive
14.20 Classification of Acidosis and Alkalosis 527 and Postabsorptive States 581
Chapter 14 Clinical Case Study 529 Insulin 581
Glucagon 585
ASSESSMENT QUESTIONS 532
Epinephrine and Sympathetic Nerves to Liver and Adipose
Tissue 585
Cortisol 586

15
Growth Hormone 586
The Digestion and Hypoglycemia 587
Absorption of Food 534 16.3 Energy Homeostasis in Exercise and Stress 588

Regulation of Total-Body Energy Balance 589

15.1  Overview of the Digestive System 535 16.4 General Principles of Energy Expenditure 589
15.2  Structure of the Gastrointestinal Tract Wall 538 Metabolic Rate 589
15.3  How Are Gastrointestinal Processes Regulated? 539 16.5 Regulation of Total-Body Energy Stores 591
Neural Regulation 540 Regulation of Food Intake 591
Hormonal Regulation 540 Overweight and Obesity 593
Phases of Gastrointestinal Control 541 What Should We Eat? 593
15.4  Mouth, Pharynx, and Esophagus 542
Regulation of Body Temperature 594
Saliva 542
Chewing 543 16.6 General Principles of Thermoregulation 594
Swallowing 543 Mechanisms of Heat Loss or Gain 595
15.5  The Stomach 545 Temperature-Regulating Reflexes 595
Anatomy 545 Temperature Acclimatization 597
Secretions of the Stomach 545 16.7 Fever and Hyperthermia 598
Gastric Motility 549 Chapter 16 Clinical Case Study 600
ASSESSMENT QUESTIONS 602

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 17.16 Additional Effects of Gonadal Steroids 635

17 Reproduction 604 17.17 Puberty (Female) 636

17.18 Female Sexual Response 637
17.19 Menopause 637
Overview and Gametogenesis, Sex Determination,
and Sex Differentiation; General Principles of Pregnancy, Contraception, Infertility, and Hormonal
Reproductive Endocrinology 605 Changes Through Life 637
17.1   Overview and Gametogenesis 605 17.20 Fertilization and Early Development 637
Gametogenesis 605 Egg Transport 637
17.2  Sex Determination 607 Intercourse, Sperm Transport, and Capacitation 638
Fertilization 638
17.3  Sex Differentiation 608 Early Development, Implantation, and Placentation 639
Differentiation of the Gonads 608 17.21 Hormonal and Other Changes During Pregnancy 643
Differentiation of Internal
Preeclampsia and the Nausea and Vomiting of Pregnancy 645
and External Genitalia 608
Fetal and Neonatal Programming 611 17.22 Parturition and Lactation 645
Sexual Differentiation of the Brain 612 Parturition 645
17.4  General Principles of Reproductive Endocrinology 612 Lactation 646
Androgens 612 17.23 Contraception and Infertility 650
Estrogens and Progesterone 612 Contraception 650
Effects of Gonadal Steroids 613 Infertility 651
Hypothalamo–Pituitary–Gonadal Control 613 17.24 Summary of Reproductive Hormones
Through Life 652
Male Reproductive Physiology 614 Fetal Life 652
17.5  Anatomy of the Male Reproductive System 614 Infancy: The Minipuberty 652
17.6  Spermatogenesis 616 Puberty 652
Adult 652
Sertoli Cells 617
Aging 652
Leydig Cells 617
Production of Mature Sperm 617 Chapter 17 Clinical Case Study 654
17.7   Transport of Sperm 618 ASSESSMENT QUESTIONS 657
Erection 619
Ejaculation 619
17.8  Hormonal Control of Male Reproductive Functions 620
Control of the Testes 620
Testosterone 620
17.9  Puberty (Male) 622
18 The Immune System 659

Secondary Sex Characteristics and Growth 622

Behavior 622 18.1  Cells and Secretions Mediating Immune Defenses 660
Anabolic Steroid Use 622 Immune Cells 660
17.10 Hypogonadism 623 Immune Cell Secretions: Cytokines 661
17.11 Andropause 624 18.2   Innate Immune Responses 663
Defenses at Body Surfaces 663
Female Reproductive Physiology 624 Inflammation 663
Interferons 667
17.12 Overview and Anatomy of the Female Reproductive
Toll-Like Receptors 668
System 624
18.3   Adaptive Immune Responses 669
Anatomy of the Female Reproductive System 624
Overview 669
17.13 Ovarian Functions 625
Lymphoid Organs and Lymphocyte Origins 669
Oogenesis 625 Humoral and Cell-Mediated Responses: Functions of B Cells
Follicle Growth 626 and T Cells 672
Formation of the Corpus Luteum 627 Lymphocyte Receptors 672
Sites of Synthesis of Ovarian Hormones 627 Antigen Presentation to T Cells 675
17.14 Control of Ovarian Function 628 NK Cells 677
Follicle Development and Estrogen Development of Immune Tolerance 677
Synthesis During the Early and Middle Follicular Phases 629 Antibody-Mediated Immune Responses: Defenses Against
LH Surge and Ovulation 630 Bacteria, Extracellular Viruses, and Toxins 677
The Luteal Phase 632 Defenses Against Virus-Infected Cells and Cancer Cells 681
17.15 Uterine Changes in the Menstrual Cycle 633 18.4  Systemic Manifestations of Infection 684
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 18.5 Factors That Alter the Resistance to Infection 685 Diagnosis 703
Acquired Immune Deficiency Syndrome (AIDS) 685 Physiological Integration 704
Antibiotics 686 Therapy 705
18.6 Harmful Immune Responses 687 19.3 Case Study of a Man with Abdominal Pain, Fever, and
Graft Rejection 687
Circulatory Failure 705
Transfusion Reactions 687 Case Presentation 705
Hypersensitivities 688 Physical Examination 705
Autoimmune Disease 690 Laboratory Tests 706
Excessive Inflammatory Responses 690 Diagnosis 706
Chapter 18 Clinical Case Study 693 Physiological Integration 707
Therapy 708
ASSESSMENT QUESTIONS 695 19.4 Case Study of a College Student with Nausea, Flushing,
and Sweating 709
Case Presentation 709

