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Principles of

HUMAN
ANATOMY
12th Edition

Gerard J. Tortora
Bergen Community College

Mark T. Nielsen
University of Utah

John Wiley & Sons, Inc.


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VP & Publisher
Executive Editor
Executive Marketing Manager
Developmental Editor
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Project Editor
Contributing Editor
Program Assistant
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Production Editor
Senior Illustration Editors
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Cover Photo

Kaye Pace
Bonnie Roesch
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Lorraina Raccuia
Joan Kalkut
Lauren Morris
Dorothy Sinclair

Sandra Dumas
Anna Melhorn/Claudia Volano
Madelyn Lesure
Brian Salisbury
Hilary Newman
Ingrao Associates
Mark Nielsen

Page layout was completed by Laura Ierardi, LCI Design.
This book was typeset in 10/12 Janson at Aptara®, Inc. and printed and bound by R. R. Donnelley ( Jefferson
City). The cover was printed by R. R. Donnelley ( Jefferson City).
Founded in 1807, John Wiley & Sons, Inc. has been a valued source of knowledge and understanding for more than
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This book is printed on acid-free paper.
Copyright © 2012, 2009, 2005. © Gerard J. Tortora, Mark T. Nielsen and Biological Sciences Textbooks, Inc. © John Wiley and
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ISBN 13
ISBN 13

978-0470-56705-0
978-0470-91746-6

Printed in the United States of America.
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HELPING TEACHERS
AND STUDENTS
SUCCEED TOGETHER

Principles of Human Anatomy, twelfth edition, is designed
for introductory courses in human anatomy. The highly successful approach of previous editions—to provide students with an

accurate, clearly written, and expertly illustrated presentation of

the structure of the human body, to offer insights into the connections between structure and function, and to explore the practical and relevant applications of anatomical knowledge to everyday life
and career development—has been enhanced in this edition by innovations designed to increase student motivation and success.
An anatomy course can be the gateway to a satisfying career in a host of health-related professions.
It can also be incredibly challenging. We have designed the organization and flow of content within
these pages based on our deep experience teaching anatomy and interacting with students over
many years. We also understand the evolving dynamics of teaching and learning in today’s world.
That is why we are so pleased to partner with Wiley to create new and innovative ways to approach
the content digitally, using a research-proven design that promotes greater engagement, which leads
to improved learning outcomes.
Principles of Human Anatomy 12e, integrated with WileyPLUS, builds students’ confidence
because it takes the guesswork out of studying by providing students with a clear roadmap (what to
do, how to do it, if they did it right). Students will take more initiative so that instructors can have
greater impact.
On the following pages students will discover the tips and tools needed to make the most of their
time studying using text and media. An overview of the changes to this edition and insights into the
resources and support available to create dynamic classroom experiences as well as build meaningful
assessment opportunities are highlighted for instructors. Both students and instructors alike will be
interested in the additional resources available—Real Anatomy and a new Photographic Atlas of
Human Anatomy—both sure to enhance your insights into anatomy.
Years of experience, and listening to teachers and students like you, have helped us to create
solutions that work. We have worked hard to integrate the teaching process with the learning
environment—helping students and teachers succeed together.

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N OT E S

TO

The challenges of learning anatomy can be complex
and time consuming. This textbook and WileyPLUS for
Anatomy have been carefully designed to maximize
your time studying by simplifying the choices you
make in deciding what to study, how to study it, and
in assessing your understanding of the content.

S T U D E N T S
Anatomy Is a Visual Science
Studying the figures in this book is as important as reading the narrative. The tools described here will help you
understand the concepts being presented in any figure
and assure you get the most out of the visuals.

LEGEND Read this first. It explains what the figure is about.
KEY CONCEPT STATEMENT Indicated by a “key” icon, this reveals a basic idea portrayed in the figure.
ORIENTATION DIAGRAM Added to many figures, this small diagram helps you understand the perspective from which you are viewing a particular piece of anatomical art.
FIGURE QUESTIONS Found at the bottom
of each figure and accompanied by a “question mark” icon, these serve as a self-check to
help you understand the material as you go
along.

FUNCTIONS BOXES Included with selected
figures, these provide brief summaries of the
functions of the anatomical structure or system
depicted.

MP3 DOWNLOADS In each chapter you
will find that several illustrations are marked
with an icon that looks like an iPod. This
indicates that an audio file which narrates
and discusses the important elements of that
particular illustration is available. You can
access the downloads on the student companion website or within WileyPLUS.

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S T U D E N T S


There are many visual resources within WileyPLUS, in addition to the art from your text. These
can help you master the topic
you are studying. Examples
closely integrated with the reading material
include animations, cadaver video clips, and
Real Anatomy Views. Anatomy Drill and
Practice lets you test your knowledge of
structures with simple-to-use drag and drop
labeling exercises, or fill-in-the-blank labeling. You can drill and practice on these
activities using illustrations from the text,
cadaver photographs, histology micrographs, or lab models.

Clinical Connections
In some cases it is easier to understand the relevance of anatomical structures and the functions they support
by considering what happens when they don’t work the way they should. The Clinical Connections, which
appear throughout the text, present a variety of clinical perspectives
related to the text discussion.

WileyPLUS offers you
opportunities for even
further Clinical Connections with animated and interactive case studies that relate specifically to one body system or another. Look for these
under additional chapter resources as an interesting and engaging break from traditional study routines.

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S T U D E N T S

Exhibits Organize
Complex Anatomy into
Manageable Modules
Many topics in this text have been organized to
bring together all the anatomical information into
a simple-to-navigate content module. You will find
Exhibits for bones, joints, skeletal muscles, nerves,
blood vessels, and surface anatomy.

