Human Physiology

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HUMAN PHYSIOLOGY, FOURTEENTH EDITION
Published by McGraw-Hill Education, 2 Penn Plaza, New York, NY 10121. Copyright © 2016
by McGraw-Hill Education. All rights reserved. Printed in the United States of America. Previous
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outside the United States.
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ISBN 978-0-07-783637-5
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Library of Congress Cataloging-in-Publication Data
Fox, Stuart Ira.
Human physiology/Stuart Ira Fox, Pierce College.—Fourteenth edition.
pages cm
Includes index.
ISBN 978-0-07-783637-5 (alk. paper)
1. Human physiology—Textbooks. I. Title.
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Brief Contents

1 The Study of Body Function 1 13 Blood, Heart, and Circulation 404

2 Chemical Composition of the Body 24 14 Cardiac Output, Blood Flow, and Blood
Pressure 450
3 Cell Structure and Genetic Control 50
15 The Immune System 493
4 Enzymes and Energy 88
16 Respiratory Physiology 532
5 Cell Respiration and Metabolism 106
17 Physiology of the Kidneys 581
6 Interactions Between Cells and the
Extracellular Environment 130 18 The Digestive System 619

7 The Nervous System 162 19 Regulation of Metabolism 661

8 The Central Nervous System 206 20 Reproduction 701

9 The Autonomic Nervous System 243 Appendix

Answers to Objective Questions A-1

10 Sensory Physiology 266 Glossary G-1

11 Endocrine Glands 316 Credits C-1

Index I-1
12 Muscle 359

iii
About the Author

Stuart Ira Fox earned a Ph.D. in human physiology

from the Department of Physiology, School of Medicine, at
the University of Southern California, after earning degrees
at the University of California at Los Angeles (UCLA);
California State University, Los Angeles; and UC Santa
Barbara. He has spent most of his professional life teach-
ing at Los Angeles City College; California State University,
Northridge; and Pierce College, where he has won numer-
ous teaching awards, including several Golden Apples.
Stuart has authored thirty-nine editions of seven textbooks,
which are used worldwide and have been translated into sev-
eral languages, and two novels. When not engaged in profes-
sional activities, he likes to hike, fly fish, and cross-country
ski in the Eastern Sierra Nevada Mountains.

I wrote the first edition of Human Physiology to provide

my students with a readable textbook to support the
lecture material and help them understand physiology
concepts they would need later in their health curricula
and professions. This approach turned out to have wide
appeal, which afforded me the opportunity to refine
and update the text with each new edition. Writing
new editions is a challenging educational experience,
and an activity I find immensely enjoyable. Although To my wife, Ellen;
changes have occurred in the scientific understanding and to Laura, Eric, Kayleigh, and Jacob Van Gilder; for
and applications of physiological concepts, the students all the important reasons.
using this fourteenth edition have the same needs as
those who used the first, and so my writing goals have
remained the same. I am thankful for the privilege of
being able to serve students and their instructors through
these fourteen editions of Human Physiology.

—Stuart Ira Fox

iv
Preface

The Cover to human health and physical performance. The scope of top-
ics included in a human physiology course is therefore wide-
William B. Westwood’s cover ranging, yet each topic must be covered in sufficient detail to
illustration of the eye and the provide a firm basis for future expansion and application.
structures and processes required Human Physiology, fourteenth edition, is written for the
for vision encompasses the study undergraduate introductory human physiology course. Based
of physiology at multiple levels. on the author’s extensive experience with teaching this course,
The physiology of vision entails the framework of the textbook is designed to provide basic
the biophysical processes of light biology and chemistry (chapters 2–5) before delving into more
becoming focused onto and inter- complex physiological processes. This approach is appreciated
acting with photoreceptors, the by both instructors and students; specific references in later
molecular and cellular constitu- chapters direct readers back to the foundational material as
ents of these receptors that enable needed, presenting a self-contained study of human physiology.
them to respond to light, and neural interactions needed for the In addition to not presupposing student’s preparedness, this
brain to meaningfully interpret this stimulation. popular textbook is known for its clear and approachable writing
Photoreceptors are located in the part of the eye and brain style, detailed realistic art, and unsurpassed clinical information.
called the retina, which is a neural layer at the back of the eye.
The front cover shows light entering the eye and becoming Acknowledgments
focused by the lens onto the retina. The outer segments of pho-
toreceptors contain stacks of membranes, shown as purple at Reviewers
the bottom of the book’s spine, which contain the photorecep-
tor pigment rhodopsin (the green structures within the mem- Patti Allen, Dixie State College
branes at the bottom left of the front cover). Dani Behonick, Canada College
The bottom middle of the front cover illustrates a plasma Justin Brown, James Madison University
membrane of a photoreceptor neuron containing ion channels Michael Burg, San Diego City College
(pink). In the dark, these channels allow Na1 ions (pink spheres)
Julia Chang, Mount St. Mary’s College Chalon
to enter the photoreceptor. Light induces a change in the rhodop-
sin that initiates a signaling pathway (not shown), which leads to Corey Cleland, James Madison University
the closing of these channels (shown by the bottom channel). This Linda Collins, University of Tennessee Chattanooga
indirectly causes the photoreceptors to stimulate other neurons in Maria Elena DeBellard, California State University–Northridge
the retina (bipolar cells, depicted in red near the bottom of the Andrew Flick, James Madison University
front cover), which then stimulate another layer of neurons (gan-
James Hoffmann, Diablo Valley College
glion cells, depicted green at the bottom of the front cover.).
The axons (nerve fibers) of the ganglion cells gather together Cynthia Kay-Nishiyama, California State University–Northridge
to form the optic nerves, which leave the eye to carry visual infor- Paul Kingston, San Diego City College
mation to the brain, as shown on the back cover. The visual fields Arnold Kondo, Citrus College
illustrated as blue and purple circles on the back cover stimulate Ann Maliszewski, Cuesta College
different regions of the retina. Because many of the axons in the
Nancy Mann, Cuesta College
optic nerves cross to the opposite side, aspects of the right visual
field are conveyed to the left cerebral cortex and vice versa, as Tim Maze, Lander University
illustrated by the blue and purple colors of the nerve tracts. Physi- Vikki Mccleary, University of North Dakota
ological processes continue within the brain, allowing it to create Cheryl Neudauer, Minneapolis Community & Technical College
images that our mind interprets as the reality of the external world. Mark Paternostro, West Virginia University–Morgantown
Erik Schweitzer, Santa Monica Community College
What Sets This Book Apart? Laura Steele, Ivy Tech Community College of Indiana–Fort Wayne
The study of human physiology provides the scientific founda- R. Douglas Watson, University of Alabama at Birmingham
tion for the field of medicine and all other professions related Allison Wilson, Benedictine University

