Anatomy and physiology, the unity of form and function 6th ed k saladin (mcgraw hill, 2012) 1

Sixth Edition

&

The Unity of Form and Function

Kenneth S. Saladin
Georgia College & State University

TM

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TM

ANATOMY & PHYSIOLOGY: THE UNITY OF FORM AND FUNCTION, SIXTH EDITION
Published by McGraw-Hill, a business unit of The McGraw-Hill Companies, Inc., 1221 Avenue of the Americas, New York, NY
10020. Copyright © 2012 by The McGraw-Hill Companies, Inc. All rights reserved. Previous editions © 2010, 2007, and 2004. 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 The McGraw-Hill Companies, Inc., 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.
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ISBN 978–0–07–337825–1
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Library of Congress Cataloging-in-Publication Data
Saladin, Kenneth S.
Anatomy & physiology : the unity of form and function / Kenneth S. Saladin. -- 6th ed.
p. cm.
Includes index.
ISBN 978–0–07–337825–1 — ISBN 0–07–337825–9 (hard copy: alk. paper) 1. Human physiology. 2. Human anatomy. I. Title.
II. Title: Anatomy and physiology.
QP34.5.S23 2012
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2010042586
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BRIEF

About the Author iv
Preface v
Reviewers xxi
Contents xxii
Letter to the Students xxvi

16
17

Contents

Sense Organs 582
The Endocrine System 633

PART FOUR
Regulation and Maintenance

PART ONE
Organization of the Body
1

2
3
4

5

Major Themes of Anatomy and Physiology 1
Atlas A General Orientation to Human
Anatomy 28
The Chemistry of Life 42
Cellular Form and Function 78
Genetics and Cellular Function 114
Histology 143

18
19
20
21
22
23
24
25
26

The Circulatory System: Blood 678
The Circulatory System: The Heart 714
The Circulatory System: Blood Vessels
and Circulation 749
The Lymphatic and Immune Systems 808
The Respiratory System 854
The Urinary System 895
Water, Electrolyte, and Acid–Base Balance 930
The Digestive System 953
Nutrition and Metabolism 1000

PART TWO
Support and Movement
6
7
8
9
10
11

The Integumentary System 180
Bone Tissue 206
The Skeletal System 233
Joints 278
The Muscular System 312
Atlas B Regional and Surface Anatomy 379
Muscular Tissue 401

PART THREE
Integration and Control
12
13
14
15

sal78259_fm_i-xxvi.indd iii

Nervous Tissue 439
The Spinal Cord, Spinal Nerves, and
Somatic Reflexes 478

The Brain and Cranial Nerves 511
The Autonomic Nervous System and
Visceral Reflexes 561

PART FIVE
Reproduction and Development
27
28
29

The Male Reproductive System 1034
The Female Reproductive System 1064
Human Development and Aging 1102

Appendix A. Periodic Table A-1
Appendix B. Answer Keys A-2
Appendix C. Symbols of Weight and Measures A-13
Appendix D. Biomedical Abbreviations A-14
Glossary G-1
Credits C-1
Index I-1

11/19/10 9:53 AM


ABOUT THE

Author

KENNETH S. SALADIN has taught since 1977 at

Georgia College and State University in Milledgeville,
Georgia. He earned a B.S. in zoology at Michigan State
University and a Ph.D. in parasitology at Florida State
University, with interests especially in the sensory ecology of freshwater invertebrates. In addition to human
anatomy and physiology, his teaching experience includes
histology, parasitology, animal behavior, sociobiology,
introductory biology, general zoology, biological etymology, and study abroad in the Galápagos Islands. Ken has
been recognized as “most significant undergraduate mentor” nine times over the years by outstanding students
inducted into Phi Kappa Phi. He received the university’s
Excellence in Research and Publication Award for the
first edition of this book, and was named Distinguished
Professor in 2001.

Ken is a member of the Human Anatomy and
Physiology Society, the Society for Integrative and Comparative Biology, the American Association of Anatomists,
and the American Association for the Advancement of
Science. He served as a developmental reviewer and wrote
supplements for several other McGraw-Hill anatomy and
physiology textbooks for a number of years before becoming a textbook writer.
Ken’s outside interests include the Big Brothers/
Big Sisters program for single-parent children, the Charles
Darwin Research Station in the Galápagos, and student
scholarships. Ken is married to Diane Saladin, a registered nurse. They have two adult children.

This book is dedicated
to the memory of
H. Kenneth Hamill
and with gratitude to
Big Brothers–Big Sisters of Greater Kalamazoo
Big Brothers–Big Sisters of America

iv

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THE EVOLUTION OF A
Storyteller
Ken Saladin’s first step into authoring was a 318-page paper on the
ecology of hydras written for his 10th-grade biology class. With his
“first book,” featuring 53 original India ink drawings and
photomicrographs, a true storyteller was born.
“When I first became a textbook writer, I found myself bringing
the same enjoyment of writing and illustrating to this book that I
first discovered back when I was 15.”
–Ken Saladin

Ken's “first book,” Hydra
Ecology, 1965

One of Ken’s drawings
from Hydra Ecology

Ken in 1964

Ken began working on his first
book for McGraw-Hill in 1993, and in
1997 the first edition of The Unity of

Form and Function was published.
In 2011 the story continues with the
sixth edition of Ken’s best-selling A&P
textbook.

The first edition (1997)

The story continues (2011)

v

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SALADIN ANATOMY & PHYSIOLOGY

A Good Story

Anatomy & Physiology: The Unity of Form and Function tells a story
made of many layers including the core science, clinical applications, the
history of medicine, and the evolution of the human body. Saladin
combines this humanistic perspective on anatomy and physiology with
vibrant photos and art to convey the beauty and excitement of the subject
to beginning students.
To help students manage the
tremendous amount of information
in this introductory course, the
narrative is broken into short

segments, each framed by expected
learning outcomes and self-testing
review questions. This
presentation strategy works as a
whole to create a more efficient
and effective way for students to
learn A&P.

“Ken Saladin’s Anatomy &
Physiology: The Unity of Form
and Function, 6th edition, provides a fresh approach to the
study of A&P, with modern pedagogy, an abundance of ancillary learning resources, and the
most up-to-date information.
Instructors and students alike
will benefit from the Saladin
experience.”

Storytelling Writing Style viii–x
Appropriate Level
Interactive Material
Interesting Reading

Artwork That Encourages
Learning xi–xii
Sets the Standard
Conducive to Learning

Pedagogical Learning Tools xiii–xiv
Engaging Chapter Layouts
Tiered Assessments Based on Key

Lists of Expected Learning Outcomes

Innovative Chapter Sequencing xv
The Saladin Digital Story xvi-xix

–David Manry, Hillsborough
Community College

What’s New in the Sixth Edition?
New Atlas Organization
Many figures of regional anatomy (former figs. A.12–A.22) are moved
from atlas A to atlas B, now titled “Regional and Surface Anatomy.”
Beside shortening atlas A and moving the student more quickly to
chapter 2, this moves some anatomical detail to a later point where
students will be better equipped to understand it and relate it to
surface anatomy.

New Deeper Insight Essays
New essays introduce contemporary issues in health science and a
fascinating historical account that underscores some principles of
respiratory physiology.

It’s not unusual to hear textbook cynics say that new editions are just the same material
bound in new covers, but that
certainly isn’t true of this one.
Just listing my sixth-edition
changes came to 50 pages
and 18,000 words.
—Ken Saladin

• Trans fats and cardiovascular disease (Deeper Insight 2.3)
• Bone marrow and cord blood transplants (Deeper Insight 18.3)
• Altitude sickness and the Zenith ballooning tragedy (Deeper Insight 22.3)

vi

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New Science

New Art

Saladin’s Anatomy & Physiology, sixth edition, stays abreast
of key developments in science. Yet, more efficient writing and illustration result in a book slightly shorter than
the fifth edition even with these additions.