Objective to focus your study
Overview narrative of structure(s)
Table summarizing key features of structure(s)
Illustrations and photographs
Checkpoint question assesses your understanding
Clinical Connection provides relevance for learning
the details

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S T U D E N T S

Chapter Resources Help You Focus and Review
Your book has a variety of special features
that will make your time studying anatomy
a more rewarding experience. These
have been developed based on feedback from students—
like you—who have used previous editions of the text.
Their effectiveness is even further enhanced within
WileyPLUS.
Chapter Introductions set the stage for the content to
come and are followed by an interesting question that
always begins with “Did you ever wonder…?” These
questions will capture your interest and encourage you to
find the answer in the chapter material to come.
Objectives at the start of each section help you focus on
what is important as you read. All of the content within
WileyPLUS is tagged to these specific learning objectives

so that you can organize your study or review what is still
not clear in simple, more meaningful ways.
Checkpoint questions at the end of each section help
you assess if you have absorbed what you have read.
Take time to review these, or answer them within the
Practice section of each WileyPLUS concept module,
where they will automatically be graded to let you know
where you stand.

Mnemonics are a memory aid that can be particularly
helpful when learning specific anatomical features.
Mnemonics are included throughout the text, some displayed in figures, tables, or Exhibits and some included
within the text discussion. We encourage you not only to
use the mnemonics provided, but also to create your own
to help you learn the multitude of terms involved in your
study of human anatomy.
Key Medical Terms at the end of chapters include
selected terms dealing with both normal and pathological
conditions.
Chapter Review and Resource Summary is a helpful
table at the end of chapters that offers you a concise
summary of the important concepts from the chapter and
links each section to the media resources available in
WileyPLUS for Anatomy.
Self-Quiz Questions give you an opportunity to evaluate your understanding of the chapter as a whole. Within
WileyPLUS, use Progress Check to quiz yourself on individual or multiple chapters in preparation for exams or
quizzes.
Critical Thinking Questions are word problems that
allow you to apply the concepts you have studied in the
chapter to specific situations.


Mastering the Language of Anatomy
Throughout the text we have included Pronunciations
and, sometimes, Word Roots for many terms that may
be new to you. These appear in parentheses immediately
following the new words, and the pronunciations are
repeated in the glossary at the back of the book. Look at
the words carefully and say them out loud several times.
Learning to pronounce a new word will help you remember it and make it a useful part of your medical vocabulary. Take a few minutes to read the pronunciation key,
found at the beginning of the Glossary at the end of this
text (page G-1), so it will be familiar as you encounter
new words.
WileyPLUS houses help for you in building your new language skills as well. The Audio Glossary which is always
available to you lets you hear all these new, unfamiliar
terms pronounced. Throughout the e-text, these terms

can be clicked on and heard pronounced
as you read. In addition, you can use the
helpful Mastering Vocabulary program
which creates electronic flash cards for
you of the key terms within each chapter for practice,
as well as the ability to take a self-quiz specifically on
the terms introduced in each chapter.
To provide more assistance in learning the language of
anatomy, a full Glossary of terms with phonetic pronunciations appears at the end of the book. The basic building blocks of medical terminology—Combining Forms,
Word Roots, Prefixes, and Suffixes—are listed inside
the back cover, as is a listing of Eponyms, traditional
terms that include reference to a person’s name, along
with the current terminology.


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NOTES TO INSTRUCTORS
Collaborating on this revision has been a rewarding experience for us. We wanted to focus on the elements of the
text that we believed would benefit you and your students the most. We are gratified that many reviewer comments that helped us shape the changes we made matched so well with our intentions. Globally, we focused on
several key areas—the all-important visuals, both drawings and photographs; helping students relate what they
are learning to their desired career goals and the world around them by increasing the focus on Clinical
Connections; revising tables to increase their effectiveness in organizing detailed content; and making narrative
and organizational changes aimed at increasing student engagement with
the material.
For a detailed list of revisions for each chapter please visit our website at
www.wiley.com/college/sc/tortora and click on the text cover.

The Art of Anatomy
Illustrations throughout the text have
been refined. The color palette for the
skulls in Chapter 7, and for the brain and
spinal cord throughout the text, has been
adjusted for greater impact. Increased
clarity has been achieved in revised drawings of joints, muscles, blood vessels,
and regional lymph nodes. In addition, new origin–insertions figures

have been added to Chapter 11.

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NOTES TO INSTRUCTORS

Cadaver Photographs are included
throughout the text. Not only have we
increased the number of photographs
paired with illustrations, but a number of
new ones are the result of dissections
done under Mark Nielsen’s direction
specifically for this text revision.

LM

LM

400x

630x


Photomicrographs have also been replaced
throughout the text. See Chapter 3 for examples
of these stunning new photomicrographs with
enlargement blow outs.

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NOTES TO INSTRUCTORS
Chapter Beginnings and Ends
Each chapter has been effectively bookended with stunning new chapter introductions designed to
grab your student’s interest and engage them in the topic at hand, and redesigned chapter summaries
which now not only highlight the important concepts of the chapter, but point students to the media
resources that will support greater understanding of those concepts.

Clinical Connections
Your students are fascinated by the clinical connections to the normal anatomy that they
are learning. In response to reviewer feedback we have greatly expanded our use of these
boxed asides in this edition. You’ll find that the text is now liberally peppered with engaging discussions of a wide variety of clinical scenarios from disease coverage to tests and procedures. A complete reference list of the Clinical Connections within each chapter follows
the Table of Contents.


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NOTES TO INSTRUCTORS
WileyPLUS and You
WileyPLUS for Anatomy is an innovative, research-based online environment
designed for effective teaching and learning. Utilizing WileyPLUS in your
course provides your students with an accessible, affordable, and active learning platform and provides you tools and resources to efficiently build presentations for a dynamic classroom experience and to create and manage effective assessment strategies. The underlying principles of design, engagement,
and measurable outcomes provide the foundation for this powerful, new release of WileyPLUS.

DESIGN
• New research-based design helps students manage their time better and develop better study
skills.
• Course Calendars help track assignments for both students and teachers.
• New Course Plan makes it easier to assign readings, activities, and assessment. Simple dragand-drop tools make it easy to assign the course plan as-is or in any way that best reflects your
course syllabus.
The new design makes it easy for students to know what it is they need to do, boosting their confidence and preparing them for greater engagement in class and lab.