19
Physical Examination 709
Medical Physiology: Integration Laboratory Tests 710
Using Clinical Cases 697 Diagnosis 710
Physiological Integration 710
Therapy 712
19.1 Case Study of a Woman with Palpitations and Heat
Intolerance 698 APPENDIX A ANSWERS TO TEST AND REVIEW QUESTIONS A-1
Case Presentation 698 APPENDIX B INDEX OF CLINICAL TERMS A-42
Physical Examination 698
APPENDIX C CONCENTRATION RANGES OF COMMONLY MEASURED
Laboratory Tests 698
VARIABLES IN BLOOD A-46
Diagnosis 699
Physiological Integration 701
Therapy 701 GLOSSARY/INDEX GI-1
19.2 Case Study of a Man with Chest Pain After a Long
Airplane Flight 702
Case Presentation 702
Physical Examination 702
Laboratory Tests 703

Table of Contents credits: Ch. 1 Andre Schoenherr/Stone/Getty Images; Ch. 2 Andrew Dunn/Alamy Stock Photo; Ch. 3 Professors Pietro
M. Motta & Tomonori Naguro/Science Source; Ch. 4 VVG/Science Photo Library/Science Source; Ch. 5 Dr. Mark J. Winter/Science
Source; Ch. 6 David Becker/Science Source; Ch. 7 Dr. Robert Fettiplace; Ch. 8 Sherbrooke Connectivity Imaging Lab (SCIL)/Getty Images;
Ch. 9 Steve Gschmeissner/Science Source; Ch. 10 Blend Images - Erik Isakson/Brand X Pictures/Getty Images; Ch. 11 Living Art
Enterprises/Science Source; Ch. 12 SPL/Science Source; Ch. 13 SPL/Science Source; Ch. 14 Steve Gschmeissner/Science Photo Library/
Getty Images; Ch. 15 Steve Gschmeissner/Science Photo Library/Science Source; Ch. 16 The Rockefeller University/AP Images; Ch. 17
David M. Phillips/Science Source; Ch. 18 Corona Borealis Studio/Shutterstock; Ch. 19 Comstock Images/Getty Images

xiv Contents

ISTUDY
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 My western windows are most beautiful in early evening. On these
lovely, cool September nights the level and quiescent dust of light
which fills the sky is as autumnal in its colouring as the earth below.
There is autumn on the earth and autumn overhead. The great isles
of tawny orange smouldering into darkness, the paths of the
channels stilled to twilight bronze, the scarlet meadows deepening to
levels of purple and advancing night—all these mount, in exhalation
of colour, to the heavens. The beam of Nauset, entering my northern
casement, brushes a recurrent pallor of light across a part of my
bedroom wall. A first flash, a second flash, a third flash, and then a
little interval as the dark sector of the lens travels between the
Fo’castle and the flame. On bright moonlit nights, I can see both the
whitewashed tower and the light; on dark nights, I can see only the
light itself suspended and secure above the earth.
It is dark to-night, and over the plains of ocean the autumnal sky
rolls up the winter stars.
 Chapter II
AUTUMN, OCEAN, AND BIRDS

I
There is a new sound on the beach, and a greater sound. Slowly,
and day by day, the surf grows heavier, and down the long miles of
the beach, at the lonely stations, men hear the coming winter in the
roar. Mornings and evenings grow cold, the northwest wind grows
cold; the last crescent of the month’s moon, discovered by chance in
a pale morning sky, stands north of the sun. Autumn ripens faster on
the beach than on the marshes and the dunes. Westward and
landward there is colour; seaward, bright space and austerity. Lifted
to the sky, the dying grasses on the dune tops’ rim tremble and lean
seaward in the wind, wraiths of sand course flat along the beach,
the hiss of sand mingles its thin stridency with the new thunder of
the sea.
I have been spending my afternoons gathering driftwood and
observing birds. The skies being clear, noonday suns take something
of the bite out of the wind, and now and then a warmish west-
sou’westerly finds its way back into the world. Into the bright, vast
days I go, shouldering home my sticks and broken boards and
driving shore birds on ahead of me, putting up sanderlings and
sandpipers, ringnecks and knots, plovers and killdeer, coveys of a
dozen, little flocks, great flocks, compact assemblies with a
regimented air. For a fortnight past, October 9th to October 23d, an
enormous population of the migrants has been “stopping over” on
my Eastham sands, gathering, resting, feeding, and commingling.
They come, they go, they melt away, they gather again; for actual
miles the intricate and inter-crisscross pattern of their feet runs
unbroken along the tide rim of Cape Cod.
 Yet it is no confused and careless horde through which I go, but
an army. Some spirit of discipline and unity has passed over these
countless little brains, waking in each flock a conscious sense of its
collective self and giving each bird a sense of himself as a member
of some migrant company. Lone fliers are rare, and when seen have
an air of being in pursuit of some flock which has overlooked them
and gone on. Swift as the wind they fly, speeding along the breakers
with the directness of a runner down a course, and I read fear in
their speed. Sometimes I see them find their own and settle down
beside them half a mile ahead, sometimes they melt away into a
vista of surf and sky, still speeding on, still seeking.
The general multitude, it would seem, consists of birds who have
spent the summer somewhere on the outer Cape and of autumn
reinforcements from the north.
I see the flocks best when they are feeding on the edge of a tide
which rises to its flood in the later afternoon. No summer blur of
breaker mist or glassiness of heat now obscures these outer
distances, and as on I stride, keeping to the lower beach when
returning with a load, I can see birds and more birds and ever more
birds ahead. Every last advance of a dissolved breaker, coursing on,
flat and seething, has those who run away before it, turning its flank
or fluttering up when too closely pursued; every retreating in-sucked
slide has those who follow it back, eagerly dipping and gleaning.
Having fed, the birds fly up to the upper beach and sit there for
hours in the luke-cold wind, flock by flock, assembly by assembly.
The ocean thunders, pale wisps and windy tatters of wintry cloud
sail over the dunes, and the sandpipers stand on one leg and dream,
their heads tousled deep into their feathers.
I wonder where these thousands spend the night. Waking the
other morning just before sunrise, I hurried into my clothes and
went down to the beach. North and then south I strolled, along an
ebbing tide, and north and south the great beach was as empty of
bird life as the sky. Far to the south, I remember now, a frightened
pair of semipalmated sandpipers did rise from somewhere on the
upper beach and fly toward me swift and voiceless, pass me on the
flank, and settle by the water’s edge a hundred yards or so behind.
They instantly began to run about and feed, and as I watched them
an orange sun floated up over the horizon with the speed and
solemnity of an Olympian balloon.
The tide being high these days late in the afternoon, the birds
begin to muster on the beach about ten o’clock in the morning.
Some fly over from the salt meadows, some arrive flying along the
beach, some drop from the sky. I startle up a first group on turning
from the upper beach to the lower. I walk directly at the birds—a
general apprehension, a rally, a scutter ahead, and the birds are
gone. Standing on the beach, fresh claw marks at my feet, I watch
the lovely sight of the group instantly turned into a constellation of
birds, into a fugitive pleiades whose living stars keep their chance
positions; I watch the spiralling flight, the momentary tilts of the
white bellies, the alternate shows of the clustered, grayish backs.
The group next ahead, though wary from the first, continues
feeding. I draw nearer; a few run ahead as if to escape me afoot,
others stop and prepare to fly; nearer still, the birds can stand no
more; another rally, another scutter, and they are following their kin
along the surges.
No aspect of nature on this beach is more mysterious to me than
the flights of these shore-bird constellations. The constellation
forms, as I have hinted, in an instant of time, and in that same
instant develops its own will. Birds which have been feeding yards
away from each other, each one individually busy for his individual
body’s sake, suddenly fuse into this new volition and, flying, rise as
one, coast as one, tilt their dozen bodies as one, and as one wheel
off on the course which the new group will has determined. There is
no such thing, I may add, as a lead bird or guide. Had I more space
I should like nothing better than to discuss this new will and its
instant of origin, but I do not want to crowd this part of my chapter,
and must therefore leave the problem to all who study the psychic
relations between the individual and a surrounding many. My special
interest is rather the instant and synchronous obedience of each
speeding body to the new volition. By what means, by what
methods of communication does this will so suffuse the living
constellation that its dozen or more tiny brains know it and obey it in
such an instancy of time? Are we to believe that these birds, all of
them, are machina, as Descartes long ago insisted, mere
mechanisms of flesh and bone so exquisitely alike that each
cogwheel brain, encountering the same environmental forces,
synchronously lets slip the same mechanic ratchet? or is there some
psychic relation between these creatures? Does some current flow
through them and between them as they fly? Schools of fish, I am
told, make similar mass changes of direction. I saw such a thing
once, but of that more anon.
We need another and a wiser and perhaps a more mystical
concept of animals. Remote from universal nature, and living by
complicated artifice, man in civilization surveys the creature through
the glass of his knowledge and sees thereby a feather magnified and
the whole image in distortion. We patronize them for their
incompleteness, for their tragic fate of having taken form so far
below ourselves. And therein we err, and greatly err. For the animal
shall not be measured by man. In a world older and more complete
than ours they move finished and complete, gifted with extensions
of the senses we have lost or never attained, living by voices we
shall never hear. They are not brethren, they are not underlings;
they are other nations, caught with ourselves in the net of life and
time, fellow prisoners of the splendour and travail of the earth.
The afternoon sun sinks red as fire; the tide climbs the beach, its
foam a strange crimson; miles out, a freighter goes north, emerging
from the shoals.