• Cis- and trans-fatty acids (fig. 2.20)

• Advances in tissue engineering (chapter 5)
• The stem-cell controversy and induced pluripotent
stem cells (chapter 5)
• Melanoma (chapter 6)
• Cola beverages and bone loss (chapter 7)
• Bases of muscle fatigue (chapter 11)
• Microglia and astrocyte functions (chapter 12)
• Neural mechanism of working memory (chapter 12)
• Hypothalamic control of hunger and satiety

(chapter 14)
• Orexins, sleep, and narcolepsy (chapter 14)
• Vascular pathogenesis in diabetes mellitus (chapter 17)
• Glycemic index of foods (chapter 26)
• Treatment of alcoholism (chapter 26)
• Vaccination against human papillomavirus (chapter 27)
• In vitro fertilization and the 2010 Nobel Prize
(chapter 29)

New Writing
Several sections have been rewritten for improved clarity,
especially:
• Carrier-mediated membrane transport (chapter 3)
• Genetic translation and ribosomal function (chapter 4)
• A better example of an anatomical second-class lever
(chapter 9)
• Muscle compartments and blood supply (chapter 10)
• Smooth muscle physiology (chapter 11)
• A view of saltatory conduction more accurate than
most textbook presentations (chapter 12)
• The adrenal cortex (chapter 17)
• Causes of arteriosclerosis and distinctions between
arteriosclerosis and atherosclerosis (chapter 20)

New Photographs

• Genetic translation (fig. 4.8)
• Types of cell junctions (fig. 5.28)
• Embryonic development of exocrine and endocrine
glands (fig. 5.29)

• Serous membrane histology (fig. 5.33b)
• The femur as a second-class lever (fig. 9.9b)
• The spinal reflex arc (fig. 13.21)
• Oxyhemoglobin dissociation curves (figs. 22.24
and 22.27)
• Connective Issues art and layouts

New Pedagogy
• Brushing Up is fleshed out and repositioned to better
catch the student’s attention and emphasize the
importance of understanding earlier material before
starting a new chapter.
• A list of Expected Learning Outcomes heads up each
chapter subdivision and exercises called Assess Your
Learning Outcomes end each chapter as a whole.
Instructors can now easily show how their courses are
outcome-driven.
• Apply What You Know questions, formerly called
Think About It, stress that these thought exercises are
analytical applications of basic anatomy and physiology knowledge to clinical situations and other new
contexts. Students can see how the basic anatomy and
physiology they are learning will be relevant to analyzing new problems.
• Building Your Medical Vocabulary, new to each endof-chapter Study Guide, focuses on familiarity with
the most common and useful biomedical word roots
and affixes. Like a mini-medical vocabulary course,
this will help students with retention, spelling, and
insight into medical terms, and ability to more comfortably approach even new terms beyond the scope
of this book.
• Muscle tables in chapter 10 are organized in a new,
more columnar format and enhanced with new color

shading for easier reading and learning.

• Male-female pelvic differences (fig. 8.37)
• Treatment of infant hip dislocation (fig. 9.27)
• External anatomy of the orbital region (fig. 16.22)
• Use of a spirometer (fig. 22.17)
vii

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STORYTELLING
Writing Style
Appropriate Level
• Plain language for A&P students early in their curricula
• Careful word selection and paragraph structure
• Appropriate for all audiences (international
readers, English as a second language, and
nontraditional students)
“The physiological mechanisms presented
throughout the text emphasize the basic
fundamental processes that occur in the
human body. I believe the information is
simplistic enough for students to comprehend yet detailed to provide important
information […] for students and for
instructors to present during lectures.”

• Avoidance of "dumbed down" content

Interactive Material
• Review activities integrated in the chapter
• Self-teaching prompts and simple experiments
liberally seeded through the narrative

—Scott Pallotta,
Baker College at Allen Park

• Learning aids such as pronunciation guides and
insights into the origins and root meanings of
medical terms

The Temporal Bones
If you palpate your skull just above and anterior
to the ear—that is, the temporal region—you
can feel the temporal bone, which forms the
lower wall and part of the floor of the cranial
cavity (fig. 8.10). The temporal bone derives its
name from the fact that people often develop
their first gray hairs on the temples with the
passage of time.9 The relatively complex shape
of the temporal bone is best understood by
dividing it into four parts:

Z

Homeostasis and Negative Feedback
e
●

Self-teaching prompts make
reading more active.
Word origins are footnoted.
Pro-NUN-see-AY-shun guides
help beginning students master
A&P.

Familiarity with word origins
helps students retain meaning
and spelling.

The human body has a remarkable capacity for selfrestoration. Hippocrates commented that it usually
returns to a state of equilibrium by itself, and people
recover from most illnesses even without the help of
a physician. This tendency results from homeostasis18
(HO-me-oh-STAY-sis), the body’s ability to detect
change, activate mechanisms that oppose it, and thereby
maintain relatively stable internal conditions.
French physiologist Claude Bernard (1813–78)
observed that the internal conditions of the body remain
quite constant even when external conditions vary greatly.
For example, whether it is freezing cold or swelteringly
hot outdoors, the internal temperature of the body stays
within a range of about 36° to 37°C (97°–99°F). American
physiologist Walter Cannon (1871–1945) coined the term
homeostasis for this tendency to maintain internal stability. Homeostasis has been one of the most enlightening
theories in physiology. We now see physiology as largely
a group of mechanisms for maintaining homeostasis, and
the loss of homeostatic control as the cause of illness

and death. Pathophysiology is essentially the study of
18

homeo = the same; stas = to place, stand, stay

viii

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STORYTELLING
Writing Style
Interesting Reading
• Students say the enlightening
analogies, clinical applications,
historical notes, biographical
vignettes, and evolutionary
insights make the book not
merely informative, but a
pleasure to read.

458

Axon

Signal
Action potential
in progress

++++–––+++++++++++
––––+++–––––––––––

Refractory
membrane
Excitable
membrane

––––+++–––––––––––
++++–––+++++++++++
+++++++++–––++++++
–––––––––+++––––––

–––––––––+++––––––
+++++++++–––++++++
+++++++++++++–––++
–––––––––––––+++––

–––––––––––––+++––
+++++++++++++–––++

FIGURE 12.16 Conduction of a Nerve Signal in an Unmyelinated
Fiber. Note that the membrane polarity is reversed in the region of the
action potential (red). A region of membrane in its refractory period
(yellow) trails the action potential and prevents the nerve signal from
going backward toward the soma. The other membrane areas (green)
are fully polarized and ready to respond.

voltage-gated channels immediately distal to the action

potential. Sodium and potassium channels open and close
just as they did at the trigger zone, and a new action potential is produced. By repetition, this excites the membrane
immediately distal to that. This chain reaction continues
until the traveling signal reaches the end of the axon.
Note that an action potential itself does not travel
along an axon; rather, it stimulates the production of a new
action potential in the membrane just ahead of it. Thus,
we can distinguish an action potential from a nerve signal.
The nerve signal is a traveling wave of excitation produced
by self-propagating action potentials. It is like a line of
falling dominoes. No one domino travels to the end of the
line, but each domino pushes over the next one and there
is a transmission of energy from the first domino to the last.
Similarly, no one action potential travels to the end of an
axon; a nerve signal is a chain reaction of action potentials.
production of a
If one action potential stimulates the produ
signal could
new one next to it, you might think that the si
also start traveling backward and return to the ssoma. This
membrane behind
does not occur,
however, because
the membra
CHAPTER
9 Joints
305
the nerve signal is still in its refractory period aand cannot
be restimulated. Only the membrane ahead is ssensitive to

Clinical applications
make the abstract science more
relevant.

Clinical Application

Knee Injuries and Arthroscopic Surgery
Although the knee can bear a lot of weight, it is highly vulnerable to
rotational and horizontal stress, especially when the knee is flexed
(as in skiing or running) and receives a blow from behind or from the
side. The most common injuries are to a meniscus or the anterior
cruciate ligament (ACL) (fig. 9.30). Knee injuries heal slowly because
ligaments and tendons have a scanty blood supply and cartilage
usually has no blood vessels at all.
The diagnosis and surgical treatment of knee injuries have been
greatly improved by arthroscopy, a procedure in which the interior
of a joint is viewed with a pencil-thin instrument, the arthroscope,
inserted through a small incision. The arthroscope has a light, a
lens, and fiber optics that allow a viewer to see into the cavity and
take photographs or video recordings. A surgeon can also withdraw samples of synovial fluid by arthroscopy or inject saline into
the joint cavity to expand it and provide a clearer view. If surgery
is required, additional small incisions can be made for the surgical instruments and the procedures can be observed through the
arthroscope or on a monitor. Arthroscopic surgery produces much
less tissue damage than conventional surgery and enables patients
to recover more quickly.
Orthopedic surgeons now often replace a damaged ACL with
a graft from the patellar ligament or a hamstring tendon. The surgeon “harvests” a strip from the middle of the patient’s ligament (or
tendon), drills a hole into the femur and tibia within the joint cavity,
threads the ligament through the holes, and fastens it with biodegradable screws. The grafted ligament is more taut and “competent” than
the damaged ACL. It becomes ingrown with blood vessels and serves

as a substrate for the deposition of more collagen, which further
strengthens it in time. Following arthroscopic ACL reconstruction,
a patient typically must use crutches for 7 to 10 days and undergo
supervised physical therapy for 6 to 10 weeks, followed by self-directed
exercise therapy. Healing is completed in about 9 months.