ENGAGEMENT
• Complete online version of the textbook for seamless integration of all content
• Relevant student study tools and learning resources ensure positive learning outcomes
• Immediate feedback boosts confidence and helps students see a return on investment for each

study session
• Precreated activities encourage learning outside of the classroom
• Course materials, including PowerPoint stacks that include animations and Wiley’s Visual Library
for Anatomy and Physiology, help you personalize lessons and optimize your time
Concept mastery in this discipline is directly related to students keeping up with the work and not
falling behind. The new Concept Modules, Activities, Self Study, and Progress Checks in WileyPLUS will
ensure that students know how to study effectively so they will remain engaged and stay on task.

MEASURABLE OUTCOMES
• Progress check enables students to hone in on areas of weakness for increased success.
• Self-assessment and remediation for all learning objectives lets students know exactly how their
efforts have paid off.
• Instant reports monitor trends in class performance, use of course materials, and student progress
toward learning objectives.
With new detailed reporting capabilities students will know that they are doing it right. With
increased confidence, motivation is sustained so students stay on task, and success will follow.
Please contact your Wiley representative for details about these and other resources or visit our website at
www.wiley.com/college/sc/tortora and click on the text cover to explore the assets more fully.

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ADDITIONAL RESOURCES
Real Anatomy
Mark Nielsen and Shawn Miller, University of Utah
Real Anatomy is 3-D imaging software that allows you to dissect through
multiple layers of a three-dimensional real human body to study and learn
the anatomical structures of all body systems.

• Dissect through up to 40 layers of the
body and discover the relationships of the
structures to the whole.
• Rotate the body as well as major
organs to view the image from multiple
perspectives.

• Use a built-in zoom feature to get a closer look at
detail.
• A unique approach to highlighting and labeling
structures does not obscure the real anatomy in
view.

• Related Images provide multiple views of
structures being studied.

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ADDITIONAL RESOURCES

• Snapshots can be saved
of any image for use in
PowerPoints, quizzes, or
handouts.

• View histology micrographs at varied levels
of magnification with
the virtual microscope.

• Audio pronunciation of all
labeled structures is readily
available.

Virtual Dissection—100% Real

NEW! THE PERFECT COMPANION TO COMPLETE YOUR STUDY
OF ANATOMY

Photographic Atlas of
Human Anatomy, First Edition
Mark Nielsen and Shawn Miller, University of Utah
This new atlas filled with outstanding photographs of meticulously executed
dissections of the human body has been developed to be a strong teaching
and learning solution, not just a catalog

of photographs. Organized around body
systems, each chapter includes a narrative
overview of the body system followed by
detailed photographs that accurately and
realistically represent the anatomical
structures. Histology is included.
Photographic Atlas of Human Anatomy
will work well in your laboratories as a
study companion to your textbook
and as a print companion to the Real
Anatomy DVD.

Like the respiratory and digestive systems, the urinary system is an environmental exchange system. Like all the exchange systems of the body, the urinary system forms an immense interface with the cardiovascular system for the single purpose of regulating the homeostatic balance of the water environment (extracellular matrix) that
surrounds every cell in the body. To make this exchange possible a large network of microscopic urinary tubes form an intimate
nary system consists of two blood processing centers called the kidneys, two transport tubes called the ureters, which move the urine,

that serves as a storage organ to hold the urine, which
is being constantly produced in the kidneys. When
it is convenient to remove the stored urine from
the body, it leaves the bladder through a single
drainage tube called the urethra.
In order to survive, every body cell
requires a water environment that is similar to the composition of the oceans in
neys help maintain this water environblood and regulating its contents so
the blood can help maintain the correct composition of the extracellular
ing the amount of water in the plasma
and the various plasma constituents,
which are either conserved for the
body or eliminated in the urine, the
kidneys are able to maintain water

and electrolyte balance within the very
narrow range compatible with life,
despite wide variations in intake and
losses of these constituents through
other avenues.

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ACKNOWLEDGMENTS
We wish to especially thank several academic colleagues for their helpful contributions to this edition. Creating and
implementing the integration of this text
with WileyPLUS for Anatomy were
possible only because of the expertise
and fine work of the following group of
people. We are very grateful to you:
Matthew Abbott
Des Moines Area Community College
Kathleen Andersen
University of Iowa
Deborah Canepa
Linfield College

Martha Dixon
Diablo Valley College
Heather Dy
Long Beach City College
Wanda Hargroder
Louisiana State University
Jacki Houghton
Moorpark College
Jon Hubbard
Hartnell College
Eric Lippincott
Lock Haven University
Izak Paul
Mount Royal University
Susan Rohde
Triton College
Sara Tolsma
Northwestern College
We are also very grateful to our colleagues who have reviewed the manuscript or participated in focus groups and
offered numerous suggestions for
improvement.
Matthew Abbott
Des Moines Area Community College
Mark Alston
University of Tennessee–Knoxville
David Babb
West Hills Community College
Debra Barnes
Contra Costa College
Kristen Bruzzini

Maryville University
Christine Byrd-Jacobs
Western Michigan University

xiv

Travis H. Carlton
Marshall University
Rosalee Carter
California State University Sacramento
Carl Christensen
Austin Community College
Lori Coble
South Dakota School of Mines & Technology
Richard Connett
Monroe Community College–Brighton
Sarah Cotton
Chaffey College
Martha Dixon
Diablo Valley College
Joel Gluck
Community College of Rhode Island
Melanie Gouzoules
University of North Carolina–Greensboro
Wanda Hardgroder
Louisiana State University
Cynthia Herbrandson
Kellogg Community College
Jane Horlings
Saddleback College

Randy Howell
Lock Haven University
Jon Hubbard
Hartnell College
Kelly Johnson
University of Kansas
Jeffrey S. Kiggins
Monroe Community College
Robert Knudsen
San Joaquin Delta College
Eric Lippincott
Lock Haven University
Shawn Miller
University of Utah
Jennifer Maze
Lander University
Angela Miller
Northwestern State University
Heather Moore
College of the Sequoias
Virginia Naples
Northern Illinois University
Gail Patt
Boston University
Ruth Peterson
Hennepin Technical College