v
GUIDED TOUR
WHAT MAKES THIS TEXT A MARKET LEADER?
Clinical Applications—No Other Human Physiology Text Has More!
The framework of this textbook is based on integrating clinically germane information with knowledge of the body’s
physiological processes. Examples of this abound throughout the book. For example, in a clinical setting we record
electrical activity from the body: this includes action potentials (chapter 7, section 7.2); EEG (chapter 8, section 8.2); and
ECG (chapter 13, section 13.5). We also record mechanical force in muscle contractions (chapter 12, section 12.3). We
note blood plasma measurements of many chemicals to assess internal body conditions. These include measurements of
blood glucose (chapter 1, section 1.2) and the oral glucose tolerance test (chapter 19, section 19.4); and measurements of
the blood cholesterol profile (chapter 13, section 13.7). These are just a few of many examples the author includes that
focus on the connections between the study of physiology and our health industry.

NEW CLINICAL INVESTIGATIONS IN

ALL CHAPTERS!

Clinical Investigation ◀ Chapter-Opening Clinical Investigations, Clues,

and Summaries are diagnostic case studies found in
Sheryl, an active 78-year-old, suddenly became greatly
fatigued and disoriented while skiing. When she was each chapter. Clues are given throughout and the case is
brought to the hospital, blood tests revealed elevated finally resolved at the end of the chapter.
levels of LDH, AST, ALT, and the MB isoform of CK.
Some of the new terms and concepts you will encoun-
ter include:
• Enzymes, isoenzymes, coenzymes, and cofactors
Clinical Investigation SUMMARY
• LDH, AST, ALT, and CK The sudden onset of Sheryl’s great fatigue and disori-
entation is cause for concern and warranted immediate
enta
Clinical Investigation CLUES medical attention. Examination of table 4.1 with refer-
med
ence to the disorders indicated by elevated levels of
Sheryl’s blood tests reveal elevated levels of CPK, LDH, enc
AST, and ALT. CK, LDH, AST, and ALT reveal that they share one pos-
CK
sible cause in common—myocardial infarction (heart
sibl
• What enzymes do these letters indicate, and what attack). This possibility is reinforced by the laboratory
atta
diseases do elevated blood levels of these enzymes test
tests demonstrating that she had elevated levels of the
suggest? CK-MB isoenzyme, which is released by damaged heart
CK-
• How might these test results relate to Sheryl’s cells, rather than the CK-BB or CK-MM isoenzymes. A
cell
symptoms? possible myocardial infarction could explain Sheryl’s
pos
sudden onset of symptom while performing the intense
sud
exercise of skiing.
exe
▶ Clinical Investigations are enhanced with even See additional chapter 4 Clinical Investigation on Enzyme
Tests to Diagnose Diseases in the Connect site for this text.
more clinical assessments available on McGraw-Hill
Connect®. These Clinical Investigations are written
by the author and are specific to each chapter. They
will offer the students great insight into that specific
chapter.

fox36375_ch04_088-105.indd 91 1/5/15 3:22 PM

vi
 ALL APPLICATION BOXES ARE
NEW OR UPDATED! C L I N I C A L A P P L I C AT I O N
When diseases damage tissues, some cells die and release
▶ Clinical Application Boxes are in-depth boxed their enzymes into the blood. The activity of these enzymes,
essays that explore relevant topics of clinical interest reflecting their concentrations in the blood plasma, can be
and are placed at key points in the chapter to support measured in a test tube by adding their specific substrates.
the surrounding material. Subjects covered include Because an increase in certain enzymes in the blood can
pathologies, current research, pharmacology, and a indicate damage to specific organs, such tests may aid the
variety of clinical diseases. diagnosis of diseases. An increase in a man’s blood levels
of the acid, phosphatase, for example, may result from dis-
ease of the prostate (table 4.1).

F I T N E S S A P P L I C AT I O N
Metabolic syndrome is a combination of abnormal mea- ◀ Fitness Application Boxes are readings that explore
surements—including central obesity (excess abdominal physiological principles as applied to well-being, sports
fat), hypertension (high blood pressure), insulin resistance medicine, exercise physiology, and aging. They are also
(prediabetes), type 2 diabetes mellitus, high plasma triglyc-
placed at relevant points in the text to highlight concepts
erides, and high LDL cholesterol—that greatly increase the
risk of coronary heart disease, stroke, diabetes mellitus, and
just covered in the chapter.
other conditions. The incidence of metabolic syndrome has
increased alarmingly in recent years because of the increase
in obesity. Eating excessive calories, particularly in the form
of sugars (including high fructose corn syrup), stimulates
insulin secretion. Insulin then promotes the uptake of blood
glucose into adipose cells, where (through lipogenesis) it is
converted into stored triglycerides (see figs. 5.12 and 5.13).
Conversely, the lowering of insulin secretion, by diets that
prevent the plasma glucose from rising sharply, promotes
lipolysis (the breakdown of fat) and weight loss.

LEARNING OUTCOMES
▶ Learning Outcomes are numbered for easy After studying this section, you should be able to:
referencing in digital material! 2. Describe the aerobic cell respiration of glucose
fox36375_ch05_106-129.indd 120 12/30/14 9:01 PM
through the citric acid cycle.
3. Describe the electron transport system and oxidative
phosphorylation, explaining the role of oxygen in this
process.

fox36375_ch04_088-105.indd 91

| CHECKPOINT
▶ Learning Outcome numbers are tied 2a. Compare the fate of pyruvate in aerobic and
directly to Checkpoint numbers! anaerobic cell respiration.
2b. Draw a simplified citric acid cycle and indicate the
high-energy products.
3a. Explain how NADH and FADH2 contribute to
oxidative phosphorylation.
3b. Explain how ATP is produced in oxidative
phosphorylation.

fox36375_ch05_106-129.indd 111

vii

fox36375_ch05_106-129.indd 116 12/30/14 9:01 PM

GUIDED TOUR
WHAT MAKES THIS TEXT A MARKET LEADER?
Writing Style—Easygoing, Logical, and Concise
The words in Human Physiology, fourteenth edition, read as if the author is explaining concepts to you in a one-on-one
conversation, pausing now and then to check and make sure you understand what he is saying. Each major section begins with
a short overview of the information to follow. Numerous comparisons (“Unlike the life of an organism, which can be viewed
as a linear progression from birth to death, the life of a cell follows a cyclical pattern”), examples (“A callus on the hand, for
example, involves thickening of the skin by hyperplasia due to frequent abrasion”), reminders (“Recall that each member of
a homologous pair came from a different parent”), and analogies (“In addition to this ‘shuffling of the deck’ of chromosomes
. . .”) lend the author’s style a comfortable grace that enables readers to easily flow from one topic to the next.