II
It chanced that on a mild September morning, as I was standing a
moment at a window looking west over the marshes and the blue
autumnal creeks, an alarm of some kind began to spread among the
gulls. The incoming tide had already crowded the birds back on the
higher gravel banks and bars, and from these isles, silvery cloud by
cloud, I saw the gulls rise and stream away to the southward in a
long, fugitive storm of wings. They were flying, I noticed, unusually
low. Interested to see what had thus disturbed them, I stepped out
a moment to the pinnacle of my dune. As I stood there staring after
the vanishing gulls and questioning the sky, I saw far above the
birds, and well behind them, an eagle advancing through the
heavens. He had just emerged from a plume of hovering cloud into
the open blue, and when I saw him first was sailing south and
seaward on motionless wings, seeming to follow in the great sky the
blue course of a channel far below.
There are sand bars at the mouth of Nauset harbour; many gulls
feed there between the tides, and the gulls from the marsh joined
forces with this gathering. As the eagle approached the bars, I
looked to see if he would descend or fly out to sea. But no; at the
harbour’s entrance he turned south, aligned his flight with the coast
line, and disappeared.
During the autumn I saw this same bird half a dozen different
times. I could tell when he was about by the terror of the gulls. Yet
this eagle—for a bald eagle, Haliætus leucocephalus leucocephalus, I
believe him to have been—is, as Mr. Forbush says, “by nature a fish
eater.” I never saw him pay the slightest attention to the fugitives;
nevertheless, he may well have a fancy for gulls when they are
plump and he is hungry. At any rate, they fear him. There are always
a few black-backed or “minister” gulls mingled in with the herring
gulls upon these flats, and these burly giants, I noticed, sought
refuge with the rest.
Eagles are by no means rare upon Cape Cod. The birds arrive here
as coast-wise visitors, find the region to their liking, and establish
themselves in various favourite domains. They fish in our sandy bays
and inlets; they have rather a fancy for the more isolated Cape Cod
ponds. Seen at close range, the bald eagle is a dusky brownish bird
with a pure-white head, neck, and tail. I never had a near view of
this Eastham visitor, but one of the coast guardsmen roused him up
one day from a thicket close by the head of a creek running up into
the moors—he heard a sudden noise of brush and great wings, he
said, and, turning round, he saw the eagle rising free of the scrub
and the bright leaves.
 Ever since I came to live here on Cape Cod, I have been amazed
at the number of land-bird migrants I have encountered on the
dunes. I expected to see sandpipers on the beach and scoters in the
surf, for they are coast-wise folk, but I did not expect to see the red-
breasted nuthatch rise out of the September dunes, or find the
charming black-and-yellow warbler sitting on the ridgepole of the
Fo’castle, his black-tipped tail feathers turned to the Atlantic. But
perhaps I had best begin at the beginning and tell how the sparrows
and the warblers came down to us this autumn by the coast.
Various new sparrows were the first strangers to arrive. There are
summer sparrows here, a great abundance of them, for the marsh
and meadow land west of the dunes is the natural habitat of many
species. Walk through these grass-lands on a summer’s day, and you
will see singles and flocks break from the sunburned stubble ahead,
some to drop and hide again farther on, others to watch you from
the coastguard wires. Song sparrows are notably abundant, for
these pleasant singers frequent both the marshes and the dunes;
but the seaside sparrow keeps more to the marsh rim and the salt-
hay mowings, the sharp-tailed sparrow fancies the wheel ruts of the
hay carts, and the odd little grasshopper sparrow, Coturniculus
savannarum passerinus, trills into the burning sundowns the two
faint notes of his curious and poignant insect song.
Early in September Hudsonian curlews arrived in Eastham marsh,
and to see them I began going to Nauset through the meadows
instead of by the beach. High September tides were then covering
both marsh and meadow land, and, as I pushed on each afternoon,
the curlews rose from close beside the inundated road, and, circling,
called to other curlews; I could hear, when I listened, the clear reply.
And then there would be silence, and I would hear the sound of
autumn and the world, and perhaps the faint withdrawing roar of
ocean beyond the dunes. When I reached the wider meadows on
these days, I found the stubble mobbed with sparrows; the
population had doubled in a week.
Flocks of fox sparrows were feeding everywhere; I whirred up
groups of savannah sparrows and families of white-throats; a
solitary white-headed sparrow watched me from the concealment of
a bush. It was a silent throng. I heard faint “tsips” and “chips” of
alarm as I passed—nothing more. Love-making was over and done,
and all were importantly busy with the importance of their lives.
On the 24th and the 25th there was wind and rain, and on the
27th I saw the first of the warblers.
The weather had cleared, and I had risen early and begun to get
breakfast. It is my custom here to sit facing the sea, and I was
moving over my table when I noticed a small bird of some kind
foraging about in the grass before the house. I could not see him
well at first, for he had entered into the grass as into a thicket, but
presently out he came, pushing through the stalks, and I watched
him from the window unsuspected. This first arrival was a Canadian
warbler. Steely ash-gray above, yellow below, and with a broad band
of black spots between his yellow throat and yellow belly, he was a
charming bit of life. Over the pale sand, in and out of the tawny-
white roots, in and out of the variegations of morning light he
moved, picking up seeds while a sea wind shook the tops of the
dying grass above his head. Presently, in search of still more food,
he turned the corner of the house, and when I went out after
breakfast he had gone.
Then came, all in one week, a Wilson’s warbler (a female,
probably), the black-and-yellow warbler, and a chestnut-sided
warbler. The birds were singles, they travelled along the dunes, they
fed on the fallen seeds. In October I saw in one day five myrtle
warblers; a pair of these lingered a week near Fo’castle dune. Then
came juncos and a raid of pigeon hawks. The juncos, like the
warblers, foraged on the dunes, and the hawks hunted them there
for an hour or so before the dawn. I went exploring one morning
while Nauset was still flashing into a sullen, cold, and overclouded
world, and saw a pigeon hawk rise unexpectedly out of the cut to
the north with a wretched junco gripped beneath him. Flying
seaward through the cut, the hawk carried his captive to the beach,