Rachel D. Smetanka
Southern Utah University
Robert Stark

California State University Bakersfield
Pamela Stein
AEC Texas Institute
Karah Street
Saddleback College
Christa Voss
Tulsa Community College
Anthony J. Weinhaus
University of Minnesota
Kira L. Wennstrom
Shoreline Community College
Andrzej Wierasko
CUNY Staten Island
John Wilkins
Ball State University
Brian Wisenden
Minnesota State University Moorhead
David A. Woodman
University of Nebraska–Lincoln
Michael Yard
Indiana University, Purdue University
Indianapolis
Scott Zimmerman
Missouri State University
Finally, our hats are off to everyone at
Wiley. We enjoy collaborating with this
enthusiastic, dedicated, and talented team of
publishing professionals. Our thanks to the
entire team—Bonnie Roesch, Executive
Editor; Karen Trost, Developmental Editor;

Lorraina Raccuia, Project Editor; Lauren
Morris, Program Assistant; Suzanne Ingrao,
Outside Production Manager; Hilary
Newman, Photo Manager; Claudia Volano,
Illustration Coordinator; Madelyn Lesure,
Designer; Laura Ierardi, LCI Design; and
Clay Stone, Executive Marketing Manager.
GERARD J. TORTORA
Department of Biology and Horticulture, S229
Bergen Community College
400 Paramus Road
Paramus, NJ 07652

MARK NIELSEN
Department of Biology
University of Utah
257 South 1400 East
Salt Lake City, UT 84112


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ABOUT THE AUTHORS
Jerry Tortora is Professor of Biology and former Biology Coordinator at Bergen Community College in

Paramus, New Jersey, where he teaches human anatomy and physiology as well as microbiology. He received
his bachelor’s degree in biology from Fairleigh Dickinson University and his master’s degree in science education from Montclair State College. He is a member of many professional organizations, including the Human
Anatomy and Physiology Society (HAPS), the American Society of Microbiology (ASM), American
Association for the Advancement of Science (AAAS), National Education Association (NEA), and the
Metropolitan Association of College and University Biologists (MACUB).
Above all, Jerry is devoted to his students and their aspirations. In recognition of this commitment, Jerry
was the recipient of MACUB’s 1992 President’s Memorial Award. In 1996, he received a National Institute for
Staff and Organizational Development (NISOD) excellence award from the University of Texas and was selected
to represent Bergen Community College in a campaign to increase awareness of the contributions of community colleges to higher education.
Courtesy of Heidi Chung.
Jerry is the author of several best-selling science textbooks and laboratory manuals, a calling that often requires
an additional 40 hours per week beyond his teaching responsibilities. Nevertheless, he still makes time for four or five weekly aerobic
workouts that include biking and running. He also enjoys attending college basketball
and professional hockey games and performances at the Metropolitan Opera House.

To my mother, Angelina M. Tortora
(August 20, 1913–August 14, 2010).
Her love, guidance, faith, support, and example will always be the
cornerstones of my personal and professional life. G.J.T.

Mark Nielsen is a Professor in the Department of Biology at the University of Utah
and for the past twenty-four years has taught anatomy and its related subjects to over
18,000 students. In addition to teaching human anatomy in the Department of Biology,
he also teaches neuroantomy, embryology, a human dissection course, a teaching human
anatomy course, and assists with the comparative vertebrate morphology course. He
developed the anatomy course for the physican assistant program at the University of
Utah School of Medicine, where he taught for five years, and taught in the cadaver lab
at the University of Utah School of Medicine. He developed and continues to help
maintain the anatomy and physiology program for the Utah College of Massage
Therapy (presently the largest massage school in the United States) and taught his

program there for twelve years during its inception and development. His graduate
training is in anatomy and his anatomy expertise has a strong basis in dissection. He has
prepared and participated in hundreds of dissections of both humans and other vertebrate animals. All his courses incorporate a cadaverbased component to the training with an outstanding exposure to cadaver anatomy. He is a member of the American Association of
Anatomists (AAA), the Human Anatomy and Physiology Society (HAPS), and the Anatomical Society of Great Britain and Ireland
(ASGBI).
Mark has a passion for teaching anatomy and sharing his knowledge with his students. In addition to the many students he has taught
anatomy, he has trained and mentored 950 students who have worked in his anatomy laboratory as teaching assistants. His concern for
students and his teaching excellence have been acknowledged through numerous awards. He received the prestigous Presidential
Teaching Scholar Award at the University of Utah, is a five-time recipient of the University of Utah Student Choice Award for
Outstanding Teacher and Mentor, a two-time winner of the Outstanding Teacher in the Physician Assistant Program, recipient of the
American Massage Therapy Association Jerome Perlinski Teacher of the Year Award, and a two-time recipient of Who’s Who Among
America’s Teachers.

To my academic mentors Professors John Legler and Dennis Bramble.
I was lucky to be nurtured academically by two great anatomists, scholars, and teachers. Thank you. I can
never thank you enough or repay you for your contributions to my academic career. M.T.N.

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BRIEF CONTENTS
1 AN INTRODUCTION TO THE HUMAN BODY
2 CELLS
3 TISSUES
4 DEVELOPMENT
5 THE INTEGUMENTARY SYSTEM
6 BONE TISSUE
7 THE SKELETAL SYSTEM: THE AXIAL SKELETON
8 THE SKELETAL SYSTEM: THE APPENDICULAR SKELETON
9 JOINTS
10 MUSCULAR TISSUE
11 THE MUSCULAR SYSTEM
12 THE CARDIOVASCULAR SYSTEM: BLOOD
13 THE CARDIOVASCULAR SYSTEM: THE HEART
14 THE CARDIOVASCULAR SYSTEM: BLOOD VESSELS
15 THE LYMPHATIC SYSTEM AND IMMUNITY
16 NERVOUS TISSUE
17 THE SPINAL CORD AND THE SPINAL NERVES
18 THE BRAIN AND THE CRANIAL NERVES
19 THE AUTONOMIC NERVOUS SYSTEM
20 SOMATIC SENSES AND MOTOR CONTROL
21 SPECIAL SENSES
22 THE ENDOCRINE SYSTEM
23 THE RESPIRATORY SYSTEM
24 THE DIGESTIVE SYSTEM
25 THE URINARY SYSTEM
26 THE REPRODUCTIVE SYSTEMS
27 SURFACE ANATOMY


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174
227
264
304
332
440
458
488
547
575
594
623
674
697
717
753
778
811
857
884
932