stimulation. The refractory period thus ensures that nerve
signals are conducted in the proper direction, from the
soma to the synaptic knobs.
A traveling nerve signal is an electrical current, but
it is not the same as a current traveling through a wire. A
current in a wire travels millions of meters per second and
is decremental—it gets weaker with distance. A nerve signal is much slower (not more than 2 m/s in unmyelinated
fibers), but it is nondecremental. Even in the longest axons,
the last action potential generated at a synaptic knob has
the same voltage
oltage as the first one generated at the trigger
la
a
zone. To clarify
this concept, we can compare the nerve
b
signal to a burning
fuse. When a fuse is lit, the heat ignites
m
powder immediately
in front of this point, and this repeats
itself in a self-propagating
fashion until the end of the fuse
e

is reached. A
At the end, the fuse burns just as hotly as it did
at the beginning.
In a fuse, the combustible powder is the
n
source of po
potential
energy that keeps the process going in
o
a nondecremental
fashion. In an axon, the potential energy
m
comes from
m the ion gradient across the plasma membrane.
Thus, the signal
does not grow weaker with distance; it is
sii
self-propagating,
like the burning of a fuse.
a

scientific content in a way
students can understand.

voltage-gated channels immediately distal to the action
potential. Sodium and potassium channels open and close
just as they did at the trigger zone, and a new action potential is produced. By repetition, this excites the membrane
immediately distal to that. This chain reaction continues
Myelinated
e Fibers until the traveling signal reaches the end of the axon.

Matters are
e somewhat different in myelinated fibers.
Voltage-gated
scarce inthat
the myelinte
e ion channels are
Note
an action potential itself does not travel
covered internodes—fewer
than 25/μm in these regions
te
compared with
2,000
to 12,000/μm
w
the nodesrather,
of
along
an ataxon;
it stimulates the production of a new
Ranvier. There
would be little point in having ion chanh
nels in thee internodes—myelin insulates the fiber from
action
potential
the ECF at these
points,
and Na from
t
the ECF could not in the membrane just ahead of it. Thus,

flow into the
th
h cell even if more channels were present.
can
an action potential from a nerve signal.
Therefore, n
no action we
potentials
can distinguish
occur in the internodes, and
d the nerve signal requires some other way of
traversing th
the
from one
node to thesignal
next.
h distance
The
nerve
is a traveling wave of excitation produced
When Na
N enters the axon at a node of Ranvier, it
diffuses forr a short distance
along the inner face of the
by self-propagating
action potentials. It is like a line of
axolemma (fig.
12.17a). Each sodium ion has an electri(
cal field around
it. When one Na moves toward another,

o
falling dominoes. No one domino travels to the end of the
its field repels
the second ion, which moves slightly and
p
repels another,
th
h and so forth—like two magnets that repel
but
domino pushes over the next one and there
each other if
to push their
northeach
poles together.
i you try line,
No one ion
on
n moves very far, but this energy transfer
ismuch
a transmission
travels down
faster and farther than any of energy from the first domino to the last.
w the axon
of the individual
ions. The signal grows weaker with
iv
distance, h
however,
partly
because the axoplasm

o
Similarly,
no resists
one action potential
ential travels to the end of an
the movement
me of the ions and partly because Na leaks
back out of
along thea
way.
Thereforesignal
with
o the axon
axon;
nerve
is a chain
n reaction of action potentials.
distance, there
is a lower and lower concentration of
th
h
Na to relay
ay
y the charge. Furthermore, with a surplus of
If one action potential stimulates
timulates the production of a
positive charges
on the inner face of the axolemma and
ha
a surplus of

o negative charges on the outer face, these
new
one
to it, you might
ht think that the signal could
cations and 
attracted
to eachnext
other through
d anions are
the membrane—like
the opposite poles of two magnets
ra
a
traveling
backward
d and return to the soma. This
attracting eeach
through astart
sheet of cardboard.
This
a other also
does not occur, however, because
ecause the membrane behind
the nerve signal is still in itss refractory period and cannot
be restimulated. Only the membrane
embrane ahead is sensitive to
2

2

+

+

+

+

+

Twisting motion

Foot fixed

Anterior cruciate
ligament (torn)
Tibial collateral
ligament (torn)
Medial
meniscus (torn)
Patellar ligament

FIGURE 9.30 Knee Injuries.

The ligaments of the ankle include (1) anterior and
posterior tibiofibular ligaments, which bind the tibia to the
fibula; (2) a multipart medial (deltoid30) ligament, which
binds the tibia to the foot on the medial side; and (3) a multipart lateral (collateral) ligament, which binds the fibula
to the foot on the lateral side. The calcaneal (Achilles)

tendon extends from the calf muscles to the calcaneus. It
plantarflexes the foot and limits dorsiflexion. Plantar flexion is limited by extensor tendons on the anterior side of
the ankle and by the anterior part of the joint capsule.
Sprains (torn ligaments and tendons) are common at
the ankle, especially when the foot is suddenly inverted or
everted to excess. They are painful and usually accompanied by immediate swelling. They are best treated by immo-

Analogies explain tough

Integration and Control

Cell body

• Even instructors say they often
learn something new and
interesting from Saladin’s
innovative perspectives.

DEEPER INSIGHT 9.4

PART THREE

Dendrites

bilizing the joint and reducing swelling with an ice pack,
but in extreme cases may require a cast or surgery. Sprains
and other joint disorders are briefly described in table 9.1.

“Saladin is a gifted author, and his conversational tone will be sure to keep students very
engaged.”

—Davonya Person,
Auburn University

Before You Go On
Answer the following questions to test your understanding of the
preceding section:
12. What keeps the mandibular condyle from slipping out of its
fossa in a posterior direction?
13. Explain how the biceps tendon braces the shoulder joint.
14. Identify the three joints found at the elbow and name the
movements in which each joint is involved.
15. What keeps the femur from slipping backward off the tibia?

delt = triangular, Greek letter delta (∆); oid = resembling

30

16. What keeps the tibia from slipping sideways off the talus?

ix

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CHAPTER 25

DEEPER INSIGHT 25.5

Medical History

The Man with a Hole in His Stomach
Perhaps the most famous episode in the history of digestive physiology began with a grave accident in 1822 on Mackinac Island in
northern Michigan. Alexis St. Martin, a 28-year-old Canadian voyageur
(fig. 25.33), was standing outside a trading post when he was accidentally hit by a shotgun blast from 3 feet away. A frontier Army doctor
stationed at Fort Mackinac, William Beaumont, was summoned to
examine St. Martin. As Beaumont later wrote, “a portion of the lung as
large as a turkey’s egg” protruded through St. Martin’s lacerated and
burnt flesh. Below that was a portion of the stomach with a puncture
in it “large enough to receive my forefinger.” Beaumont did his best
to pick out bone fragments and dress the wound, though he did not
expect St. Martin to survive.
Surprisingly, he lived. Over a period of months the wound
extruded pieces of bone, cartilage, gunshot, and gun wadding. As
the wound healed, a fistula (hole) remained in the stomach, so large
that Beaumont had to cover it with a compress to prevent food from
coming out. The opening remained, covered only by a loose flap of
skin, for the rest of St. Martin's life. A fold of tissue later grew over the
fistula, but it was easily opened. A year later, St. Martin was still feeble.
Town authorities decided they could no longer support him on public
funds and wanted to ship him 1,500 miles to his home. Beaumont,
however, was imbued with a passionate sense of destiny. Very little
was known about digestion, and he saw the accident as a unique
opportunity to learn. He took St. Martin in at his personal expense and
performed 238 experiments on him over several years. Beaumont had
never attended medical school and had little idea how scientists work,
yet he proved to be an astute experimenter. Under crude frontier
conditions and with almost no equipment, he discovered many of the
basic facts of gastric physiology discussed in this chapter.