found himself a sheltered nook close along the dune wall, stood at
attention for a moment, and then unbent and ate.
I saw various other migrant land birds as well, but I shall not dwell
upon them, for the listing and cataloguing of species seems to me of
less interest than their arrival by sea. This outer arm of Cape Cod, as
I have already explained, stands thirty miles or so out from the
continental main, yet there are land birds, little birds, going south
along it as casually as so many arctic geese. Writing here this cloudy
morning, with a great confused roaring of breakers in my ears, I call
to mind the Wilson’s warbler, the female, I saw a fortnight ago, and I
wonder where it was that she forsook her familiar earth for the grey
ocean, an ocean she perhaps had never seen. What a gesture of
ancient faith and present courage such a flight is, what a defiance of
circumstance and death—land wing and hostile sea, the fading land
behind, the unknown and the distant articulate and imperious in the
bright, aërial blood.
But who shall say by what sea routes these landsmen reach the
Cape? Some species, I imagine, cross Massachusetts Bay, their
jumping-off place being north of Boston (Cape Ann or Ipswich
perhaps); some may cross over from the South Shore at a point well
north of Cape Cod Bay, others undoubtedly come directly down from
Maine. The wooded archipelago of Maine is a famous place for
warblers. It is quite possible that the species I have mentioned may
have followed some great river to the sea, the Kennebec or the
Penobscot, perhaps, and crossed from the river mouth directly over
to Cape Cod. The Highland Light bears south ¾ west (true) from
Seguin at the mouth of the Kennebec, and is separated from it by
only 101 miles of open water. The birds could manage this easily.
 Herring Gulls
All over the world land migrants go great distances over open
water. Numbers of birds, for instance, migrating back and forth
between Europe and North Africa cross the Mediterranean twice a
year, and in our own hemisphere there are flights across the Gulf of
Mexico and movements between the West Indies and our south
Atlantic states.
Late in October there came an easterly gale, and in the afternoon,
when the tide was high, I put on oilskins and went out to see the
surf. About a mile to the north of the Fo’castle, as I was trudging on
across the rain, I saw just ahead and close over the breakers a flying
speck sailing landward out of the wrack, and even as I stared, it fell
to the beach in danger of the waves. I ran ahead, then, and picked
the thing up just as a slide of foam was about to overflow it, and
found it to be an autumn leaf, a maple leaf flat and drenched and
red.
Mid-October and the land birds have gone. A few sparrows linger
in the marshes. The plum bushes have lost their leaves. Walking the
beach, I read winter in the new shapes of the clouds.
 III
Western cloud, dark substance of cloud, gathered at the wintry
horizon of the short-lived days making them even shorter with the
false sundown of its rim. Now come the sea fowl and the wild fowl
to the beach from the lonely and darkening north, from the Arctic
Ocean and the advancing pack, from the continental fragments and
great empty islands that lie between the continent and the pole,
from the tundra and the barrens, from the forests, from the bright
lakes, from the nest-strewn crevices and ledges of Atlantic rocks no
man has ever named or scaled. Over the round of earth, down from
the flattened summit, pour the living streams, bearing south the
tribes and gathered nations, the peoples and flocks, the clans and
families, the young and the old. And the dying grasslands, the
October snows, and the forests fall behind, and presently the nations
behold a first far glint of the sea.
There are many streams, and it is said that two of the greatest
bear down upon Cape Cod. A first river, rising in the interior of
Alaska, flows southeast across Canada to the Atlantic; on this stream
move birds from the north woods and the Canadian lakes together
with birds from the north barrens and the arctic isles and half lands;
a second stream, rising in the shadow of the pole, flows south along
the coast past Greenland and the bays of Labrador—on this move
the hardy arctic folk who get their living from the tides. Many
species are common to both streams. Somewhere north of the Cape,
perhaps round and about the mouth of the St. Lawrence, these
streams immix their multitudes, and south to New England moves
the great united flood, peopling with primeval life the seacoasts and
the sky.
Ducks enter the channels, some flying in from the bay, others
from the outer ocean, geese settle at sundown in the golden skin of
the western coves, coveys of winter yellow-legs circle in the gloom,
and hide when disturbed in the taller salt grasses between the
meadows and the creeks. At nightfall and at daybreak I hear birds
talking. Strangers in rubber boots and khaki uniforms now visit my
domain, and every Saturday afternoon I look with philosophy
through my western windows to a number of tufts of grass disguised
as gunners.
Now that I have settled down here for the winter, I find myself
becoming something of a beach comber. Every once in a while,
when I chance to look seaward, I spy an unknown something or
other rising and falling, appearing and disappearing in the offshore
surges, and at this sight the beach comber in me wakes. All kinds of
things “come ashore” on these vast sands, and even the most
valueless have an air of being treasure trove. The mysterious
something moving from the swells into the breakers may be nothing
but a smelly bait tub washed overboard from some Gloucester
fisherman, or a lobster pot, or a packing case stencilled with a liner’s
name; but in the sea or on the beach a mile ahead it is something
for nothing, it is the unknown, it is hope springing eternal in the
human breast. The other day I found myself thoughtfully examining
a U. S. Navy blue undress jumper which lay flat and soggy and
solitary on the lower beach. It was not, in its day, an unfamiliar
garment, and I have an old friend in the village who occasionally
dons a rather good one he found just south of the light. Alas, the
cloth was rotted and the jumper much too small. But I cut off and
saved the buttons.
 Semipalmated Sandpiper
Now, while I stood there slicing off the buttons, I chanced to look
up a moment at the southern sky, and there for the first and still the
only time in my life, I saw a flight of swans. The birds were passing
along the coast well out to sea; they were flying almost cloud high
and travelling very fast, and their course was as direct as an arrow’s
from a bow. Glorious white birds in the blue October heights over
the solemn unrest of ocean—their passing was more than music,
and from their wings descended the old loveliness of earth which
both affirms and heals.