APPENDIX A: MEASUREMENTS


A-1

APPENDIX B: ANSWERS

B-1

GLOSSARY

G-1

CREDITS

C-1

INDEX

I-1

xvii


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1 AN INTRODUCTION TO THE HUMAN

3 TISSUES

BODY 2

3.1 Types of Tissues 63
3.2 Cell Junctions 64
Tight Junctions 65
Adherens Junctions 65
Desmosomes 65
Hemidesmosomes 65
Gap Junctions 65
3.3 Comparison Between Epithelial and Connective
Tissues 66
3.4 Epithelial Tissue 66
Classification of Epithelial Tissues 67
Covering and Lining Epithelium 69
Glandular Epithelium 74

1.1
1.2
1.3
1.4

Anatomy Defined 3
Levels of Body Organization and Body Systems 4
Life Processes 10
Basic Anatomical Terminology 10


Anatomical Position 10
Regional Names 10
Planes and Sections 12
Overview 14

1.5 Body Cavities 16
Thoracic and Abdominal Cavity Membranes 17

1.6
1.7
1.8
1.9

Abdominopelvic Regions and Quadrants 20
The Human Body and Disease 21
Medical Imaging 22
Measuring the Human Body 25

Chapter Review and Resource Summary 25 / Self-Quiz
Questions 26 / Critical Thinking Questions 27 /
Answers to Figure Questions 27

62

Structural Classification of Exocrine Glands 74
Functional Classification of Exocrine Glands 75

3.5 Connective Tissue 76
General Features of Connective Tissue 77
Connective Tissue Cells 77

Connective Tissue Extracellular Matrix 78
Ground Substance 78 / Fibers 79

2 CELLS

28

2.1 A Generalized Cell 29
2.2 The Plasma Membrane 30
Structure of the Membrane 30
Functions of Membrane Proteins 31
Membrane Permeability 31
Transport Across the Plasma Membrane 31
Kinetic Energy Transport 32 / Transport by Transporter
Proteins 32 / Transport in Vesicles 32
2.3 Cytoplasm 35
Cytosol 35
Organelles 36
Centrosome 37 / Cilia and Flagella 37 / Ribosomes 38
Endoplasmic Reticulum 39 / Golgi Complex 39 / Lysosomes 41
Peroxisomes 42 / Proteasomes 42 / Mitochondria 42

2.4 Nucleus 44
2.5 Cell Division 47
Somatic Cell Division 47
Interphase 47
Mitotic Phase 48
Control of Cell Destiny 50
Reproductive Cell Division 51
Meiosis 51


2.6 Cellular Diversity 54
2.7 Aging and Cells 56
Key Medical Terms Associated with Cells 57 / Chapter Review
and Resource Summary 57 / Self-Quiz Questions 59 / Critical
Thinking Questions 61 / Answers to Figure Questions 61

xviii

Classification of Connective Tissues 79
Embryonic Connective Tissue 79
Mature Connective Tissue 80
Loose Connective Tissue 80
Dense Connective Tissue 83
Cartilage 84
Bone Tissue 86
Liquid Connective Tissue 86

3.6 Membranes 88
Epithelial Membranes 88
Mucous Membranes 88 / Serous Membranes 88
Cutaneous Membrane 88
Synovial Membranes 88

3.7 Muscular Tissue 90
3.8 Nervous Tissue 92
3.9 Aging and Tissues 92
Key Medical Terms Associated with Tissues 93 / Chapter Review
and Resource Summary 93 / Self-Quiz Questions 95 / Critical
Thinking Questions 97 / Answers to Figure Questions 97


4 DEVELOPMENT

98

4.1 Embryonic Period 100
First Week of Development 100
Fertilization 100 / Cleavage of the Zygote 101 / Blastocyst
Formation 101 / Implantation 102
Second Week of Development 104
Development of the Trophoblast 104 / Development of the
Bilaminar Embryonic Disc 104 / Development of the


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Amnion 104 / Development of the Yolk Sac 104 / Development
of Sinusoids 106 / Development of the Extraembryonic
Coelom 106 / Development of the Chorion 106
Third Week of Development 106
Gastrulation 106 / Neurulation 108 / Development of
Somites 109 / Development of the Intraembryonic
Coelom 110 / Development of the Cardiovascular

System 110 / Development of the Chorionic Villi
and Placenta 110
Fourth Week of Development 112
Fifth Through Eighth Weeks of Development 114

4.2 Fetal Period 115
4.3 Maternal Changes During Pregnancy 117
4.4 Labor 117

123

5.1 Structure of the Skin 124
Epidermis 124
Stratum Basale 126 / Stratum Spinosum 128 / Stratum
Granulosum 128 / Stratum Lucidum 128 / Stratum
Corneum 128 / Keratinization and Growth of the
Epidermis 128
Dermis 129
The Structural Basis of Skin Color 130 / Tattooing and Body
Piercing 131
Subcutaneous Layer or Hypodermis 131

5.2 Accessory Structures of the Skin 132
Hair 132
Anatomy of a Hair 133 / Hair Growth 135 / Types of
Hairs 135 / Hair Color 135
Skin Glands 136
Sebaceous Glands 136 / Sudoriferous Glands 137 /
Ceruminous Glands 137
Nails 138


5.3 Types of Skin 140
5.4 Function of the Skin 140
5.5 Blood Supply of the Integumentary
System 141
5.6 Development of the
Integumentary System 143
5.7 Aging and the Integumentary
System 145
Key Medical Terms Associated with the
Integumentary System 145 / Chapter
Review and Resource Summary 146 / SelfQuiz Questions 148 / Critical Thinking
Questions 149 / Answers to Figure Questions 149

6 BONE TISSUE

6.3 Anatomy of a Bone 152
6.4 Bone Surface Markings 153
6.5 Histology of Bone Tissue 154
Compact Bone Tissue 156
Spongy Bone Tissue 158
6.6 Blood and Nerve Supply of Bone 158
6.7 Bone Formation 159
Initial Bone Formation in an Embryo and Fetus 159
Intramembranous Ossification 159
Endochondral Ossification 161
Bone Growth During Infancy, Childhood, and
Adolescence 162
Growth in Length 162 / Growth in Thickness 163
Remodeling of Bone 163