“I can look directly into the cavity of the stomach, observe its
motion, and almost see the process of digestion,” Beaumont wrote.
“I can pour in water with a funnel and put in food with a spoon, and
draw them out again with a siphon.” He put pieces of meat on a string
into the stomach and removed them hourly for examination. He sent
vials of gastric juice to the leading chemists of America and Europe,
who could do little but report that it contained hydrochloric acid. He
proved that digestion required HCl and could even occur outside the
stomach, but he found that HCl alone did not digest meat; gastric
juice must contain some other digestive ingredient. Theodor Schwann,
one of the founders of the cell theory, identified that ingredient as
pepsin. Beaumont also demonstrated that gastric juice is secreted
only in response to food; it did not accumulate between meals as
previously thought. He disproved the idea that hunger is caused by
the walls of the empty stomach rubbing against each other.
Now disabled from wilderness travel, St. Martin agreed to
participate in Beaumont's experiments in exchange for room and
board—though he felt helpless and humiliated by it all. The fur trappers taunted him as “the man with a hole in his stomach,” and he

The Digestive System

995

longed to return to his work in the wilderness. He had a wife and
daughter in Canada whom he rarely got to see, and he ran away
repeatedly to join them. He was once gone for 4 years before poverty made him yield to Beaumont’s financial enticement to come
back. Beaumont despised St. Martin's drunkenness and profanity
and was quite insensitive to his embarrassment and discomfort over
the experiments. Yet St. Martin’s temper enabled Beaumont to make
the first direct observations of the relationship between emotion and

digestion. When St. Martin was particularly distressed, Beaumont
noted little digestion
occurring—as
we now know, the sympathetic
g
g
y p
nervous system
em inhibits digestive activity.
Beaumont
nt published a book in 1833 that laid the foundation
for modern gastric physiology and dietetics. It was enthusiastically
received by the medical community and had no equal until Russian
physiologist Ivan
van Pavlov (1849–1936) performed his celebrated experiments on digestion
gestion in animals. Building on the methods pioneered by
Beaumont, Pavlov received the 1904 Nobel Prize for Physiology or
Medicine.
In 1853, Beaumont slipped on some ice, suffered
suffer
f ed a blow to the
base of his skull, and died a few weeks later. St. Martin continued
to tour medical
ical schools and submit to experiments by other physiologists, whose
ose conclusions were often less correct than Beaumont’s.
Some, for example, attributed chemical digestion to lactic acid
instead of hydrochloric
drochloric acid. St. Martin lived in wretched poverty in a
tiny shack with
th his wife and several children, and died 28 years after

Beaumont. Byy then he was senile and believed he had been to Paris,
where Beaumont
mont had often promised to take him.

Medical History

Saladin “puts the human
in human A&P” with his occasional vignettes on the
people behind the science. Students say these stories
make learning A&P more fun and stimulating.

y
p
p
conditions and with almost no equipment, he discovered many of the
basic facts of gastric physiology discussed in this chapter.
“I can look directly into the cavity of the stomach, observe its
motion, and almost see the process of digestion,” Beaumont wrote.
“I can pour in water with a funnel and put in food with a spoon, and
draw them out again with a siphon.” He put pieces of meat on a string
into the stomach and removed them hourly for examination. He sent
vials of gastric juice to the leading chemists of America and Europe,
who could do little but report that it contained hydrochloric acid. He
proved that digestion required HCl and could even occur outside the
stomach, but he found that HCl alone did not digest meat; gastric
juice must contain some other digestive ingredient. Theodor Schwann,
one of the founders of the cell theory, identified that ingredient as
pepsin. Beaumont also demonstrated that gastric juice is secreted
only in response to food; it did not accumulate between meals as
previously thought. He disproved the idea that hunger is caused by

the walls of the empty stomach rubbing against each other.
Now disabled from wilderness travel, St. Martin agreed to
participate in Beaumont's experiments in exchange for room and
board—though he felt helpless and humiliated by it all. The fur trappers taunted him as “the man with a hole in his stomach,” and he

umont (1785–1853)
William Beaumont

William Beaumont (1785–1853)

Alexis St. Martin (1794–1880)

FIGURE 25.33 Doctor and Patient in a Pioneering Study of
Digestion.

Alexis St. Martin (1794–1880)

FIGURE 25.33
5.33
33
3 Doctor and Patient in a Pioneering Study of
Digestion.

More than a few distinguished scientists and clinicians say they found their inspiration in reading of
the lives of their predecessors. Maybe these stories
will inspire some of our own students to go on to
do great things.
–Ken Saladin

1076

Evolutionary Medicine

Rapidly growing,

increasingly fascinating

Evolutionary medicine provides
novel and intriguing ways of
looking at:
• menopause
• the sweet tooth
• bipedalism
• the origin of mitochondria
• skin color
• body hair
• lactose intolerance
• the kidney and life on dry land
• the palate
• theories of aging and death

DEEPER INSIGHT 28.2

PART FIVE

Reproduction and Development

stimulates gonadotropin secretion. Therefore, if body fat
and leptin levels drop too low, gonadotropin secretion
declines and a girl’s or woman’s menstrual cycle may

cease. Adolescent girls with very low body fat, such as
avid dancers and gymnasts, tend to begin menstruating at
a later age than average.
Menarche does not necessarily signify fertility. A
girl’s first few menstrual cycles are typically anovulatory
(no egg is ovulated). Most girls begin ovulating regularly
about a year after they begin menstruating.
Estradiol stimulates many other changes of puberty. It
causes the vaginal metaplasia described earlier. It stimulates growth of the ovaries and secondary sex organs. It
stimulates growth hormone secretion and causes a rapid
increase in height and widening of the pelvis. Estradiol is
largely responsible for the feminine physique because it
stimulates fat deposition in the mons pubis, labia majora,
hips, thighs, buttocks, and breasts. It makes a girl’s skin
thicken, but the skin remains thinner, softer, and warmer
than in males of corresponding age.
Progesterone27 acts primarily on the uterus, preparing it for possible pregnancy in the second half of each
menstrual cycle and playing roles in pregnancy discussed
later. Estrogens and progesterone also suppress FSH and
LH secretion through negative feedback inhibition of the
anterior pituitary. Inhibin selectively suppresses FSH
secretion.
Thus, we see many hormonal similarities in males
and females from puberty onward. The sexes differ less
in the identity of the hormones that are present than in
their relative amounts—high levels of androgens and
low levels of estrogens in males and the opposite in
females. Another difference is that these hormones are
secreted more or less continually and simultaneously
in males, whereas in females, secretion is distinctly

cyclic and the hormones are secreted in sequence. This
will be very apparent as you read about the ovarian and
menstrual cycles.

Evolutionary Medicine

The Evolution of Menopause

There has been considerable speculation about why women do not
remain fertile to the end of their lives, as men do. Some theorists
argue that menopause served a biological purpose for our prehistoric
foremothers. Human offspring take a long time to rear. Beyond a certain point, the frailties of age make it unlikely that a woman could rear
another infant to maturity or even survive the stress of pregnancy. She
might do better in the long run to become infertile and finish rearing
her last child, or help to rear her grandchildren, instead of having
more. In this view, menopause was biologically
advantageous for our
Climacteric and Menopause
ancestors—in other words, an evolutionary
adaptation.
Women,
like men, go through a midlife change in hormone secretion called the climacteric. In women, it is
Others argue against this hypothesis onaccompanied
the grounds
thatthe
Pleistocene
by menopause,
cessation of menstruation (see Deeper Insight 28.2).
A female israrely
born with lived

about 2 million
in her ova(Ice Age) skeletons indicate that early hominids
pasteggsage
ries, each in its own follicle. The older she gets, the fewer
specific age,
Climacteric
begins not at
40. If this is true, menopause setting infollicles
at remain.
45 to
55  years
ofanyage
but when she has only about 1,000 follicles left. Even the
remaining
follicles
are less responsive to
gonadotropins, so
could have served little purpose. In this
view,
Pleistocene
women
they secrete less estrogen and progesterone. Without these
steroids,
uterus, vagina,
breasts atrophy. Intercourse
may indeed have been fertile to the end
of thetheir
lives;andmenopause
may become uncomfortable, and vaginal infections more
now may be just an artifact of modern nutrition and medicine, which

have made it possible for us to live much longer than our ancestors did.
27

common, as the vagina becomes thinner, less distensible,
and drier. The skin becomes thinner, cholesterol levels
rise (increasing the risk of cardiovascular disease), and
bone mass declines (increasing the risk of osteoporosis).
Blood vessels constrict and dilate in response to shifting
hormone balances, and the sudden dilation of cutaneous arteries may cause hot flashes—a spreading sense of
heat from the abdomen to the thorax, neck, and face. Hot
flashes may occur several times a day, sometimes accompanied by headaches resulting from the sudden vasodilation
of arteries in the head. In some people, the changing hormonal profile also causes mood changes. Many physicians
prescribe hormone replacement therapy (HRT)—low doses
of estrogen and progesterone usually taken orally or by a
skin patch—to relieve some of these symptoms. The risks
and benefits of HRT are still being debated.

Apply What You Know
FSH and LH secretion rise at climacteric and these hormones attain high concentrations in the blood. Explain this
using the preceding information and what you know about
the pituitary–gonadal relationship.

Menopause is the cessation of menstrual cycles, usually occurring between the ages of 45 and 55. The average
age has increased steadily in the last century and is now
about 52. It is difficult to precisely establish the time of
menopause because the menstrual periods can stop for
several months and then begin again. Menopause is generally considered to have occurred when there has been
no menstruation for a year or more.