IV
The last two weeks in October see the peak of the autumnal
visitations. In November and December the stream from the inland
shrinks, but the coast-wise stream, continuing to flow, brings us
down a rare and curious world. Of this I shall write at greater length,
for I found it of enormous interest.
Here, approaching the end of my notes on birds and autumn, I
chance to remember that one of the strangest and most beautiful of
the migrations over the dunes was not a movement of birds at all
but of butterflies. There came a morning early in October which
ripened, as the sun rose higher, into a rather mild and September-
like day; the wind was autumnal, I recall, and from the north by
west, but the current was both mildly warm and light. As it was a
day to be spent out-of-doors, soon after ten o’clock I went out round
the back of the Fo’castle into the sunlight and began to work there
on a bin I was putting together out of driftage. I looked about, as I
always do, but nothing in the landscape chanced to take my eye.
Sawing and hammering, I worked for about three quarters of an
hour, and then downed tools to take a moment’s rest.
During the hour, a flight of twenty or more large orange-and-black
butterflies had arrived in the region of the dunes. It was a flight, yet
were the individuals far apart. There was at least an eighth of a mile
between any two; some were on the dunes, some were on the salt
meadows, three were on the beach. Their movements were casual
as the wind, yet there was an unmistakable southerly pull drawing
them on. I tried to catch one of the travellers on the beach, and
though I count myself a fair runner, it was no easy work keeping up
with his turns and erratic doublings. I wished him no ill; I simply
wanted to have a better look at him, but he escaped me by rising
and disappearing over the top of a dune. When I reached the same
top after a scramble up a steep of sand, the fugitive was already a
good eighth of a mile away. I went back to my carpentry with an
increased respect for butterflies as fliers.
An entomologist with whom I have been in correspondence tells
me that my visitors were undoubtedly specimens of the monarch or
milkweed butterfly, Anosia plexippus. In early autumn adults gather
in great swarms and move in a generally southward direction, and it
is believed (but not proved) that New England specimens go as far
as Florida. The following spring individuals (not swarms) appear in
the North apparently coming from the South. We do not know—I am
quoting this paragraph almost verbatim—whether these are
returning migrants or whether they are individuals that had not
previously been in the North. We do know that none of the fall
migrants had previously been in the South.
 The butterflies of Eastham remained upon the dunes the rest of
the morning. I imagine that they were in search of food. Between
half-past twelve and half-past one they melted away as mysteriously
as they had come, and with them went the last echo of summer and
the high sun from the dunes. And that day I finished my bin and
filled it and began to build a wall of seaweed round the foundation
of my house. A cricket sang as I worked in the mild afternoon, alive
and hardy in his cave under my driftwood mountain, and beyond this
little familiar sound of earth I heard the roar of ocean filling the
hollow space of day with its inexorable warning.
 Chapter III
THE HEADLONG WAVE