Key Medical Terms Associated with Development 119 / Chapter
Review and Resource Summary 120 / Self-Quiz Questions 121 /
Critical Thinking Questions 122 / Answers to Figure Questions 122

5 THE INTEGUMENTARY SYSTEM

xix

150

6.1 Functions of Bone and the Skeletal System 151
6.2 Types of Bones 151

6.8 Fractures 166
6.9 Exercise and Bone Tissue 169
6.10 Aging and Bone Tissue 169
6.11 Factors Affecting Bone Growth 169
Key Medical Terms Associated with Bone Tissue 171 /
Chapter Review and Resource Summary 171 / Self-Quiz
Questions 172 / Critical Thinking Questions 173 / Answers
to Figure Questions 173

7 THE SKELETAL SYSTEM: THE AXIAL
SKELETON 174
7.1 Divisions of the Skeletal System 175
7.2 Skull 177
General Features and Functions 177
Nasal Septum 198
Orbits 198

Foramina 200
Unique Features of the Skull 200
Sutures 200 / Paranasal Sinuses 201 /
Fontanels 201
Cranial Fossae 204
Age-related Changes in the Skull 204
Sexual Differences in the Skull 205

7.3 Hyoid Bone 206
7.4 Vertebral Column 207
Normal Curves of the Vertebral Column 207
Intervertebral Discs 207
Parts of a Typical Vertebra 209
Vertebral Body 209 / Vertebral Arch 209 / Processes 210
Regions of the Vertebral Column 211
Age-related Changes in the Vertebral Column 211

7.5 Thorax 220
Key Medical Terms Associated with the Axial Skeleton 224 /
Chapter Review and Resource Summary 224 / Self-Quiz
Questions 225 / Critical Thinking Questions 226 / Answers to
Figure Questions 226

8 THE SKELETAL SYSTEM: THE
APPENDICULAR SKELETON 227
8.1 Skeleton of the Upper Limb 228


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8.3
8.4
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Skeleton of the Lower Limb 242
False and True Pelves 247
Comparison of Female and Male Pelves 248
Comparison of Pectoral and Pelvic Girdles 248
Development of the Skeletal System 259

Key Medical Terms Associated with Appendicular Skeleton 261
Chapter Review and Resource Summary 261
Self-Quiz Questions 262 / Critical Thinking Questions 263
Answers to Figure Questions 263

9 JOINTS

264


9.1 Joint Classifications 265
Ligaments 266
9.2 Fibrous Joints 266
Sutures 266
Syndesmoses 266
Interosseous Membranes 266
9.3 Cartilaginous Joints 268
Synchondroses 268
Symphyses 268
9.4 Synovial Joints 268
Structure of Synovial Joints 268
Articular Capsule 269 / Synovial Fluid 269 / Accessory
Ligaments, Articular Discs, and Labra 270 / Nerve and Blood
Supply 270
Bursae and Tendon Sheaths 271
Types of Synovial Joints 271
Plane Joints 271 / Hinge Joints 271 / Pivot Joints 271 /
Condyloid Joints 271 / Saddle Joints 271 / Ball-and-Socket
Joints 271

9.5 Types of Movements at Synovial Joints 274
Gliding 274
Angular Movements 274
Flexion, Extension, Lateral Flexion, and Hyperextension 274 /
Abduction, Adduction, and Circumduction 274
Rotation 276
Special Movements 277

9.6 Factors Affecting Contact and Range of Motion at
Synovial Joints 277

9.7 Selected Joints of the Body 280
9.8 Aging and Joints 298
Key Medical Terms Associated with Joints 299
Chapter Review and Resource Summary 299
Self-Quiz Questions 301 / Critical Thinking Questions 302
Answers to Figure Questions 302

10 MUSCULAR TISSUE

304

10.1 Overview of Muscular Tissue 305
Types of Muscular Tissue 305
Functions of Muscular Tissue 305
Properties of Muscular Tissue 305
10.2 Skeletal Muscular Tissue 306
Structure of a Skeletal Muscle 306

Connective Tissue Coverings 306
Nerve and Blood Supply 308
Microscopic Anatomy of a Skeletal Muscle Fiber (Cell) 308
Sarcolemma, T Tubules, and Sarcoplasm 308 / Myofibrils
and Sarcoplasmic Reticulum 310 / Filaments and the
Sarcomere 310
Muscle Proteins 310

Contraction and Relaxation of Skeletal Muscle Fibers 314
Sliding Filament Mechanism 314 / The Neuromuscular
Junction 314 / The Contraction Cycle 317 /
Excitation–Contraction Coupling 318

Muscle Tone 319
Isotonic and Isometric Contractions 319

10.3 Types of Skeletal Muscle Fibers 320
Slow Oxidative Fibers 320
Fast Oxidative-Glycolytic Fibers 320
Fast Glycolytic Fibers 320
10.4 Exercise and Skeletal Muscle Tissue 321
Effective Stretching 322
Strength Training 322
10.5 Cardiac Muscle Tissue 322
10.6 Smooth Muscle Tissue 324
10.7 Development of Muscles 326
10.8 Aging and Muscular Tissue 326
Key Medical Terms Associated with Muscular Tissue 327
Chapter Review and Resource Summary 328
Self-Quiz Questions 329 / Critical Thinking Questions 330
Answers to Figure Questions 331

11 THE MUSCULAR SYSTEM

332

11.1 How Skeletal Muscles Produce
Movements 333
Muscle Attachment Sites: Origin and
Insertion 333
Lever Systems 334
Effects of Fascicle Arrangement 336
Muscle Actions 337

Coordination Among Muscles 337
Structure and Function of Muscle
Groups 337
11.2 How Skeletal Muscles are Named 338
11.3 Principal Skeletal Muscles 338
Key Medical Terms Associated with the Muscular System 436 /
Chapter Review and Resource Summary 436 / Self-Quiz
Questions 437 / Critical Thinking Questions 438 / Answers to
Figure Questions 439

12 THE CARDIOVASCULAR SYSTEM:
BLOOD 440
12.1 Functions of Blood 441
12.2 Physical Characteristics of Blood 441


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12.3 Components of Blood 441
Blood Plasma 443
Formed Elements 444
12.4 Formation of Blood Cells 445