Exceptional Art—Designed from the Outer mitochondrial

membrane

Student’s Point of View Inner mitochondrial

membrane

What better way to support such unparalleled writing

H+
than with high-quality art? Large, bright illustrations H+ 2

Intermembrane
demonstrate the physiological processes of the human body space
Third
Second pump
beautifully in a variety of ways. H+ pump

1
2 H+
► Stepped-out art clearly depicts various First pump
H 2O ATP
synthase

stages or movements with numbered 4 H+ 2 H + 1/2 O2 3

e–
explanations. 1 ADP
+
Pi H+ ATP

4 H+ NAD+

Matrix
NADH

Nucleus
Nucleus Basement
membrane
Basement Nucleus
Connective
membrane tissue
Basement
membrane Goblet cell
Connective
tissue ◀ Labeled photos placed side by side
with illustrations allow diagrammatic
detail and realistic application.

(a) (b) (c)

Muscle fiber nucleus

Nerve fiber branches
Motor end plate

► Macro-to-micro art helps Myofibril

students put context around

Mitochondria
detailed concepts. Folded sarcolemma

Synaptic vesicles

Neuromuscular cleft

Motor end plate

(a)

viii
FOURTEENTH EDITION Chapter 5: Cell Respiration and Metabolism
■ Updated description of the respiratory assemblies and their

CHANGES functions.
■ New discussion of inherited mitochondrial diseases.

What’s New? ■ Updated discussion of metabolic syndrome.

■ Updated and expanded discussion of brown fat.
Human Physiology, fourteenth edition, incorporates a number
of new and recently modified physiological concepts. This may Chapter 6: Interactions Between Cells and the
surprise people who are unfamiliar with the subject; indeed, Extracellular Environment
the author sometimes is asked if the field really changes much ■ New figure 6.22b.
■ Updated discussion of dialysis and hemodialysis.
from one edition to the next. It does; that’s one of the reasons
physiology is so much fun to study. Stuart has tried to impart Chapter 7: The Nervous System: Neurons and
this sense of excitement and fun in the book by indicating, in a Synapses
manner appropriate for this level of student, where knowledge ■ Updated and expanded discussions of microglia, axon
is new and where gaps in our knowledge remain. regeneration, neurotrophins, astrocytes, and of microglia.
The list that follows indicates only the larger areas of text ■ Discussion of the structure and function of gap junctions updated
and figure revisions and updates. It doesn’t indicate instances and expanded.
where passages were rewritten to improve the clarity or accu- ■ Figure 7.23 updated and revised.
racy of the existing material, or smaller changes made in ■ Explanation of synaptic vesicle docking and exocytosis updated
response to information from recently published journals and and expanded.
from the reviewers of the previous edition. ■ Expanded Table 7.4.
■ New discussion of different subtypes of muscarinic ACh
GLOBAL CHANGES: receptors.
■ Each Clinical Investigation in every chapter of the textbook is ■ Updated and expanded discussion of dopamine receptors and new
new. discussion of atypical antipsychotic drugs.
■ Each of the Clinical Investigation Clues, in every chapter, is new. ■ Updated discussion of inhibitory neurotransmitters.
■ The Clinical Investigation Summaries at the ends of all chapters ■ Expanded discussion of endocannabinoid neurotransmitters.
are new. ■ New discussion of hydrogen sulfide as a neurotransmitter.
■ Every Clinical Application box, in each and every chapter, has
been rewritten and updated. Chapter 8: The Central Nervous System
■ Every Fitness Application box, in each and every chapter, has ■ New photos in figures 8.9, 8.17, and 8.18.

been rewritten and updated. ■ Updated and expanded discussion of CSF formation and

circulation.
MAJOR CHANGES IN CHAPTERS ■ Updated discussion of neurogenesis in the adult brain.

These are specific changes made in the individual chapters in ■ Updated discussion of the origin of the electroencephalogram.

addition to the global changes described above. ■ New discussion of transient ischemic attack and stroke.
■ Updated description of brain areas involved in memory storage.
Chapter 1: The Study of Body Function ■ Updated and expanded discussion of Alzheimer’s disease.
■ Discussions of exfoliative cytology and Pap smear added.
■ Updated and expanded discussion of the molecular mechanisms
■ Discussions of embryonic stem cells, totipotency, and
involved in memory formation.
pluripotency added. ■ Updated and expanded discussion of the roles of dendritic spines

Chapter 3: Cell Structure and Genetic Control and neurogenesis in memory formation.
■ Updated discussion of the regulation of circadian rhythms.
■ New figures 3.3, 3.4, 3.7, 3.9a, and 3.18.
■ Updated discussion of the role of the nucleus accumbens in the
■ Descriptions of microtubules and autophagosomes updated.
■ Updated discussion of mitochondria, including hereditary reward pathway.
■ Updated discussion of orexin and new discussion of hypnotic drugs.
mitochondrial diseases.
■ Updated and expanded discussion of the agranular endoplasmic
Chapter 9: The Autonomic Nervous System
reticulum and drug tolerance. ■ New discussion of b3-adrenergic receptors added.
■ Updated and expanded discussion of genes, including new

description of retrotransposons. Chapter 10: Sensory Physiology

■ Updated discussion of microRNA and new description of circular ■ New figures 10.10 and 10.14a.