I
This morning I am going to try my hand at something that I do
not recall ever having encountered either in a periodical or in a
book, namely, a chapter on the ways, the forms, and the sounds of
ocean near a beach. Friends are forever asking me about the surf on
the great beach and if I am not sometimes troubled or haunted by
its sound. To this I reply that I have grown unconscious of the roar,
and though it sounds all day long in my waking ears, and all night
long in my sleeping ones, my ears seldom send on the long tumult
to the mind. I hear the roar the instant I wake in the morning and
return to consciousness, I listen to it a while consciously, and then
accept and forget it; I hear it during the day only when I stop again
to listen, or when some change in the nature of the sound breaks
through my acceptance of it to my curiosity.
They say here that great waves reach this coast in threes. Three
great waves, then an indeterminate run of lesser rhythms, then
three great waves again. On Celtic coasts it is the seventh wave that
is seen coming like a king out of the grey, cold sea. The Cape
tradition, however, is no half-real, half-mystical fancy, but the truth
itself. Great waves do indeed approach this beach by threes. Again
and again have I watched three giants roll in one after the other out
of the Atlantic, cross the outer bar, break, form again, and follow
each other in to fulfilment and destruction on this solitary beach.
Coast guard crews are all well aware of this triple rhythm and take
advantage of the lull that follows the last wave to launch their boats.
It is true that there are single giants as well. I have been roused
by them in the night. Waked by their tremendous and unexpected
crash, I have sometimes heard the last of the heavy overspill,
sometimes only the loud, withdrawing roar. After the roar came a
briefest pause, and after the pause the return of ocean to the night’s
long cadences. Such solitary titans, flinging their green tons down
upon a quiet world, shake beach and dune. Late one September
night, as I sat reading, the very father of all waves must have flung
himself down before the house, for the quiet of the night was
suddenly overturned by a gigantic, tumbling crash and an
earthquake rumbling; the beach trembled beneath the avalanche,
the dune shook, and my house so shook in its dune that the flame of
a lamp quivered and pictures jarred on the wall.
The three great elemental sounds in nature are the sound of rain,
the sound of wind in a primeval wood, and the sound of outer ocean
on a beach. I have heard them all, and of the three elemental
voices, that of ocean is the most awesome, beautiful, and varied. For
it is a mistake to talk of the monotone of ocean or of the
monotonous nature of its sound. The sea has many voices. Listen to
the surf, really lend it your ears, and you will hear in it a world of
sounds: hollow boomings and heavy roarings, great watery
tumblings and tramplings, long hissing seethes, sharp, rifle-shot
reports, splashes, whispers, the grinding undertone of stones, and
sometimes vocal sounds that might be the half-heard talk of people
in the sea. And not only is the great sound varied in the manner of
its making, it is also constantly changing its tempo, its pitch, its
accent, and its rhythm, being now loud and thundering, now almost
placid, now furious, now grave and solemn-slow, now a simple
measure, now a rhythm monstrous with a sense of purpose and
elemental will.
Every mood of the wind, every change in the day’s weather, every
phase of the tide—all these have subtle sea musics all their own.
Surf of the ebb, for instance, is one music, surf of the flood another,
the change in the two musics being most clearly marked during the
first hour of a rising tide. With the renewal of the tidal energy, the
sound of the surf grows louder, the fury of battle returns to it as it
turns again on the land, and beat and sound change with the
renewal of the war.
 Sound of surf in these autumnal dunes—the continuousness of it,
sound of endless charging, endless incoming and gathering, endless
fulfilment and dissolution, endless fecundity, and endless death. I
have been trying to study out the mechanics of that mighty
resonance. The dominant note is the great spilling crash made by
each arriving wave. It may be hollow and booming, it may be heavy
and churning, it may be a tumbling roar. The second fundamental
sound is the wild seething cataract roar of the wave’s dissolution and
the rush of its foaming waters up the beach—this second sound
diminuendo. The third fundamental sound is the endless dissolving
hiss of the inmost slides of foam. The first two sounds reach the ear
as a unisonance—the booming impact of the tons of water and the
wild roar of the up-rush blending—and this mingled sound dissolves
into the foam-bubble hissing of the third. Above the tumult, like
birds, fly wisps of watery noise, splashes and counter splashes,
whispers, seethings, slaps, and chucklings. An overtone sound of
other breakers, mingled with a general rumbling, fells earth and sea
and air.
 The Edge of Foam
Here do I pause to warn my reader that although I have
recounted the history of a breaker—an ideal breaker—the surf
process must be understood as mingled and continuous, waves
hurrying after waves, interrupting waves, washing back on waves,
overwhelming waves. Moreover, I have described the sound of a
high surf in fair weather. A storm surf is mechanically the same
thing, but it grinds, and this same long, sepulchral grinding—sound
of utter terror to all mariners—is a development of the second
fundamental sound; it is the cry of the breaker water roaring its way
ashore and dragging at the sand. A strange underbody of sound
when heard through the high, wild screaming of a gale.
Breaking waves that have to run up a steep tilt of the beach are
often followed by a dragging, grinding sound—the note of the
baffled water running downhill again to the sea. It is loudest when
the tide is low and breakers are rolling beach stones up and down a
slope of the lower beach.
 I am, perhaps, most conscious of the sound of surf just after I
have gone to bed. Even here I read myself to drowsiness, and,
reading, I hear the cadenced trampling roar filling all the dark. So
close is the Fo’castle to the ocean’s edge that the rhythm of sound I
hear oftenest in fair weather is not so much a general tumult as an
endless arrival, overspill, and dissolution of separate great seas.
Through the dark, mathematic square of the screened half window, I
listen to the rushes and the bursts, the tramplings, and the long,
intermingled thunderings, never wearying of the sonorous and
universal sound.
Away from the beach, the various sounds of the surf melt into one
great thundering symphonic roar. Autumnal nights in Eastham village
are full of this ocean sound. The “summer people” have gone, the
village rests and prepares for winter, lamps shine from kitchen
windows, and from across the moors, the great levels of the marsh,
and the bulwark of the dunes resounds the long wintry roaring of
the sea. Listen to it a while, and it will seem but one remote and
formidable sound; listen still longer and you will discern in it a
symphony of breaker thunderings, an endless, distant, elemental
cannonade. There is beauty in it, and ancient terror. I heard it last as
I walked through the village on a starry October night; there was no
wind, the leafless trees were still, all the village was abed, and the
whole sombre world was awesome with the sound.