12.5 Red Blood Cells 447
RBC Anatomy 447
RBC Functions 447
RBC Life Cycle 449
Erythropoiesis: Production of RBCs 449
Blood Group Systems 449
12.6 White Blood Cells 450
WBC Anatomy and Types 450
Granular Leukocytes 451 / Agranular Leukocytes 451

WBC Functions 451
12.7 Platelets 452
12.8 Stem Cell Transplants from Bone Marrow
and Cord-Blood 453
Key Medical Terms Associated with Blood 454
Chapter Review and Resource Summary 455
Self-Quiz Questions 456 / Critical Thinking Questions 457
Answers to Figure Questions 457

13 THE
CARDIOVASCULAR
SYSTEM:
THE HEART 458
13.1 Location and Surface
Projection of the Heart 459
13.2 Structure and Function of the
Heart 461
Pericardium 461
Layers of the Heart Wall 463
Chambers of the Heart 464

Right Atrium 464 / Right Ventricle 464 / Left Atrium 466 /
Left Ventricle 468
Myocardial Thickness and Function 468
Fibrous Skeleton of the Heart 468
Heart Valves 468
Atrioventricular Valves 469 / Semilunar Valves 469

13.3 Circulation of Blood 471
Systemic and Pulmonary Circulations 471
Coronary Circulation 471
Coronary Arteries 472 / Coronary Veins 474

13.4 Cardiac Conduction System and Innervation 475
Cardiac Conduction System 475
Cardiac Nerves 476
13.5 Cardiac Cycle (Heartbeat) 477
13.6 Heart Sounds 478
13.7 Exercise and the Heart 478
13.8 Development of the Heart 483
Key Medical Terms Associated with The Heart 484
Chapter Review and Resource Summary 485
Self-Quiz Questions 486 / Critical Thinking Questions 487
Answers to Figure Questions 487

xxi

14 THE CARDIOVASCULAR
SYSTEM: BLOOD VESSELS 488
14.1 Anatomy of Blood Vessels 489
Basic Structure of a Blood Vessel 489

Tunica Interna (Intima) 489 / Tunica Media 491 / Tunica
Externa 491
Arteries 491
Elastic Arteries 491 / Muscular
Arteries 491
Anastomoses 492
Arterioles 492
Capillaries 492
Venules 494
Veins 494
Blood Distribution 496

14.2 Circulatory Routes 497
Systemic Circulation 497
The Hepatic Portal Circulation 537
The Pulmonary Circulation 538
The Fetal Circulation 538
14.3 Development of Blood Vessels and Blood 542
14.4 Aging and the Cardiovascular System 542
Key Medical Terms Associated with Blood Vessels 544
Chapter Review and Resource Summary 544
Self-Quiz Questions 545 / Critical Thinking Questions 546
Answers to Figure Questions 546

15 THE LYMPHATIC
SYSTEM AND
IMMUNITY 547
15.1 Lymphatic System Structure
and Functions 548
Functions of the Lymphatic

System 548
Lymphatic Vessels and Lymph
Circulation 550
Lymphatic Capillaries 551 / Lymph Trunks
and Ducts 551 / Formation and Flow of Lymph 552
Lymphatic Organs and Tissues 553
Thymus 553 / Lymph Nodes 555 / Spleen 557 / Lymphatic
Nodules 559

15.2 Principal Groups of Lymph Nodes 559
15.3 Development of Lymphatic Tissues 570
15.4 Aging and the Lymphatic System 570
Key Medical Terms Associated with Lymphatic System and
Immunity 572
Chapter Review and Resource Summary 572
Self-Quiz Questions 573 / Critical Thinking Questions 574
Answers to Figure Questions 574

16 NERVOUS TISSUE

575

16.1 Overview of the Nervous System 576
Structures of the Nervous System 576


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Organization of the Nervous System 576
Anatomical Organization 576 / Functional Organization 577

16.2 Histology of Nervous Tissue 579
Neurons 579
Parts of a Neuron 579 / Cell Body 579 / Nerve Fibers 580

Synapses 581
Neuromuscular Junction 582 / Synapses Between Neurons
582 / Neurotransmitters 582 / Structural Diversity in
Neurons 583
Neuroglia 584
Neuroglia of the CNS 584 / Neuroglia of the PNS 586
Myelination 586
Gray and White Matter 587

16.3 Neural Circuits 589
16.4 Regeneration and Neurogenesis 590
Key Medical Terms Associated with Nervous Tissue 591 /
Chapter Review and Resource Summary 591 / Self-Quiz
Questions 592 / Critical Thinking Questions 593 / Answers to
Figure Questions 593


17 THE SPINAL CORD
AND THE SPINAL
NERVES 594
17.1 Spinal Cord Anatomy 595
Protective Structures 595
Vertebral Column 595 / Meninges 595

External Anatomy of the
Spinal Cord 595
Internal Anatomy of the Spinal Cord 599
17.2 Spinal Nerves 602
Structure of a Single Nerve 602
Organization of Spinal Nerves 603
Branches of Spinal Nerves 604
Plexuses 604 / Intercostal Nerves 616

Dermatomes Versus Cutaneous Fields 616
17.3 Spinal Cord Functions 616
Sensory and Motor Tracts 616
Reflexes and Reflex Arcs 618
Key Medical Terms Associated with the Spinal Cord and the
Spinal Nerves 619 / Chapter Review and Resource Summary 620 /
Self-Quiz Questions 621 / Critical Thinking Questions 622
Answers to Figure Questions 622

18 THE BRAIN AND THE CRANIAL
NERVES 623
18.1 Development and General Structure of the Brain 624
Brain Development 624

Major Parts of the Brain 624
18.2 Protection and Blood Supply 627
Protective Coverings of the Brain 627
Cerebrospinal Fluid 627
Formation of CSF in the Ventricles 627 / Functions of
CSF 627 / Circulation of CSF 627
Brain Blood Flow and the Blood–Brain Barrier 631