RNA. ■ Updated and expanded discussions of nociceptors, afferent fiber

■ Updated discussion of the medical uses of RNA interference. categories, and spinal cord lamina.
■ Updated discussion of epigenetic regulation and its significance. ■ Discussion of salty taste updated.

ix
■ Updated and expanded discussion of olfactory processing. Chapter 14: Cardiac Output, Blood Flow, and Blood
■ Discussion of the structure and function of the cochlea updated Pressure
and expanded. ■ New comparison of the pulmonary and systemic circulations.
■ New discussion of the role of microsaccades in vision. ■ Updated discussion of the effects of sympathetic and
■ New discussion of direction sensitive ganglion cells in vision. parasympathetic nerves on the cardiac rate.
■ Expanded discussion on the resting cardiac rate.
Chapter 11: Endocrine Glands: Secretion and Action
■ New discussion of the Anrep effect.
of Hormones
■ New discussion of neurovascular coupling and functional
■ New photos in figures 11.24 and 11.26.
■ Updated and expanded discussion of the different drugs used to
hyperemia.
■ New goals for the treatment of hypertension discussed.
treat breast cancer.
■ Updated discussion of the mechanisms responsible for
■ Updated and expanded discussion of insulin receptor structure

and function. hypertension.

■ Updated discussion of the role of dietary salt in hypertension.
■ Revised rendering of insulin receptor in figure 11.11.
■ Updated and expanded discussion of anterior pituitary cells and
Chapter 15: The Immune System
the hormones they produce.
■ Updated and expanded discussion of epithelial membranes and
■ Updated and expanded discussion of stress and glucocorticoid effects.
■ Updated discussions of calcitonin and the pancreatic islets.
immunity.
■ New discussion of NOD-like receptors and immunity.
■ New discussion of adipokines and myokines.
■ Updated and expanded discussion of opsonization and phagocytosis.

Chapter 12: Muscle: Mechanisms of Contraction and ■ Updated discussions of interferons and of secondary lymphoid

Neural Control organs.

■ Expanded discussion of motor end plates and new explanation of ■ Updated discussion of the effects of mast cell cytokines in local

end plate potential. inflammation.

■ New figure 12.9a. ■ Updated discussion of the roles of resident macrophages and

■ New discussion of the SERCA pumps in muscle contraction and neutrophils in an inflammation.
■ New figure 15.9.
relaxation.
■ New discussion of muscle glycogen and exercise. ■ Updated discussions of helper and regulatory T cells and antigen-

■ Updated discussion of muscle metabolism of fat during exercise. presenting cells.

■ New discussion of myokines and irisin. ■ Updated discussion of MHC class-1 and class-2 molecules.

■ Updated and expanded discussion of satellite cells in muscle ■ Updated discussion of immune response to viral infections.
■ Figures 15.15, 15.17, and 15.18 revised.
regeneration and sarcopenia.
■ Updated and expanded discussion of calcium-induced calcium ■ Updated and expanded discussions of memory T cells and of

release in cardiac muscle. adjuvants.

■ New discussion of calcium puffs and sparks in smooth muscle ■ New discussion of intravenous immunoglobulin.
■ New discussion of humanized monoclonal antibodies and
contraction.
■ New discussion of myosin light-chain phosphatase in smooth adoptive cell transfer.
■ New discussion of natural killer T cells.
muscle relaxation.
■ Updated discussion of autoimmune and allergic reactions.
Chapter 13: Blood, Heart, and Circulation ■ Updated and expanded discussion of contact dermatitis.
■ New discussion of the dietary need for iron in erythropoiesis.
■ Updated discussions of hepcidin and the intrinsic clotting pathway. Chapter 16: Respiratory Physiology
■ Updated discussion of the role of platelets in blood clotting and ■ Updated description of alveoli structure and function.

the use of warfarin to inhibit blood clotting. ■ New figures 16.3 and 16.5.
■ Updated and expanded discussion of the origin of the pacemaker ■ Revised discussion of surfactant and respiratory distress

potential. syndrome.
■ New discussion of sinoatrial conduction pathways and ectopic foci. ■ Updated and expanded discussion of the function of the
■ Updated discussion of calcium pumping in the regulation of the diaphragm in ventilation.
heartbeat. ■ Updated discussions of asthma and of the pulmonary capillaries.
■ New figure 13.31. ■ Updated and expanded discussion of the mechanisms of
■ Updated discussion of atherosclerosis. ventilation/perfusion matching.
■ Updated discussion of myocardial infarction and diet. ■ Revised discussion of pulmonary hypertension and cor
■ Updated and expanded discussion of blood tests to detect pulmonale.
myocardial infarction. ■ Updated and expanded discussion of the central regulation of
■ New discussion of interstitial fluid and the extracellular matrix. breathing.

x
■ Updated and expanded discussions of the carotid bodies and the ■ Updated discussion of the formation of the superoxide radical.
central chemoreceptors. ■ Updated discussions of adipocyte turnover, and adipose tissue in
■ Updated discussion of the role of nitric oxide in acclimatization starvation and obesity.
to high altitude. ■ Discussion of weight-loss medications updated.
■ Updated and expanded discussion of hypothalamic neurons and
Chapter 17: Physiology of the Kidneys neurotransmitters involved in the regulation of eating.
■ Updated discussion of glomerular structure and function. ■ Updated discussion of leptin and its regulation of appetite.
■ New figure 17.9. ■ New discussion of beige (or brite) adipocytes.
■ Updated discussion of the renal tubule transport of sodium and ■ Updated discussion of the mechanisms of beta cell insulin
chloride. secretion.
■ Revised discussion of the countercurrent multiplier system. ■ Updated discussion of how autonomic nerves and somatostatin
■ Updated discussion of urea transporters and aquaporin channels
regulate insulin secretion.
in the vasa recta. ■ Updated and expanded discussions of type 1 and type 2 diabetes
■ Updated discussion of countercurrent exchange in the renal medulla.
and their treatments.
■ Updated and expanded discussion of the role of urea in ■ New discussion on the roles of ectopic fat and visceral obesity in
concentrating the urine. impaired glucose tolerance and type 2 diabetes.
■ New discussion of arginine vasopressin as the antidiuretic ■ New discussion of soluble and insoluble fiber and its affect on
hormone, and updated discussion of its secretion. insulin resistance.
■ Revised organization of the sections on renal plasma clearance. ■ Updated discussion of dwarfism and new discussion of
■ Updated discussion of renal tubule potassium secretion.
achondroplasia.
■ Updated discussion of the roles of kidney-generated angiotensin II. ■ Updated discussion of the regulation of osteoclast formation.
■ New discussion of B-type natriuretic peptide. ■ New discussion of articular cartilage regeneration.
■ Updated discussion of ammonia produced by the renal tubules. ■ Discussion of calcitonin updated.
Chapter 18: The Digestive System
■ New discussion of osteocalcin and updated discussion of leptin
■ Revised figure 18.7 and new fig. 18.11.
actions on bone.
■ Updated discussion of the lower esophageal sphincter.
■ Updated and expanded discussion of intestinal calcium
■ New discussion of parietal cells and potassium recycling.
absorption and the actions of vitamin D.
■ Updated discussion of Paneth cells and intestinal stem cells.
■ Updated discussion of the actions of parathyroid hormone on
■ Updated and expanded discussion of the enteric nervous system.
renal phosphate excretion.
■ Updated discussion of intestinal slow waves and action
Chapter 20: Reproduction
potentials.
■ New figures 20.3, 20.40, and 20.42c.
■ Updated and expanded discussion of the origin and function of
■ Updated discussion of X chromosome inactivation and SRY.
the intestinal microbiota.
■ New discussion of kisspeptins and the regulation of GnRH
■ Updated and expanded discussion of the antimicrobial properties

of the intestinal mucosa. secretion.