DEEPER INSIGHT 28.2

Evolutionary Medicine

The Evolution of Menopause
There has been considerable speculation about why women do not
remain fertile to the end of their lives, as men do. Some theorists
argue that menopause served a biological purpose for our prehistoric
foremothers. Human offspring take a long time to rear. Beyond a certain point, the frailties of age make it unlikely that a woman could rear
another infant to maturity or even survive the stress of pregnancy. She
might do better in the long run to become infertile and finish rearing
her last child, or help to rear her grandchildren, instead of having
more. In this view, menopause was biologically advantageous for our
ancestors—in other words, an evolutionary adaptation.
Others argue against this hypothesis on the grounds that Pleistocene
(Ice Age) skeletons indicate that early hominids rarely lived past age
40. If this is true, menopause setting in at 45 to 55  years of age
could have served little purpose. In this view, Pleistocene women
may indeed have been fertile to the end of their lives; menopause
now may be just an artifact of modern nutrition and medicine, which
have made it possible for us to live much longer than our ancestors did.

pro = favoring; gest = pregnancy; sterone = steroid hormone

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ARTWORK THAT
Inspires Learning
CHAPTER 3

Cellular Form and Function

103

Microvilli

Sets the Standard
• Stunning portfolio of art and photos

Microfilaments

• Hundreds of accuracy reviews
Terminal web

Secretory
vesicle in
transport

• Art focus groups

Lysosome

Desmosome

Kinesin
Microtubule

Vivid Illustrations

Rich textures
and shading, and bold, bright colors
bring structures to life.

Intermediate
filaments

Intermediate
filaments

Microtubule
in the process
of assembly

Centrosome
Microtubule
undergoing
disassembly

Nucleus
Mitochondrion

Axoneme:
Peripheral microtubules
Central microtubules
Dynein arms

(a)

Basement
membrane

Cilia

Hemidesmosome

Shaft of cilium

(b)

FIGURE 3.25 The C
Cytoskeleton. (a) Components of the cytoskeleton.
Basal body
(a)

Plasma membrane

10 μm

rre show
Few organelles are
shown in order to emphasize the cytoskeleton. Note that all
microtubules radiate
from the centrosome; they often serve as trackways for motor
iate fro
proteins (kinesin) transp
transporting organelles. (b) Cells with their cytoskeletons labeled

with fluorescent aantibod
antibodies, photographed through a fluorescence microscope.
The density of a ttypical
cytoskeleton far exceeds even that shown in part (a).
yp

(b)

15 μm

Page 103

Cilia

Page 721
Axoneme

Microvilli

Dynein
arm
Central
microtubule
Peripheral
microtubules

(c)

0.15 μm

(d)

Page 89

The visual appeal of nature is immensely important
in motivating one to study it. We certainly see this
at work in human anatomy—in the countless students who describe themselves as ‘visual learners’;
in the many laypeople who find anatomy atlases so
intriguing; and in the enormous popularity of Body
Worlds and similar exhibitions of human anatomy.

(a)

ocardiall V
o
Vortex. (a) Anterior view of

–Ken Saladin uum rendered
rend
de
transparent to expose

m
muscle.
(b)
( View from the apex to
oils around
o
arou
u the heart. This results in a

ventricles
vventricle
es contract.

(b)

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Page 193

“The diagrams and the photographs of the human body
structure bring real elements into the text and
excitement for students who
are being introduced to A&P
for the first time.”

Old club hair
Epidermis
Sebaceous
gland

Club hair
(detached
from matrix)

Piloerector
New hair

Club

Bulge
Hair matrix
Hair bulb

Degeneration
of lower follicle

Dermis

1 Anagen (early)
Anagen (mature)
(Growing phase, 6–8 years)
Stem cells multiply and follicle grows deeper into dermis; hair matrix
cells multiply and keratinize, causing hair to grow upward; old club hair
may persist temporarily alongside newly growing hair.

Dermal papilla

—Charmaine Irvin, Baker
College of Allen Park

3 Telogen
(Resting phase, 1–3 months)
Dermal papilla has ascended
to level of bulge; club hair falls

out, usually in telogen or
next anagen.

2 Catagen
(Degenerative phase, 2–3 weeks)
Hair growth ceases; hair bulb
keratinizes and forms club hair;
lower follicle degenerates.

FIGURE 6 9

Page 764
TABLE 10.1

Conducive to Learning

Muscles of Facial Expression (continued)

Risorius24
(rih-SOR-ee-us)
Depressor Anguli Oris

25

Depressor Labii Inferioris26

• Easy-to-understand
process figures

Draws angle of mouth laterally in expressions of laughing,

horror, or disdain

O: Zygomatic arch; fascia near ear
I: Modiolus

Facial nerve

Draws angle of mouth laterally and downward in opening
mouth or sad expressions

O: Inferior margin of mandibular body
I: Modiolus

Facial nerve

Draws lower lip downward and laterally in chewing and
expressions of melancholy or doubt

O: Mandible near mental protuberance
I: Skin and mucosa of lower lip

Facial nerve

The Mental and Buccal Regions. Adjacent to the oral orifice are the mental region (chin) and buccal region (cheek). In addition to muscles already discussed that
act directly on the lower lip, the mental region has a pair of small mentalis muscles extending from the upper margin of the mandible to the skin of the chin. In some
people, these muscles are especially thick and have a visible dimple between them called the mental cleft (see fig. 4.18, p. 135). The buccinator is the muscle in the
cheek. It has multiple functions in chewing, sucking, and blowing. If the cheek is inflated with air, compression of the buccinator blows it out. Sucking is achieved
by contracting the buccinators to draw the cheeks inward, and then relaxing them. This action is especially important to nursing infants. To feel this action, hold
your fingertips lightly on your cheeks as you make a kissing noise. You will notice the relaxation of the buccinators at the moment air is sharply drawn in through
the pursed lips. The platysma is a thin superficial muscle of the upper chest and lower face. It is relatively unimportant, but when men shave they tend to tense the

platysma to make the concavity between the jaw and neck shallower and the skin tauter.

• Tools for students to easily
orient themselves
Page 390
Anterior
Pectoralis
major m.

Mentalis
(men-TAY-lis)

Elevates and protrudes lower lip in drinking, pouting, and
expressions of doubt or disdain; elevates and wrinkles skin
of chin

O: Mandible near inferior incisors
I: Skin of chin at mental protuberance

Facial nerve

Buccinator27
(BUC-sin-AY-tur)

Compresses cheek against teeth and gums; directs food
between molars; retracts cheek from teeth when mouth is
closing to prevent biting cheek; expels air and liquid

O: Alveolar processes on lateral surfaces of
mandible and maxilla

I: Orbicularis oris; submucosa of cheek
and lips

Facial nerve

Platysma28
(plah-TIZ-muh)

Draws lower lip and angle of mouth downward in
expressions
of horror or surprise; may aid in opening
Sternum
mouth widely

O: Fascia of deltoid and pectoralis major
I: Mandible; skin and subcutaneous
tissue of lower face

Facial nerve

Fat of breast

Ventricles
of heart

Ribs

Pericardial
cavity

Muscles tables are organized in new
columnar format and enhanced with
new shading for easier reading and
learning.

Right lung
Esophagus

Atria of heart

Aorta
Vertebra

Left lung

Spinal cord
Pleural cavity
10

1 Blood enters right atrium from superior
and inferior venae cavae.

Posterior
Aorta
Left pulmonary
artery

11
5

Orientation Tools Saladin art integrates tools
to help students quickly orient themselves within a
figure and make connections between ideas.

5
9

Pulmonary trunk

Superior
vena cava
Right
pulmonary
veins

4

6

6

Left pulmonary
veins
Left atrium

1

Aortic valve
7

3
Right
atrium

Left AV
(bicuspid) valve
8

2

Right AV
(tricuspid) valve

Left ventricle

3 Contraction of right ventricle forces
pulmonary valve open.
4 Blood flows through pulmonary valve
into pulmonary trunk.
5 Blood is distributed by right and left
pulmonary arteries to the lungs, where it
unloads CO2 and loads O2.
6 Blood returns from lungs via pulmonary
veins to left atrium.
7 Blood in left atrium flows through left AV
valve into left ventricle.
8 Contraction of left ventricle (simultaneous with
3 ) forces aortic valve open.
step 3)
9 Blood flows through aortic valve into

ascending aorta.

Right
ventricle
Inferior
vena cava

2 Blood in right atrium flows through right
AV valve into right ventricle.

11

10 Blood in aorta is distributed to every organ in
the body, where it unloads O2 and loads CO2.
11 Blood returns to heart via venae cavae.

Process Figures Saladin breaks complicated

Page 721

physiological processes into numbered steps for a manageable introduction to difficult concepts.
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PEDAGOGICAL
Learning Tools

Engaging Chapter Layouts
• Chapters are structured around the
way students learn.
• Frequent subheadings and
expected learning outcomes
help students plan their study
time and review strategies.