II
The seas are the heart’s blood of the earth. Plucked up and
kneaded by the sun and the moon, the tides are systole and diastole
of earth’s veins.
The rhythm of waves beats in the sea like a pulse in living flesh. It
is pure force, forever embodying itself in a succession of watery
shapes which vanish on its passing.
I stand on my dune top watching a great wave coursing in from
sea, and know that I am watching an illusion, that the distant water
has not left its place in ocean to advance upon me, but only a force
shaped in water, a bodiless pulse beat, a vibration.
Consider the marvel of what we see. Somewhere in ocean,
perhaps a thousand miles and more from this beach, the pulse beat
of earth liberates a vibration, an ocean wave. Is the original force
circular, I wonder? and do ocean waves ring out from the creative
beat as they do on a quiet surface broken by a stone? Are there,
perhaps, ocean circles so great and so intricate that they are
unperceived? Once created, the wave or the arc of a wave begins its
journey through the sea. Countless vibrations precede it, countless
vibrations follow after. It approaches the continent, swings into the
coast line, courses ashore, breaks, dissolves, is gone. The innermost
waters it last inhabited flow back in marbly foam to become a body
to another beat, and to be again flung down. So it goes night and
day, and will go till the secret heart of earth strikes out its last slow
beat and the last wave dissolves upon the last forsaken shore.
 Slide and Seethe
As I stand on my dune top, however, I do not think of the illusion
and the beat of earth, for I watch the waves with my outer rather
than my inner eye. After all, the illusion is set off by an
extraordinary, an almost miraculous thing—the embodiment of the
wave beat in an almost constant shape. We see a wave a quarter of
a mile off, then a few hundred yards nearer in, then just offshore;
we seem to have been watching the same travelling mass of water—
there has been no appreciable change in mass or in shape—yet all
the while the original beat has taken on a flowing series of liquid
bodies, bodies so alike, so much the same, that our eye will
individualize them and follow them in—the third wave, we say, or the
second wave behind the great wave. How strange it is that this beat
of earth, this mysterious undulation of the seas, moving through and
among the other forces stirring the waters close off the continent,
should thus keep its constancy of form and mass, and how odd a
blend of illusion and reality it all is! On the whole, the outer eye has
the best of it.
Blowing all day long, a northwest wind yesterday swept the sky
clear of every tatter and wisp of cloud. Clear it still is, though the
wind has shifted to the east. The sky this afternoon is a harmony of
universal blue, bordered with a surf rim of snowiest blue-white. Far
out at sea, in the northeast and near the horizon, is a pool of the
loveliest blue I have ever seen here—a light blue, a petal blue, blue
of the emperor’s gown in a Chinese fairy tale. If you would see
waves at their best, come on such a day, when the ocean reflects a
lovely sky, and the wind is light and onshore; plan to arrive in the
afternoon so that you will have the sun facing the breakers. Come
early, for the glints on the waves are most beautiful and interesting
when the light is oblique and high. And come with a rising tide.
The surf is high, and on the far side of it, a wave greater than its
fellows is shouldering out of the blue, glinting immensity of sea.
Friends tell me that there are certain tropic beaches where waves
miles long break all at once in one cannonading crash: a little of this,
I imagine, would be magnificent; a constancy of it, unbearable. The
surf here is broken; it approaches the beach in long intercurrent
parallels, some a few hundred feet long, some an eighth of a mile
long, some, and the longest, attaining the quarter-mile length and
perhaps just over. Thus, at all times and instants of the day, along
the five miles of beach visible from the Fo’castle deck, waves are to
be seen breaking, coursing in to break, seething up and sliding back.
But to return to the blue wave rolling in out of the blue
spaciousness of sea. On the other side of the world, just opposite
the Cape, lies the ancient Spanish province of Galicia, and the town
of Pontevedra and St. James Compostella, renowned of pilgrims.
(When I was there they offered me a silver cockle shell, but I would
have none of it, and got myself a sea shell from some Galician
fisherfolk.) Somewhere between this Spanish land and Cape Cod the
pulse of earth has engendered this wave and sent it coursing
westward through the seas. Far off the coast, the spray of its
passing has, perhaps, risen on the windward bow of some rusty
freighter and fallen in rainbow drops upon her plates; the great
liners have felt it course beneath their keels.
A continent rises in the west, and the pulse beat approaches this
bulwark of Cape Cod. Two thirds of a mile out, the wave is still a sea
vibration, a billow. Slice it across, and its outline will be that of a
slightly flattened semi-circle; the pulse is shaped in a long,
advancing mound. I watch it approach the beach. Closer and closer
in, it is rising with the rise of the beach and the shoaling of the
water; closer still, it is changing from a mound to a pyramid, a
pyramid which swiftly distorts, the seaward side lengthening, the
landward side incurving—the wave is now a breaker. Along the ridge
of blue forms a rippling crest of clear, bright water; a little spray flies
off. Under the racing foam churned up by the dissolution of other
breakers the beach now catches at the last shape of sea inhabited
by the pulse—the wave is tripped by the shoaling sand—the giant
stumbles, crashes, and is pushed over and ahead by the sloping line
of force behind. The fall of a breaker is never the work of gravity
alone.
It is the last line of the wave that has captured the decorative
imagination of the world—the long seaward slope, the curling crest,
the incurved volute ahead.
 Summer Breakers
Toppling over and hurled ahead, the wave crashes, its mass of
glinting blue falling down in a confusion of seething, splendid white,
the tumbling water rebounding from the sand to a height almost
always a little above that of the original crest. Out of the wild,
crumbling confusion born of the dissolution of the force and the last
great shape, foamy fountains spurt, and ringlets of spray. The mass
of water, still all furiously a-churn and seething white, now rushes for
the rim of the beach as it might for an inconceivable cataract. Within
thirty-five feet the water shoals from two feet to dry land. The edge
of the rush thins, and the last impulse disappears in inch-deep slides
of foam which reflect the sky in one last moment of energy and
beauty and then vanish all at once into the sands.
Another thundering, and the water that has escaped and
withdrawn is gathered up and swept forward again by another
breaking wave. Night and day, age after age, so works the sea, with
infinite variation obeying an unalterable rhythm moving through an
intricacy of chance and law.
 I can watch a fine surf for hours, taking pleasure in all its wild
plays and variations. I like to stand on my beach, watching a long
wave start breaking in many places, and see the curling water run
north and south from the several beginnings, and collide in furious
white pyramids built of the opposing energies. Splendid fountains
often delight the eye. A towering and deep-bellied wave, toppling,
encloses in its volute a quantity of air, and a few seconds after the
spill this prisoned and compressed vapour bursts up through the
boiling rush in feathery, foamy jets and geyser plumes. I have seen
fountains here, on a September day, twenty and twenty-five and
even thirty feet high. Sometimes a curious thing happens. Instead of
escaping vertically, the rolled-up air escapes horizontally, and the
breaker suddenly blows, as from a dragon’s mouth, a great lateral
puff of steamy spray. On sunny days, the toppling crest is often
mirrored in the glassy volute as the wave is breaking. One lovely
autumn afternoon, I saw a beautiful white gull sailing along the
volute of a breaker accompanied by his reflection in the wave.
I add one curious effect of the wind. When the wind is directly
offshore or well offshore, the waves approach fighting it; when the
wind is offshore but so little off that its angle with the coast line is
oblique—say an angle never greater than twenty-two degrees and
never less than about twelve—the waves that approach the coast do
not give battle, but run in with their long axis parallel to the wind.
Sitting in the Fo’castle, I can often tell the exact quarter of an
offshore wind simply by looking at this oblique alignment of the
waves.
The long miles of beach are never more beautiful than when
waves are rolling in fighting a strong breeze. Then do the breakers
actually seem to charge the coast. As they approach, the wind
meets them in a shock of war, the chargers rear but go on, and the
wind blows back their manes. North and south, I watch them
coursing in, the manes of white, sun brilliant spray streaming behind
them for thirty and even forty feet. Sea horses do men call such
waves on every coast of the world. If you would see them at their
best, come to this beach on a bright October day when a northwest
wind is billowing off to sea across the moors.
 III
I will close my chapter with a few paragraphs about heavy surf.
It is best to be seen, I think, when the wind is not too high. A gale
blows up a surf, but it also flattens out the incoming rollers, making
monstrous, foamy travelling mounds of them much like those visible
from a ship at sea. Not until the wind has dropped do the breakers
gather form. The finest surf I have ever seen here—it was a
Northern recoil of the great Florida hurricane—broke on three
pleasant and almost windless autumn days. The storm itself had
passed us, but our seas had been stirred to their deeps. Returning to
the Cape at night from a trip to town, I heard the roar of the ocean
in Orleans, and on arriving at Nauset, found the beach flooded to
the dunes, and covered with a churn of surf and moonlight.
Dragging a heavy suitcase and clad in my go-to-town clothes, I had
an evil time getting to the Fo’castle over the dune tops and along
the flooded marsh.
Many forces mingle in the surf of a storm—the great earth rhythm
of the waves, the violence of wind, the struggle of water to obey its
own elemental law. Out of the storm at sea come the giants and,
being giants, trip far out, spilling first on the outer bar. Shoreward
then they rush, breaking all the way. Touching the beach, they
tumble in a roar lost in a general noise of storm. Trampled by the
wind and everlastingly moved and lifted up and flung down by the
incoming seas, the water offshore becomes a furious glassiness of
marbly foam; wild, rushing sheets of seethe fifty feet wide border it;
the water streams with sand.
Under all this move furious tidal currents, the longshore undertow
of outer Cape Cod. Shore currents here move in a southerly
direction; old wreckage and driftwood is forever being carried down
here from the north. Coast guard friends often look at a box or stick
I have retrieved, and say, “Saw that two weeks ago up by the light.”
After an easterly, I find things on the beach which have been
blown down from the Gulf of Maine—young, uprooted spruce trees,
lobster buoys from Matinicus, and, after one storm, a great strewing
of empty sea-urchin shells. Another easterly washed up a strewing
of curious wooden pebbles shaped by the sea out of the ancient
submerged forests which lie just off the present coast. They were
brown-black, shaped like beach stones, and as smooth as such
stones.
The last creature I found in the surf was a huge horseshoe crab,
the only one I have ever chanced to find on the outside. Poor
Limulus polyphemus! The surf having turned him upside down, he
had as usual doubled up, and the surf had then filled with sand the
angle of his doubling. When I discovered him, he was being bullied
by a foam slide, and altogether in a desperate way. So I picked him
up, rinsed the sand out of his waving gills, held him up all dripping
by the tail, and flung him as far as I could to seaward of the
breakers. A tiny splash, and I had seen the last of him, a moment
more, and the surf had filled the hollow in which he had lain.
Autumnal easterlies and November tides having scoured from the
beach its summer deeps of sand, the high seasonal tides now run
clear across to the very foot of the dunes. Under this daily overflow
of cold, the last of the tide-rim hoppers and foragers vanish from the
beach. An icy wind blusters; I hear a dry tinkle of sand against my
western wall; December nears, and winter closes in upon the coast.
 Chapter IV
MIDWINTER