■ Updated discussion of DHT and estradiol in male physiology.
■ New discussion of the gut-associated lymphoid tissue.
■ Updated discussion of spermatogenesis and the blood-testis
■ New discussions of Clostridium difficile infections and fecal

microbiota transplantation. barrier.

■ Updated and expanded discussions of the mechanisms of penile
■ Updated discussions of liver fibrosis and cirrhosis.
■ Updated and expanded discussion of transport processes in the
erection and of male contraception.
■ Updated and expanded discussion of ovarian follicle hormone
pancreatic acini.
■ New discussion of the function of somatostatin secreted by the D
production and its regulation.
■ Updated and expanded discussion of female contraception.
cells of the pancreatic islets.
■ Updated and expanded discussion of sperm capacitation and
■ New discussion of incretins in the regulation of insulin secretion.
■ Updated discussion of CCK in the regulation of pancreatic juice
hyperactivation.
■ New discussion of CatSper channels in sperm.
secretion.
■ Updated discussion of fertilization.
■ Updated discussion of secretin action.
■ Updated and expanded discussion of cloning and pluripotency.
■ Updated discussions of fat transport and fatty acid uptake.
■ Updated discussion of stem cells in regenerative medicine.
Chapter 19: Regulation of Metabolism ■ Updated discussion of adult stem cells and transdifferentiation.
■ New figures 19.17 and 19.20a. ■ Updated and expanded discussion of the pituitary-like hormones
■ New discussion of hypothermia and hypothermic circulatory secreted by the placenta.
arrest. ■ Table 20.7 updated and expanded.

xi
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xii
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have requested and we are providing five Physiology: Concepts and Clinical
new exercises (Respiratory Quotient, Applications, 13th edition, contains
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xiv
Contents
Preface v

C H AP TE R 1 2.4 Nucleic Acids 44

Deoxyribonucleic Acid 44
The Study of Body Function 1
Ribonucleic Acid 46
1.1 Introduction to Physiology 2
Summary 47
Scientific Method 2
Review Activities 48
1.2 Homeostasis and Feedback Control 4
History of Physiology 4
Negative Feedback Loops 6 CHAPTER 3
Positive Feedback 8
Cell Structure and Genetic Control 50
Neural and Endocrine Regulation 8
3.1 Plasma Membrane and Associated
Feedback Control of Hormone Secretion 9
Structures 51
1.3 The Primary Tissues 10
Structure of the Plasma Membrane 52
Muscle Tissue 11
Phagocytosis 54
Nervous Tissue 12
Endocytosis 54
Epithelial Tissue 12
Exocytosis 55
Connective Tissue 16
Cilia and Flagella 55
1.4 Organs and Systems 18
Microvilli 56
An Example of an Organ: The Skin 18
3.2 Cytoplasm and Its Organelles 56
Systems 20
Cytoplasm and Cytoskeleton 57
Body-Fluid Compartments 20
Lysosomes 58
Summary 21
Peroxisomes 58
Review Activities 22
Mitochondria 59
Ribosomes 60

C H AP TE R 2 Endoplasmic Reticulum 60
Golgi Complex 61
Chemical Composition of the Body 24 3.3 Cell Nucleus and Gene Expression 62
2.1 Atoms, Ions, and Chemical Bonds 25 Genome and Proteome 63
Atoms 25 Chromatin 63
Chemical Bonds, Molecules, and Ionic RNA Synthesis 64
Compounds 26 RNA Interference 67
Acids, Bases, and the pH Scale 29 3.4 Protein Synthesis and Secretion 68
Organic Molecules 30 Transfer RNA 68
2.2 Carbohydrates and Lipids 33 Formation of a Polypeptide 69
Carbohydrates 33 Functions of the Endoplasmic Reticulum
Lipids 36 and Golgi Complex 70
2.3 Proteins 41 Protein Degradation 70
Structure of Proteins 41 3.5 DNA Synthesis and Cell Division 72
Functions of Proteins 44 DNA Replication 72

xv
xvi Contents

The Cell Cycle 74 5.4 Metabolism of Lipids and Proteins 119

Mitosis 76 Lipid Metabolism 119
Meiosis 78 Amino Acid Metabolism 122
Epigenetic Inheritance 79 Uses of Different Energy Sources 123
Interactions 83 Interactions 126
Summary 84 Summary 127
Review Activities 85 Review Activities 128

C H AP T ER 4 CHAPTER 6
Enzymes and Energy 88 Interactions Between Cells and the
Extracellular Environment 130
4.1 Enzymes as Catalysts 89
6.1 Extracellular Environment 131
Mechanism of Enzyme Action 89
Body Fluids 131
Naming of Enzymes 91
Extracellular Matrix 132
4.2 Control of Enzyme Activity 92
Categories of Transport Across the Plasma
Effects of Temperature and pH 92
Membrane 132
Cofactors and Coenzymes 93
6.2 Diffusion and Osmosis 133
Enzyme Activation 94
Diffusion Through the Plasma Membrane 135
Substrate Concentration and Reversible
Rate of Diffusion 136
Reactions 94
Osmosis 136
Metabolic Pathways 95
Regulation of Blood Osmolality 141
4.3 Bioenergetics 97
6.3 Carrier-Mediated Transport 142
Endergonic and Exergonic Reactions 98
Facilitated Diffusion 143
Coupled Reactions: ATP 98
Active Transport 144
Coupled Reactions: Oxidation-Reduction 99
Bulk Transport 148
Summary 102
6.4 The Membrane Potential 149
Review Activities 104
Equilibrium Potentials 150
Resting Membrane Potential 152
C H AP T ER 5 6.5 Cell Signaling 153
Second Messengers 155
Cell Respiration and Metabolism 106
G-Proteins 155
5.1 Glycolysis and the Lactic Acid Pathway 107 Interactions 157
Glycolysis 107 Summary 158
Lactic Acid Pathway 109 Review Activities 159
5.2 Aerobic Respiration 111
Citric Acid Cycle 111
Electron Transport and Oxidative CHAPTER 7
Phosphorylation 112 The Nervous System 162
Coupling of Electron Transport to ATP
Production 113 7.1 Neurons and Supporting Cells 163
ATP Balance Sheet 115 Neurons 163
5.3 Interconversion of Glucose, Lactic Acid, Classification of Neurons and Nerves 165
and Glycogen 117 Neuroglial Cells 166
Glycogenesis and Glycogenolysis 117 Neurilemma and Myelin Sheath 167
Cori Cycle 117 Functions of Astrocytes 170
 Contents xvii