CHAPTER

12
NERVOUS TISSUE

A Purkinje cell, a neuron from the cerebellum of the brain

Chapter Outline provides

CHAPTER OUTLINE

a quick overview of the content.

12.1 Overview of the Nervous System 440

Deeper Insights highlight areas

12.2 Properties of Neurons 441
• Universal Properties 441
• Functional Classes 442
• Structure of a Neuron 442
• Axonal Transport 445

of interest for students.

12.3 Supportive Cells (Neuroglia) 446
• Types of Neuroglia 446
• Myelin 448
• Unmyelinated Nerve Fibers 450
• Conduction Speed of Nerve Fibers 450
• Regeneration of Nerve Fibers 450

• Neurotransmitters and Related
Messengers 461
• Synaptic Transmission 463
• Cessation of the Signal 465
• Neuromodulators 465
12.6 Neural Integration 466
• Postsynaptic Potentials 466
• Summation, Facilitation, and
Inhibition 467
• Neural Coding 468
• Neural Pools and Circuits 469
• Memory and Synaptic Plasticity 471

DEEPER INSIGHTS
12.1 Clinical Application: Glial Cells and Brain
Tumors 447
12.2 Clinical Application: Diseases of the Myelin
Sheath 448
12.3 Medical History: Nerve Growth Factor—
From Home Laboratory to Nobel Prize 452

12.4 Clinical Application: Alzheimer and
Parkinson Diseases 472

Connective Issues 474
Study Guide 475

12.4 Electrophysiology of Neurons 451
• Electrical Potentials and Currents 452
• The Resting Membrane Potential 453
• Local Potentials 454
• Action Potentials 455
• The Refractory Period 457
• Signal Conduction in Nerve Fibers 457
12.5 Synapses 460
• The Discovery of Neurotransmitters 460
• Structure of a Chemical Synapse 461

Module 7: Nervous System

439

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750

Tiered Assessments Based on Key
Learning Outcomes
• Chapters are divided into easily manageable
chunks, which help students
budget study time effectively.
• Section-ending questions allow students to
check their understanding before moving on.

New! Each chapter begins with Brushing
Up to emphasize the interrelatedness of concepts and also provides an aid to returning,
nontraditional students.
Each numbered section begins with Expected
Learning Outcomes to help focus the reader’s
attention on the larger concepts and make the
course outcome-driven.

282

PART TWO

Support and Movement

fibers that extend from the bone matrix of the jaw into
the dental tissue (see fig. 9.2b). The periodontal ligament
allows the tooth to move or give a little under the stress of
chewing. This allows us to sense how hard we are biting or
to sense a particle of food stuck between the teeth.

the forearm. A less movable syndesmosis is the one that
binds the distal ends of the tibia and fibula together, side

by side (see fig. 9.2c).

Syndesmoses

A cartilaginous joint is also called an amphiarthrosis7
(AM-fee-ar-THRO-sis) or amphiarthrodial joint. In these
joints, two bones are linked by cartilage (fig. 9.4). The
two types of cartilaginous joints are synchondroses and
symphyses.

6

A syndesmosis (SIN-dez-MO-sis) is a fibrous joint at
which two bones are bound by relatively long collagenous
fibers. The separation between the bones and length of
the fibers give these joints more mobility than a suture
or gomphosis has. An especially movable syndesmosis
exists between the shafts of the radius and ulna, which
are joined by a broad fibrous interosseous membrane. This
permits such movements as pronation and supination of

Cartilaginous Joints

Synchondroses
A synchondrosis8 (SIN-con-DRO-sis) is a joint in which
the bones are bound by hyaline cartilage. An example is
7

syn = together; desm = band; osis = condition

6

Clavicle

8

PART FOUR

Regulation and Maintenance

Brushing Up...
• The concepts of homeostatic set point and dynamic equilibrium
should be reviewed (p. 17) as background for understanding the
control of blood pressure.
• The principles of blood volume, pressure, and flow discussed in this
chapter hinge on the reasons behind the osmolarity and viscosity of
blood introduced on page 682.
• Familiarity with cardiac systole and diastole (p. 728) is necessary for
understanding blood pressure in this chapter.
• Blood flow is regulated by variations in cardiac output and blood
vessel diameter, which are governed in part by the autonomic
nervous system as discussed on page 576.
• The exchange of materials between the blood capillaries and
surrounding tissues is based on the principles of filtration, osmosis
and osmotic pressure, diffusion, and transcytosis introduced earlier
(pp. 91–100).

crackpot because his conclusion flew in the face of common sense—
if the blood was continually recirculated and not consumed by the
tissues, they reasoned, then what purpose could it serve? We now

know, of course, that he was right. Harvey’s case is one of the most
interesting in biomedical history, for it shows how empirical science
overthrows old theories and spawns better ones, and how common
sense and blind allegiance to authority can interfere with the
acceptance of truth. But most importantly, Harvey’s contributions
represent the birth of experimental physiology.

20.1 General Anatomy of
the Blood Vessels
Expected Learning Outcomes
When you have completed this section, you should be able to
a. describe the structure of a blood vessel;

T

he route taken by the blood after it leaves the heart was a
point of much confusion for many centuries. In traditional
Chinese medicine as early as 2650 BCE, blood was believed
to flow in a complete circuit around the body and back to the
heart, just as we know today. But in the second century CE, Roman
physician Claudius Galen (129–c. 199) argued that it flowed
back and forth in the veins, like air in the bronchial tubes. He
believed that the liver received food directly from the esophagus
and converted it to blood, the heart pumped the blood through
the veins to all other organs, and those organs consumed it.
The arteries were thought to contain only a mysterious vapor or
“vital spirit.”
The Chinese view was right, but the first experimental
demonstration of this did not come for another 4,000 years. English
physician William Harvey (1578–1657) (see p. 5) studied the filling

and emptying of the heart in snakes, tied off the vessels above
aand be
below the heart to observe the effects on cardiac filling and
output
o
output, measured cardiac output in a variety of living animals,
aand est
estimated cardiac output in humans. He concluded that (1)
th
he hea
the
heart pumps more blood in half an hour than there is in the
entire b
e
body, (2) not enough food is consumed to account for the
ccontinu
continual production of so much blood, and therefore (3) the blood
re
eturns to the heart rather than being consumed by the peripheral
returns
organs
o
organs. He could not explain how, since the microscope had yet to
be dev
b
developed to the point that enabled Antony van Leeuwenhoek
1632–
1
(1632–1723)
and Marcello Malpighi (1628–94) to discover the blood

ccapillar
capillaries.
Ha
Harvey’s
work was the first experimental study of animal
physiol
p
physiology and a landmark in the history of biology and medicine.
But so entrenched were the ideas of Aristotle and Galen in
B
tthe
he me
medical community, and so strange was the idea of doing
experim
e
experiments on living animals, that Harvey’s contemporaries
re
ejecte his ideas. Indeed, some of them regarded him as a
rejected

b. describe the different types of arteries, capillaries,
and veins;
c. trace the general route usually taken by the blood from
the heart and back again; and
d. describe some variations on this route.
There are three principal categories of blood vessels:
arteries, veins, and capillaries (fig. 20.1). Arteries are the
efferent vessels of the cardiovascular system—that is,
vessels that carry blood away from the heart. Veins are
the afferent vessels that carry blood back to the heart.

Capillaries are microscopic, thin-walled vessels that connect the smallest arteries to the smallest veins. Aside from
their general location and direction of blood flow, these
three categories of vessels also differ in the histological
structure of their walls.

The Vessel Wall
The walls of arteries and veins are composed of three
layers called tunics (fig. 20.2):
1. The tunica interna (tunica intima) lines the inside of
the vessel and is exposed to the blood. It consists of a
simple squamous epithelium called the endothelium
overlying a basement membrane and a sparse layer
of loose connective tissue; it is continuous with the
endocardium of the heart. The endothelium acts as
a selectively permeable barrier to materials entering
or leaving the bloodstream; it secretes chemicals that
stimulate dilation or constriction of the vessel; and it
normally repels blood cells and platelets so that they
flow freely without sticking to the vessel wall. When
the endothelium is damaged, however, platelets may
adhere to it and form a blood clot; and when the
tissue around a vessel is inflamed, the endothelial

amphi = on all sides; arthr = joined; osis = condition
syn = together; chondr = cartilage; osis = condition

Sternum

Rib 1
Costal

cartilage

Intervertebral
disc (fibrocartilage)
(a)

rim. Dislocations of the hip are rare, but some infants suffer
congenital dislocations because the acetabulum is not deep
enough
to hold
Body
vertebrahead of the femur in place. If detected
B d of the
(c)
early, this condition can be treated with a harness, worn
for 2 to 4 months, that holds the head of the femur in the
proper position until the joint is stronger (fig. 9.27).