I
A year indoors is a journey along a paper calendar; a year in outer
nature is the accomplishment of a tremendous ritual. To share in it,
one must have a knowledge of the pilgrimages of the sun, and
something of that natural sense of him and feeling for him which
made even the most primitive people mark the summer limits of his
advance and the last December ebb of his decline. All these autumn
weeks I have watched the great disk going south along the horizon
of moorlands beyond the marsh, now sinking behind this field, now
behind this leafless tree, now behind this sedgy hillock dappled with
thin snow. We lose a great deal, I think, when we lose this sense
and feeling for the sun. When all has been said, the adventure of the
sun is the great natural drama by which we live, and not to have joy
in it and awe of it, not to share in it, is to close a dull door on
nature’s sustaining and poetic spirit.
The splendour of colour in this world of sea and dune ebbed from
it like a tide; it shallowed first without seeming to lose ground, and
presently vanished all at once, almost, so it seemed, in one grey
week. Warmth left the sea, and winter came down with storms of
rushing wind and icy, pelting rain. The first snow fell early in
November, just before the dawn of a grey and bitter day. I had
written a letter the night before, intending to give it to the coast
guardsman who came south at seven o’clock, but somehow or other
I missed him; and no welcome light flashed an answer to mine as I
stood on the crest of my dune looking into the darkness of the
beach and listening to the sombre thunder of a rising sea. Unwilling
to stay up till after midnight for the next patrol, I went out and put a
note on the coast guard key post just south of me asking the last
man south in the morning to wake me up and come in and get the
letter. At about half-past five I woke to a stamping of feet and a
knock on the door, and in came John Blood, the tall, light-haired
New Yorker, very much buttoned up into his blue pea jacket, and
with his watch cap well pulled down upon his ears.

The Burst of a Wave

“Ahoy, John—thanks for coming in. What’s it like outside?”
“It’s snowing. Winter’s come, I guess,” he said, with a meditative
grin.
We talked, I gave him the letter, and he went out into the dark
break of dawn, the wind, and the snow.
My fire had gone out, the Fo’castle was raw and cold, but my
wood was ready, and I soon had a fire crackling. All winter long, I
kept a basket of little sticks and fragments of driftwood ready for the
morning, and began the day with a bonfire in the fireplace. A
hearthful of high, leaping flame sends out a quick and plentiful heat.
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