7.2 Electrical Activity in Axons 172 Hindbrain 230

Ion Gating in Axons 173 Reticular Activating System in Sleep and
Action Potentials 174 Arousal 231
Conduction of Nerve Impulses 178 8.5 Spinal Cord Tracts 232
7.3 The Synapse 180 Ascending Tracts 233
Electrical Synapses: Gap Junctions 181 Descending Tracts 233
Chemical Synapses 181 8.6 Cranial and Spinal Nerves 236
7.4 Acetylcholine as a Neurotransmitter 184 Cranial Nerves 236
Chemically Regulated Channels 185 Spinal Nerves 236
Acetylcholinesterase (AChE) 189 Summary 239
Acetylcholine in the PNS 190 Review Activities 240
Acetylcholine in the CNS 190
7.5 Monoamines as Neurotransmitters 191
Serotonin as a Neurotransmitter 192 CHAPTER 9
Dopamine as a Neurotransmitter 192 The Autonomic Nervous System 243
Norepinephrine as a Neurotransmitter 194
9.1 Neural Control of Involuntary
7.6 Other Neurotransmitters 194 Effectors 244
Amino Acids as Neurotransmitters 194 Autonomic Neurons 244
Polypeptides as Neurotransmitters 196 Visceral Effector Organs 245
Endocannabinoids as Neurotransmitters 197 9.2 Divisions of the Autonomic Nervous
Gases as Neurotransmitters 198 System 246
ATP and Adenosine as Neurotransmitters 198 Sympathetic Division 246
7.7 Synaptic Integration 199 Parasympathetic Division 247
Synaptic Plasticity 199 9.3 Functions of the Autonomic Nervous
Synaptic Inhibition 200 System 251
Summary 201 Adrenergic and Cholinergic Synaptic
Review Activities 203 Transmission 251
Responses to Adrenergic Stimulation 252

C H AP TE R 8 Responses to Cholinergic Stimulation 256

Other Autonomic Neurotransmitters 257
The Central Nervous System 206 Organs with Dual Innervation 258
8.1 Structural Organization of the Brain 207 Organs Without Dual Innervation 260
8.2 Cerebrum 209 Control of the Autonomic Nervous System
by Higher Brain Centers 260
Cerebral Cortex 209
Interactions 262
Basal Nuclei 215
Summary 263
Cerebral Lateralization 216
Review Activities 264
Language 218
Limbic System and Emotion 219
Memory 220
Emotion and Memory 224
CHAPTER 10
8.3 Diencephalon 225 Sensory Physiology 266
Thalamus and Epithalamus 225 10.1 Characteristics of Sensory Receptors 267
Hypothalamus and Pituitary Gland 226 Categories of Sensory Receptors 267
8.4 Midbrain and Hindbrain 228 Law of Specific Nerve Energies 268
Midbrain 228 Generator (Receptor) Potential 269
xviii Contents

10.2 Cutaneous Sensations 270 Effects of Hormone Concentrations on Tissue

Neural Pathways for Somatesthetic Response 321
Sensations 271 11.2 Mechanisms of Hormone Action 323
Receptive Fields and Sensory Acuity 272 Hormones That Bind to Nuclear Receptor
Lateral Inhibition 273 Proteins 323
10.3 Taste and Smell 274 Hormones That Use Second Messengers 326
Taste 274 11.3 Pituitary Gland 331
Smell 276 Pituitary Hormones 331
10.4 Vestibular Apparatus and Equilibrium 278 Hypothalamic Control of the Posterior
Pituitary 333
Sensory Hair Cells of the Vestibular
Apparatus 279 Hypothalamic Control of the Anterior
Pituitary 333
Utricle and Saccule 280
Feedback Control of the Anterior Pituitary 335
Semicircular Canals 280
Higher Brain Function and Pituitary Secretion 336
10.5 The Ears and Hearing 282
11.4 Adrenal Glands 337
Outer Ear 283
Functions of the Adrenal Cortex 337
Middle Ear 283
Functions of the Adrenal Medulla 339
Cochlea 284
Stress and the Adrenal Gland 340
Spiral Organ (Organ of Corti) 286
11.5 Thyroid and Parathyroid Glands 341
10.6 The Eyes and Vision 290
Production and Action of Thyroid Hormones 342
Refraction 294
Parathyroid Glands 344
Accommodation 295
11.6 Pancreas and Other Endocrine Glands 345
Visual Acuity 296
Pancreatic Islets (Islets of Langerhans) 345
10.7 Retina 297
Pineal Gland 346
Effect of Light on the Rods 299
Gastrointestinal Tract 349
Electrical Activity of Retinal Cells 300
Gonads and Placenta 349
Cones and Color Vision 301
11.7 Paracrine and Autocrine Regulation 349
Visual Acuity and Sensitivity 304
Examples of Paracrine and Autocrine
Neural Pathways from the Retina 304
Regulation 350
10.8 Neural Processing of Visual Information 307
Prostaglandins 351
Ganglion Cell Receptive Fields 307
Interactions 354
Lateral Geniculate Nuclei 308
Summary 355
Cerebral Cortex 308
Review Activities 356
Interactions 310
Summary 311
Review Activities 314
CHAPTER 12
Muscle 359
C H AP T ER 11 12.1 Skeletal Muscles 360
Endocrine Glands 316 Structure of Skeletal Muscles 360
11.1 Endocrine Glands and Hormones 317 Motor End Plates and Motor Units 361
Chemical Classification of Hormones 318 12.2 Mechanisms of Contraction 364
Prohormones and Prehormones 320 Sliding Filament Theory of Contraction 367
Common Aspects of Neural and Endocrine Regulation of Contraction 369
Regulation 321 12.3 Contractions of Skeletal Muscles 374
Hormone Interactions 321 Twitch, Summation, and Tetanus 374
 Contents xix