FIGURE 9.4 Cartilaginous Joints.
(a) A synchondrosis, represented by the
costal cartilage joining rib 1 to the sternum.
(b) The pubic symphysis. (c) Intervertebral
discs, which join adjacent vertebrae to each
other by symphyses.
● What is the difference between the pubic
symphysis and the interpubic disc?

Interpubic disc
(fibrocartilage)

(b)

Pubic symphysis

Questions in figure legends and
Apply What You Know items
prompt students to think more
deeply about the implications and
applications of what they have
learned.

Apply What You Know
Where else in the body is there a structure similar to the
acetabular labrum? What do those two locations have in
common?

Ligaments that support the coxal joint include the
iliofemoral (ILL-ee-oh-FEM-oh-rul) and pubofemoral
(PYU-bo-FEM-or-ul) ligaments on the anterior side and
the ischiofemoral (ISS-kee-oh-FEM-or-ul) ligament on the
posterior side. The name of each ligament refers to  the
bones to which it attaches—the femur and the ilium,
pubis, or ischium. When you stand up, these ligaments
become twisted and pull the head of the femur tightly
into the acetabulum. The head of the femur has a conspicuous pit called the fovea capitis. The round ligament, or
ligamentum teres27 (TERR-eez), arises here and attaches

End-of-chapter questions
build on all levels of Bloom's
taxonomy in sections that:

1. assess learning outcomes
2. test simple recall and
analytical thought
3. build medical vocabulary
4. apply the basic knowledge to
new clinical problems and
other situations

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INNOVATIVE
Chapter Sequencing
Innovative Chapter
Order
Some chapters and topics are presented
in a sequence that is more instructive
than the conventional order.

Early Presentation of Heredity
Fundamental principles of heredity are
presented in the last few pages of
chapter 4 rather than at the back of the
book to better integrate molecular and
mendelian genetics. This organization
also prepares students to learn about

such genetic traits and conditions as
cystic fibrosis, color blindness, blood
types, hemophilia, cancer genes, or
sickle-cell disease by first teaching
them about dominant and recessive
alleles, genotype and phenotype, and
sex linkage.

BRIEF

About the Author iv
Preface v
Reviewers xxi
Contents xxii
Letter to the Students xxvi

The functional morphology of the
skeleton, joints, and muscles is treated
in three consecutive chapters, 8 through
10, so when students learn muscle
origins and insertions, these come only
two chapters after the names of the
relevant bone features. When they learn
muscle actions, it is in the first chapter
after learning the terms for the joint
movements. This order brings another
advantage: the physiology of muscle and
nerve cells is treated in two consecutive
chapters (11 and 12), which are thus
closely integrated in their treatment of

synapses, neurotransmitters, and
membrane electrophysiology.

Sense Organs 582
The Endocrine System 633

PART FOUR
Regulation and Maintenance

PART ONE
Organization of the Body
1

2
3
4
5

Major Themes of Anatomy and Physiology 1
Atlas A General Orientation to Human
Anatomy 28
The Chemistry of Life 42
Cellular Form and Function 78
Genetics and Cellular Function 114
Histology 143

18
19

The Circulatory System: Blood 678

The Circulatory System: The Heart 714

20

The Circulatory System: Blood Vessels
and Circulation 749
The Lymphatic and Immune Systems 808
The Respiratory System 854
The Urinary System 895
Water, Electrolyte, and Acid–Base Balance 930
The Digestive System 953
Nutrition and Metabolism 1000

21
22
23
24
25
26

PART TWO
Support and Movement
6
7
8
9
10
11

The Integumentary System 180

Bone Tissue 206
The Skeletal System 233
Joints 278
The Muscular System 312
Atlas B Regional and Surface Anatomy 379
Muscular Tissue 401

PART THREE

Muscle Anatomy and
Physiology Follow Skeleton
and Joints

16
17

Contents

Integration and Control
12
13
14
15

Nervous Tissue 439
The Spinal Cord, Spinal Nerves, and
Somatic Reflexes 478
The Brain and Cranial Nerves 511
The Autonomic Nervous System and
Visceral Reflexes 561

PART FIVE
Reproduction and Development
27
28
29

The Male Reproductive System 1034
The Female Reproductive System 1064
Human Development and Aging 1102

Appendix A. Periodic Table A-1
Appendix B. Answer Keys A-3
Appendix C. Symbols of Weight and Measures A-15
Appendix D. Biomedical Abbreviations A-17
Glossary G-1
Credits C-1
Index I-1

Urinary System Presented Close to Circulatory and
Respiratory Systems
Most textbooks place this system near the end of the book because
of its anatomical and developmental relationships with the
reproductive system. However, its physiological ties to the
circulatory and respiratory systems are much more important.
Except for a necessary digression on lymphatics and immunity, the
circulatory system is followed almost immediately with the
respiratory and urinary systems.

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ANOTHER LAYER TO ENHANCE THE CONNECTION

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Ph.I.L.S. (physiology
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Student study guide

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Engaging
Presentation Materials
for Lecture and Lab

New! ConnectPlus with LearnSmart makes
teaching easier and learning smarter

McGraw-Hill ConnectPlus Anatomy & Physiology is a
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Anatomy & Physiology Revealed 3.0

Anatomy & Physiology Revealed is the ultimate online
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Anatomy & Physiology Revealed also offers animations,
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New to this edition, the text, images, and artwork in the Saladin’s Anatomy & Physiology 6th edition are
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they can simply click the logo and they will be taken specifically to the dissection photos, animations,
histology slides, and radiological images in APR that support and enrich their understanding of the text.
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This comprehensive study guide written by experienced
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Saladin contains vocabulary-building and contenttesting exercises, labeling exercises, and practice exams.

The Anatomy & Physiology
Laboratory Manual by Eric Wise of
Santa Barbara City College is
expressly written to coincide with
chapters of Saladin's Anatomy &
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Reviewers
Tami Asplin, North Dakota State
University

Jody Johnson, Arapahoe Community
College

Charles J. Venglarik, Jefferson State
Community College

Seher Atamturktur, Bronx Community
College of CUNY

Jamie Joseph, University of Waterloo,
School of Pharmacy

Janice Webster, Ivy Tech Community
College

Vincent Austin, Bluegrass Community
and Technical College

Roman Kondratov, Cleveland State
University

Van Wheat, South Texas College

Melissa M. Bailey, Emporia State
University

Raymond Larsen, Bowling Green State
University

Jeanne K. Barnett, University of Southern
Indiana

Sarah Liechty, Ivy Tech Community
College

Board of Advisors

Jerry D. Barton II, Tarrant County
College-South

Jo Anne Lucas, Wayne County
Community College District

Dr. Peter G. Bushnell, Indiana University
South Bend

Moges Bizuneh, Ivy Tech Community
College

Paul Luyster, Tarrant County College
District

Cindy Prentice-Craver, Chemeketa
Community College

Barbara A. Coles, Wake Technical
Community College

David E. Manry, Hillsborough
Community College

Dr. Timothy A Ballard, University of
North Carolina Wilmington

Teresa Cowan, Baker College of Clinton
Township

Margaret McMichael, Baton Rouge
Community College

Dr. Jane L. Johnson-Murray, Houston
Community College

Melissa A. Deadmond, Truckee Meadows
Community College

Kristina Miranda, Tarrant County College

Vladimir Jurukovski, PhD, Suffolk
County Community College

Heather J. Evans Anderson, Winthrop
University
Greg Feitelson, Ivy Tech Community
College
Dean Furbish, Wake Technical
Community College
Deborah Furbish, Wake Technical
Community College
Michael Gaetz, University of Fraser
Valley
Anthony Gaudin, Ivy Tech Community
College
Matthew Geddis, Borough of Manhattan
Community College-City Univof NY
Elmer Godeny, Baton Rouge Community
College
Sylvester Hackworth, Bishop State
Community College
Elizabeth Hoffman, Baker College of
Clinton Township
Charmaine Irvin, Baker College of Allen

Park
Jean Jackson, Bluegrass Community and
Technical College

Lucia New, Saskatchewan Institute of
Applied Arts & Sciences, Kelsey Campus

Shirley Whitescarver, Bluegrass
Community and Technical College

Dale Smoak, Piedmont Technical College

Scott Pallotta, Baker College at Allen
Park

Dr. Wanda Hargroder, Louisiana State
University

Glenn H. Parker, Laurentian University

Martha J. Ross, Jefferson State
Community College

Ajay Patel, Langara College
Davonya Person, Auburn University
Gilbert Pitts, Austin Peay State
University
Dr. William A. Said, University of
Georgia
Ronald Slavin, Borough of Manhattan

Community College-City Univ of NY
Ken Smith, Arapahoe Community
College
Kerry L. Smith, Oakland Community
College