18.3 The Brain Stem and Reticular Formation 634
Medulla Oblongata 634
Pons 636
Midbrain 636
Reticular Formation 637
18.4 The Cerebellum 639
18.5 The Diencephalon 641
Thalamus 641
Hypothalamus 642
Epithalamus 643
Circumventricular Organs 643
18.6 The Cerebrum 644
Structure of the Cerebrum 644
Cerebral White Matter 646
Basal Nuclei 646
The Limbic System 648
18.7 Functional Organization of the Cerebral Cortex 650
Sensory Areas 650
Motor Areas 651
Association Areas 651
Hemispheric Lateralization 653
Memory 653

Brain Waves 653
18.8 Aging and the Nervous System 654
18.9 Cranial Nerves 654
Key Medical Terms Associated with the Brain and the Cranial
Nerves 669 / Chapter Review and Resource Summary 669
Self-Quiz Questions 671 / Critical Thinking Questions 673
Answers to Figure Questions 673

19 THE AUTONOMIC NERVOUS
SYSTEM 674
19.1 Comparison of Somatic and Autonomic
Nervous Systems 675
Somatic Nervous System 675
Autonomic Nervous System 675
Comparison of Somatic and Autonomic
Motor Neurons 676
19.2 Anatomy of Autonomic Motor Pathways 677
Understanding Autonomic Motor Pathways 677
Migration of the Neural Crest Tissue 678

Shared Anatomical Components of an Autonomic
Motor Pathway 679
Motor Neurons and Autonomic Ganglia 679 /
Autonomic Plexuses 679

19.3 Structure of the Sympathetic Division 681
Sympathetic Preganglionic Neurons 681
Sympathetic Ganglia and Postganglionic Neurons 684
Sympathetic Trunk Ganglia 684 / Prevertebral Ganglia 684


19.4 Structure of the Parasympathetic Division 685
Parasympathetic Preganglionic Neurons 685
Parasympathetic Ganglia and Postganglionic Neurons 685
19.5 Structure of the Enteric Division 687
19.6 ANS Neurotransmitters and Receptors 687
Cholinergic Neurons and Receptors 687


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Adrenergic Neurons and Receptors 688
19.7 Functions of the ANS 688
Sympathetic Responses 689
Parasympathetic Responses 690
19.8 Integration and Control of Autonomic Functions 692
Autonomic Reflexes 692
Autonomic Control by Higher Centers 692
Key Medical Terms Associated with the Autonomic Nervous
System 693 / Chapter Review and Resource
Summary 693 / Self-Quiz Questions 695 /
Critical Thinking Questions 696 /
Answers to Figure Questions 696


20 SOMATIC SENSES
AND MOTOR
CONTROL 697
20.1 Overview of Sensations 698
Definition of Sensations 698
Characteristics of Sensations 698
Classification of Sensations 698
Types of Sensory Receptors 698
20.2 Somatic Sensations 699
Tactile Sensations 699
Touch 699 / Pressure and Vibration 700 / Itch and Tickle 700

Thermal Sensations 701
Pain Sensations 701
Types of Pain 701 / Localization of Pain 701

Proprioceptive Sensations 701
Muscle Spindles 702 / Tendon Organs 703 / Joint Kinesthetic
Receptors 703

20.3 Somatic Sensory Pathways 704
Posterior Column–Medial Lemniscus Pathway to the
Cortex 705
Anterolateral Pathways to the Cortex 706
Mapping the Primary Somatosensory Area 706
Somatic Sensory Pathways to the Cerebellum 706
20.4 Somatic Motor Pathways 707
Mapping the Motor Areas 709
Direct Motor Pathways 709

Indirect Motor Pathways 710
Roles of the Basal Nuclei 710
Roles of the Cerebellum 711
20.5 Integration of Sensory Input and Motor Output 712
Key Medical Terms Associated with Somatic Senses and Motor
Control 713 / Chapter Review and Resource Summary 713 /
Self-Quiz Questions 714 / Critical Thinking Questions 716 /
Answers to Figure Questions 716

21 SPECIAL SENSES

717

21.1 Olfaction: Sense of Smell 718
Anatomy of Olfactory Receptors 719
The Olfactory Pathway 720
21.2 Gustation: Sense of Taste 720
Anatomy of Gustatory Receptors 720

xxiii

The Gustatory Pathway 722
21.3 Vision 722
Accessory Structures of the Eye 722
Eyelids 722 / Eyelashes and Eyebrows 724 / The Lacrimal
Apparatus 724 / Extrinsic Eye Muscles 724
Anatomy of the Eyeball 724
Fibrous Tunic 724 / Vascular Tunic 724 / Retina 726 /
Lens 729 / Interior of the Eyeball 729
The Visual Pathway 730

Processing of Visual Input in the Retina 731 / Pathway in the
Brain 731

21.4 Hearing and Equilibrium 732
Anatomy of the Ear 732
External (Outer) Ear 732 / Middle Ear 733 / Internal (Inner)
Ear 734
Mechanism of Hearing 739
The Auditory Pathway 740
Mechanism of Equilibrium 740
Otolithic Organs: Saccule and Utricle 741 / Semicircular
Ducts 741
Equilibrium Pathways 741

21.5 Development of the Eyes and Ears 746
Development of the Eyes 746
Development of the Ears 747
21.6 Aging and the Special Senses 748
Key Medical Terms Associated with Special Senses 748 / Chapter
Review and Resource Summary 749 / Self-Quiz Questions 750 /
Critical Thinking Questions 752 / Answers to Figure
Questions 752

22 THE ENDOCRINE SYSTEM

753

22.1 Endocrine Glands Defined 754
22.2 Hormones 754
22.3 Hypothalamus and Pituitary Gland 756

Anterior Pituitary 757
Posterior Pituitary 758
22.4 Pineal Gland and Thymus 761
22.5 Thyroid Gland and Parathyroid Glands 761
22.6 Adrenal Glands 765
Adrenal Cortex 766
Adrenal Medulla 766
22.7 Pancreas 768
22.8 Ovaries and Testes 770
22.9 Other Endocrine Tissues 772
22.10 Development of the
Endocrine System 772
22.11 Aging and the Endocrine
System 773
Key Medical Terms Associated with the
Endocrine System 774 / Chapter Review and Resource
Summary 774 / Self-Quiz Questions 776 / Critical Thinking
Questions 777 / Answers to Figure Questions 777


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