Types of Muscle Contractions 375 13.5 Electrical Activity of the Heart and the
Series-Elastic Component 376 Electrocardiogram 425
Length-Tension Relationship 376 Electrical Activity of the Heart 425
12.4 Energy Requirements of Skeletal The Electrocardiogram 428
Muscles 377 13.6 Blood Vessels 431
Metabolism of Skeletal Muscles 378 Arteries 431
Slow- and Fast-Twitch Fibers 380 Capillaries 433
Muscle Fatigue 381 Veins 435
Adaptations of Muscles to Exercise Training 382 13.7 Atherosclerosis and Cardiac
Muscle Damage and Repair 384 Arrhythmias 436
12.5 Neural Control of Skeletal Muscles 384 Atherosclerosis 436
Muscle Spindle Apparatus 386 Arrhythmias Detected by the
Electrocardiograph 440
Alpha and Gamma Motoneurons 387
13.8 Lymphatic System 442
Coactivation of Alpha and Gamma
Motoneurons 387 Summary 445
Skeletal Muscle Reflexes 387 Review Activities 447
Upper Motor Neuron Control of Skeletal
Muscles 390
12.6 Cardiac and Smooth Muscles 391 CHAPTER 14
Cardiac Muscle 392
Cardiac Output, Blood Flow,
Smooth Muscle 393
and Blood Pressure 450
Interactions 398
14.1 Cardiac Output 451
Summary 399
Regulation of Cardiac Rate 451
Review Activities 401
Regulation of Stroke Volume 452
Venous Return 454

C H AP TE R 13 14.2 Blood Volume 456

Exchange of Fluid Between Capillaries and
Blood, Heart, and Circulation 404 Tissues 457
Regulation of Blood Volume by the
13.1 Functions and Components of the
Kidneys 459
Circulatory System 405
14.3 Vascular Resistance to Blood
Functions of the Circulatory System 405
Flow 463
Major Components of the Circulatory System 405
Physical Laws Describing Blood Flow 464
13.2 Composition of the Blood 406
Extrinsic Regulation of Blood Flow 465
Plasma 406
Paracrine Regulation of Blood Flow 466
The Formed Elements of Blood 407
Intrinsic Regulation of Blood Flow 467
Hematopoiesis 409
14.4 Blood Flow to the Heart and Skeletal
Red Blood Cell Antigens and Blood Typing 412 Muscles 468
Blood Clotting 414 Aerobic Requirements of the Heart 468
Dissolution of Clots 417 Regulation of Coronary Blood Flow 469
13.3 Structure of the Heart 418 Regulation of Blood Flow Through Skeletal
Pulmonary and Systemic Circulations 418 Muscles 470
Atrioventricular and Semilunar Valves 419 Circulatory Changes During Exercise 470
Heart Sounds 420 14.5 Blood Flow to the Brain and Skin 473
13.4 Cardiac Cycle 422 Cerebral Circulation 473
Pressure Changes During the Cardiac Cycle 423 Cutaneous Blood Flow 474
xx Contents

14.6 Blood Pressure 475

Baroreceptor Reflex 477 CHAPTER 16
Atrial Stretch Reflexes 479 Respiratory Physiology 532
Measurement of Blood Pressure 479 16.1 The Respiratory System 533
Pulse Pressure and Mean Arterial Pressure 481 Structure of the Respiratory System 533
14.7 Hypertension, Shock, and Congestive Heart Thoracic Cavity 536
Failure 482
16.2 Physical Aspects of Ventilation 536
Hypertension 482
Intrapulmonary and Intrapleural Pressures 537
Circulatory Shock 484
Physical Properties of the Lungs 538
Congestive Heart Failure 486
Surfactant and Respiratory Distress
Interactions 488 Syndrome 540
Summary 489 16.3 Mechanics of Breathing 540
Review Activities 490 Inspiration and Expiration 541
Pulmonary Function Tests 542
C H AP T ER 15 Pulmonary Disorders 544
The Immune System 493 16.4 Gas Exchange in the Lungs 547
Calculation of PO 547
15.1 Defense Mechanisms 494 2

Partial Pressures of Gases in Blood 548

Innate (Nonspecific) Immunity 495
Significance of Blood PO and PCO
Adaptive (Specific) Immunity 497 2 2

Measurements 550
Lymphocytes and Lymphoid Organs 499
Pulmonary Circulation and Ventilation/Perfusion
Local Inflammation 500 Ratios 550
15.2 Functions of B Lymphocytes 503 Disorders Caused by High Partial Pressures of
Antibodies 504 Gases 552
The Complement System 506 16.5 Regulation of Breathing 553
15.3 Functions of T Lymphocytes 507 Brain Stem Respiratory Centers 553
Killer, Helper, and Regulatory Effects of Blood PCO and pH on
2

T Lymphocytes 507 Ventilation 555

Interactions Between Antigen-Presenting Cells and Effects of Blood PO on Ventilation 557
2

T Lymphocytes 511 Effects of Pulmonary Receptors on

15.4 Active and Passive Immunity 514 Ventilation 558
Active Immunity and the Clonal Selection 16.6 Hemoglobin and Oxygen Transport 559
Theory 515 Hemoglobin 559
Immunological Tolerance 517 The Oxyhemoglobin Dissociation Curve 561
Passive Immunity 518 Effect of pH and Temperature on Oxygen
15.5 Tumor Immunology 519 Transport 562
Natural Killer Cells 520 Effect of 2,3-DPG on Oxygen Transport 563
Effects of Aging and Stress 521 Inherited Defects in Hemoglobin Structure and
15.6 Diseases Caused by the Immune Function 564
System 521 Muscle Myoglobin 564
Autoimmunity 521 16.7 Carbon Dioxide Transport 565
Immune Complex Diseases 522 The Chloride Shift 566
Allergy 523 The Reverse Chloride Shift 566
Interactions 527 16.8 Acid-Base Balance of the Blood 567
Summary 528 Principles of Acid-Base Balance 568
Review Activities 529 Ventilation and Acid-Base Balance 569
 Contents xxi