Marcia Bradley, Ocean County College
Cliff Fontenot, Southeastern Louisiana
University
Dr. James Junker, University of Maryland
Eastern Shore
James Horwitz, Palm Beach Community
College – Lake Worth
Sonya Williams, Oklahoma City
Community College
Teresa Gillian, Virginia Tech

Thomas Snyder, Augusta State
University
Bonnie J. Tarricone, Ivy Tech Community
College
Christine Terry, Augusta State University
James F. Thompson, Austin Peay State
University

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Contents
PART ONE

Chapter 2

PART TWO

Organization of the Body

The Chemistry of Life 42

Support and Movement

2.1 Atoms, Ions, and Molecules 43
2.2 Water and Mixtures 50
2.3 Energy and Chemical Reactions 56
2.4 Organic Compounds 59
Study Guide 75

Chapter 3
Cellular Form and Function 78

Chapter 1

3.1 Concepts of Cellular Structure 79
3.2 The Cell Surface 82
3.3 Membrane Transport 91
3.4 The Cell Interior 101

Study Guide 111

Major Themes of Anatomy and
Physiology 1

Chapter 4

1.1

The Scope of Anatomy and
Physiology 2
1.2 The Origins of Biomedical
Science 3
1.3 Scientific Method 7
1.4 Human Origins and Adaptations 9
1.5 Human Structure 12
1.6 Human Function 14
1.7 The Language of Medicine 20
1.8 Review of Major Themes 22
Study Guide 25

Genetics and Cellular Function 114

Atlas A

5.1
5.2
5.3
5.4

General Orientation to Human
Anatomy 28
A.1 General Anatomical Terminology 29
A.2 Major Body Regions 31
A.3 Body Cavities and Membranes 34
A.4 Organ Systems 37
Study Guide 40

Chapter 6
The Integumentary System 180

4.1

DNA and RNA—The Nucleic
Acids 115
4.2 Genes and Their Action 120
4.3 DNA Replication and the Cell
Cycle 129
4.4 Chromosomes and Heredity 134
Study Guide 140

6.1

The Skin and Subcutaneous
Tissue 181
6.2 Hair and Nails 190
6.3 Cutaneous Glands 195
6.4 Skin Disorders 197
Connective Issues 202
Study Guide 203

Chapter 7
Bone Tissue 206

Chapter 5
Histology 143
The Study of Tissues 144
Epithelial Tissue 146
Connective Tissue 153
Nervous and Muscular Tissues—
Excitable Tissues 162
5.5 Cell Junctions, Glands, and
Membranes 164
5.6 Tissue Growth, Development,
Repair, and Degeneration 171
Study Guide 177

7.1

Tissues and Organs of the Skeletal
System 207
7.2 Histology of Osseous Tissue 209
7.3 Bone Development 214
7.4 Physiology of Osseous Tissue 220
7.5 Bone Disorders 225
Connective Issues 229
Study Guide 230

Chapter 8
The Skeletal System 233

8.1
8.2

Overview of the Skeleton 234
The Skull 236

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8.3

The Vertebral Column and Thoracic
Cage 250
8.4 The Pectoral Girdle and Upper
Limb 259
8.5 The Pelvic Girdle and Lower
Limb 265
Study Guide 275

Chapter 9
Joints 278
9.1
9.2
9.3

Joints and Their Classification 279

Synovial Joints 283
Anatomy of Selected
Diarthroses 298
Study Guide 309

11.2 Microscopic Anatomy of Skeletal
Muscle 403
11.3 The Nerve–Muscle Relationship 408
11.4 Behavior of Skeletal Muscle
Fibers 411
11.5 Behavior of Whole Muscles 418
11.6 Muscle Metabolism 423
11.7 Cardiac and Smooth Muscle 428
Connective Issues 435
Study Guide 436

PART THREE
Integration and Control

Chapter 10
The Muscular System 312
10.1 The Structural and Functional
Organization of Muscles 313
10.2 Muscles of the Head and Neck
322
10.3 Muscles of the Trunk 333
10.4 Muscles Acting on the Shoulder and
Upper Limb 343
10.5 Muscles Acting on the Hip and
Lower Limb 359

Study Guide 375

Atlas B
Regional and Surface Anatomy 379
B.1
B.2
B.3

Regional Anatomy 380
The Importance of Surface
Anatomy 380
Learning Strategy 380

Chapter 11
Muscular Tissue 401
11.1 Types and Characteristics of
Muscular Tissue 402

Chapter 13
The Spinal Cord, Spinal Nerves, and
Somatic Reflexes 478
13.1 The Spinal Cord 479
13.2 The Spinal Nerves 487
13.3 Somatic Reflexes 500
Study Guide 508

Chapter 14
The Brain and Cranial Nerves 511
14.1 Overview of the Brain 512
14.2 Meninges, Ventricles, Cerebrospinal

Fluid, and Blood Supply 516
14.3 The Hindbrain and Midbrain 521
14.4 The Forebrain 528
14.5 Integrative Functions of the
Brain 534
14.6 The Cranial Nerves 546
Study Guide 558

Chapter 15
The Autonomic Nervous System and
Visceral Reflexes 561

Chapter 12
Nervous Tissue 439
12.1 Overview of the Nervous
System 440
12.2 Properties of Neurons 441
12.3 Supportive Cells (Neuroglia) 446
12.4 Electrophysiology of Neurons 451
12.5 Synapses 460
12.6 Neural Integration 466
Connective Issues 474
Study Guide 475

15.1 General Properties of the
Autonomic Nervous System 562
15.2 Anatomy of the Autonomic Nervous
System 565
15.3 Autonomic Effects on Target
Organs 572

15.4 Central Control of Autonomic
Function 577
Study Guide 579

Chapter 16
Sense Organs 582
16.1 Properties and Types of Sensory
Receptors 583
16.2 The General Senses 585
16.3 The Chemical Senses 591

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16.4 Hearing and Equilibrium 596
16.5 Vision 610
Study Guide 629

Chapter 17
The Endocrine System 633
17.1 Overview of the Endocrine
System 634
17.2 The Hypothalamus and Pituitary
Gland 637
17.3 Other Endocrine Glands 645
17.4 Hormones and Their Actions 655

17.5 Stress and Adaptation 665
17.6 Eicosanoids and Paracrine
Signaling 666
17.7 Endocrine Disorders 667
Connective Issues 674
Study Guide 675

18.3 Blood Types 691
18.4 Leukocytes 696
18.5 Platelets and Hemostasis—The
Control of Bleeding 702
Study Guide 711

Chapter 19
The Circulatory System: The Heart 714
19.1 Overview of the Cardiovascular
System 715
19.2 Gross Anatomy of the Heart 717
19.3 Cardiac Muscle and the Cardiac
Conduction System 725
19.4 Electrical and Contractile Activity of
the Heart 728
19.5 Blood Flow, Heart Sounds, and the
Cardiac Cycle 734
19.6 Cardiac Output 740
Study Guide 746

PART FOUR

Chapter 20

Regulation and Maintenance

The Circulatory System: Blood Vessels
and Circulation 749

Chapter 18
The Circulatory System: Blood 678
18.1 Introduction 679
18.2 Erythrocytes 684

20.1 General Anatomy of the Blood
Vessels 750
20.2 Blood Pressure, Resistance, and
Flow 758
20.3 Capillary Exchange 765
20.4 Venous Return and Circulatory
Shock 769
20.5 Special Circulatory Routes 771
20.6 Anatomy of the Pulmonary
Circuit 772
20.7 Systemic Vessels of the Axial
Region 773
20.8 Systemic Vessels of the
Appendicular Region 792
Connective Issues 803
Study Guide 804

Chapter 21
The Lymphatic and Immune

Systems 808
21.1 The Lymphatic System 809
21.2 Nonspecific Resistance 822
21.3 General Aspects of Specific
Immunity 830
21.4 Cellular Immunity 834
21.5 Humoral Immunity 837
21.6 Immune System Disorders 843
Connective Issues 849
Study Guide 850

Chapter 22
The Respiratory System 854
22.1 Anatomy of the Respiratory
System 855
22.2 Pulmonary Ventilation 866
22.3 Gas Exchange and Transport 877
22.4 Respiratory Disorders 887
Connective Issues 891
Study Guide 892

Chapter 23
The Urinary System 895
23.1 Functions of the Urinary
System 896
23.2 Anatomy of the Kidney 898
23.3 Urine Formation I: Glomerular
Filtration 904
23.4 Urine Formation II: Tubular
Reabsorption and Secretion 910

23.5 Urine Formation III: Water
Conservation 914
23.6 Urine and Renal Function Tests 918
23.7 Urine Storage and Elimination 920
Connective Issues 926
Study Guide 927

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