16.9 Effect of Exercise and High Altitude on

Respiratory Function 570 CHAPTER 18
Ventilation During Exercise 570 The Digestive System 619
Acclimatization to High Altitude 571
18.1 Introduction to the Digestive System 620
Interactions 575
Layers of the Gastrointestinal Tract 621
Summary 576
Regulation of the Gastrointestinal Tract 622
Review Activities 578
18.2 From Mouth to Stomach 623
Esophagus 624
Stomach 625
C H AP TE R 17 Pepsin and Hydrochloric Acid Secretion 626
Physiology of the Kidneys 581 18.3 Small Intestine 628
Villi and Microvilli 629
17.1 Structure and Function of the
Kidneys 582 Intestinal Enzymes 630
Gross Structure of the Urinary System 582 Intestinal Contractions and Motility 631
Control of Micturition 584 18.4 Large Intestine 632
Microscopic Structure of the Kidney 585 Intestinal Microbiota 633
17.2 Glomerular Filtration 587 Fluid and Electrolyte Absorption in the
Intestine 635
Glomerular Ultrafiltrate 588
Defecation 636
Regulation of Glomerular Filtration Rate 588
18.5 Liver, Gallbladder, and Pancreas 636
17.3 Reabsorption of Salt and Water 590
Structure of the Liver 636
Reabsorption in the Proximal Tubule 590
Functions of the Liver 638
The Countercurrent Multiplier System 592
Gallbladder 641
Collecting Duct: Effect of Antidiuretic Hormone
(ADH) 595 Pancreas 643
17.4 Renal Plasma Clearance 598 18.6 Regulation of the Digestive System 645
Transport Process Affecting Renal Regulation of Gastric Function 645
Clearance 599 Regulation of Intestinal Function 648
Renal Clearance of Inulin: Measurement of Regulation of Pancreatic Juice and Bile
GFR 600 Secretion 648
Renal Clearance Measurements 601 Trophic Effects of Gastrointestinal Hormones 650
Reabsorption of Glucose 603 18.7 Digestion and Absorption of Food 650
17.5 Renal Control of Electrolyte and Acid-Base Digestion and Absorption of Carbohydrates 650
Balance 604 Digestion and Absorption of Proteins 651
Role of Aldosterone in Na1/K1 Balance 604 Digestion and Absorption of Lipids 652
Control of Aldosterone Secretion 606 Interactions 656
Juxtaglomerular Apparatus 606 Summary 657
Natriuretic Peptides 607 Review Activities 658
Relationship Between Na1, K1, and H1 608
Renal Acid-Base Regulation 608
17.6 Diuretics and Renal Function Tests 611
Use of Diuretics 611
CHAPTER 19
Renal Function Tests and Kidney Disease 613 Regulation of Metabolism 661
Interactions 614 19.1 Nutritional Requirements 662
Summary 615 Metabolic Rate and Caloric Requirements 662
Review Activities 616 Anabolic Requirements 664
xxii Contents

Vitamins and Minerals 666 Pineal Gland 712

Free Radicals and Antioxidants 668 Human Sexual Response 712
19.2 Regulation of Energy Metabolism 669 20.3 Male Reproductive System 712
Regulatory Functions of Adipose Tissue 670 Control of Gonadotropin Secretion 713
Regulation of Hunger and Metabolic Rate 672 Endocrine Functions of the Testes 714
Caloric Expenditures 674 Spermatogenesis 715
Hormonal Regulation of Metabolism 675 Male Accessory Sex Organs 718
19.3 Energy Regulation by the Pancreatic Erection, Emission, and Ejaculation 719
Islets 677 Male Fertility 721
Regulation of Insulin and Glucagon Secretion 677 20.4 Female Reproductive System 722
Insulin and Glucagon: Absorptive State 679 Ovarian Cycle 724
Insulin and Glucagon: Postabsorptive State 679 Ovulation 725
19.4 Diabetes Mellitus and Hypoglycemia 681 Pituitary-Ovarian Axis 727
Type 1 Diabetes Mellitus 681 20.5 Menstrual Cycle 728
Type 2 Diabetes Mellitus 682 Phases of the Menstrual Cycle: Cyclic Changes in
Hypoglycemia 685 the Ovaries 728
19.5 Metabolic Regulation by Adrenal Hormones, Cyclic Changes in the Endometrium 731
Thyroxine, and Growth Hormone 686 Effects of Pheromones, Stress, and Body Fat 732
Adrenal Hormones 686 Contraceptive Methods 733
Thyroxine 686 Menopause 734
Growth Hormone 687 20.6 Fertilization, Pregnancy, and
19.6 Regulation of Calcium and Phosphate Parturition 734
Balance 690 Fertilization 735
Bone Deposition and Resorption 690 Cleavage and Blastocyst Formation 737
Hormonal Regulation of Bone 692 Implantation of the Blastocyst and Formation of
1,25-Dihydroxyvitamin D3 693 the Placenta 740
Negative Feedback Control of Calcium Exchange of Molecules Across the Placenta 742
and Phosphate Balance 695 Endocrine Functions of the Placenta 743
Summary 696 Labor and Parturition 744
Review Activities 698 Lactation 746
Interactions 749

C H AP T ER 20 Concluding Remarks 750

Summary 750
Reproduction 701 Review Activities 752
20.1 Sexual Reproduction 702
Sex Determination 702
Appendix
Development of Accessory Sex Organs and
External Genitalia 705 Answers to Test Your Knowledge Questions A-1
Disorders of Embryonic Sexual Development 706 Glossary G-1
20.2 Endocrine Regulation of Reproduction 708
Interactions Between the Hypothalamus, Pituitary Credits C-1
Gland, and Gonads 709
Onset of Puberty 710 Index I-1
C H A P TE R O UTLI N E
1.1 Introduction to Physiology 2
CHAPTER

1
Scientific Method 2
1.2 Homeostasis and Feedback Control 4
History of Physiology 4
Negative Feedback Loops 6
Positive Feedback 8
Neural and Endocrine Regulation 8
Feedback Control of Hormone Secretion 9
1.3 The Primary Tissues 10
Muscle Tissue 11
Nervous Tissue 12
Epithelial Tissue 12 The Study of
Connective Tissue 16
1.4 Organs and Systems 18
An Example of an Organ: The Skin 18
Body Function
Systems 20
Body-Fluid Compartments 20

Summary 21
Review Activities 22