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Gerard J. Tortora / Bryan Derrickson

14th Edition
Principles of

ANATOMY &
PHYSIOLOGY
14th Edition

Gerard J. Tortora
Bergen Community College

Bryan Derrickson
Valencia College
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10 9 8 7 6 5 4 3 2 1
 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 bachelors degree in biology from Fairleigh Dickinson University and his masters 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), the
Courtesy of Heidi Chung

American Association for the Advancement of Science (AAAS), the 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 MACUBs 1992 Presidents 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.
Jerry is the author of several best-selling science textbooks and laboratory manuals, a calling that often requires an additional
40hours 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.

Courtesy of Gerard J. Tortora

To Reverend Dr. James F. Tortora, my brother, my friend, and my role model.
His life of dedication has inspired me in so many ways, both personally and professionally,
and I honor him and pay tribute to him with this dedication. G.J.T.

Bryan Derrickson is Professor of Biology at Valencia College in Orlando, Florida, where he teaches
human anatomy and physiology as well as general biology and human sexuality. He received his bachelors
Courtesy of Bryan Derrickson

degree in biology from Morehouse College and his Ph.D. in cell biology from Duke University. Bryans
study at Duke was in the Physiology Division within the Department of Cell Biology, so while his degree
is in cell biology, his training focused on physiology. At Valencia, he frequently serves on faculty hiring
committees. He has served as a member of the Faculty Senate, which is the governing body of the college,
and as a member of the Faculty Academy Committee (now called the Teaching and Learning Academy),
which sets the standards for the acquisition of tenure by faculty members. Nationally, he is a member of
the Human Anatomy and Physiology Society (HAPS) and the National Association of Biology Teachers
(NABT). Bryan has always wanted to teach. Inspired by several biology professors while in college, he
decided to pursue physiology with an eye to teaching at the college level. He is completely dedicated to the success of his students.
He particularly enjoys the challenges of his diverse student population, in terms of their age, ethnicity, and academic ability, and finds
being able to reach all of them, despite their differences, a rewarding experience. His students continually recognize Bryans efforts and
care by nominating him for a campus award known as the Valencia Professor Who Makes Valencia a Better Place to Start. Bryan
has received this award three times.

To my family: Rosalind, Hurley, Cherie, and Robb.

Your support and motivation have been invaluable to me. B.H.D.

iii
 PREFACE

An anatomy and physiology course can be the gateway to a gratifying career in a host of health-related
professions. It can also be an incredible challenge. Principles of Anatomy and Physiology,14th
edition continues to offer a balanced presentation of content under the umbrella of our primary and
unifying theme of homeostasis, supported by relevant discussions of disruptions to homeostasis. Through
years of collaboration with students and instructors alike, this new edition of the textintegrated with
WileyPLUS with ORIONbrings together deep experience and modern innovation to provide solutions
for students greatest challenges.

We have designed the organization and flow of content within these pages to provide students with
an accurate, clearly written, and expertly illustrated presentation of the structure and function of
the human body. We are also cognizant of the fact that the teaching and learning environment has
changed significantly to rely more heavily on the ability to access the rich content in this printed text
in a variety of digital ways, anytime and anywhere. We are pleased that this 14th edition meets these
changing standards and offers dynamic and engaging choices to make this course more rewarding
and fruitful. Students can start here, and armed with the knowledge they gain through a professors
guidance using these materials, be ready to go anywhere with their careers.

New for This Edition

The 14th edition of Principles of Anatomy and Physiology has been updated throughout, paying
careful attention to include the most current medical terms in use (based on Terminologia Anatomica)
and including an enhanced glossary. The design has been refreshed to ensure that the content is
clearly presented and easy to access. Clinical Connections that help students understand the relevance
of anatomical structures and functions have been updated throughout and in some cases are now
placed alongside related illustrations to strengthen these connections for students.

The all-important illustrations that support this most visual of sciences have been scrutinized and
revised as needed throughout. Nearly every chapter of the text has a new or revised illustration or
photograph.

ANTERIOR ANTERIOR

PULMONARY
VALVE (closed) Right coronary PULMONARY
artery VALVE (open)
Left coronary
artery
AORTIC AORTIC
VALVE VALVE
BICUSPID (closed) (open)
VALVE BICUSPID
(open) VALVE
(closed)
TRICUSPID
TRICUSPID
VALVE
VALVE
(closed)
(open)

POSTERIOR POSTERIOR

Superior view with atria removed: pulmonary and aortic Superior view with atria removed: pulmonary and aortic
valves closed, bicuspid and tricuspid valves open valves open, bicuspid and tricuspid valves closed

iv
 Crista galli
Axodendritic
Perpendicular
plate Frontal
sinus Axoaxonic

Superior nasal Left

concha orbit

Superior nasal
meatus

Middle nasal Maxillary

meatus Dendrites
sinus

Middle nasal Vomer

concha

Inferior nasal Oral cavity Axon

Axosomatic
concha
Maxilla
Cell body
Inferior nasal
meatus
Frontal section through ethmoid bone in skull

Thyroid cartilage of larynx

Cricoid cartilage of larynx

RIGHT LATERAL LOBE OF THYROID GLAND
LEFT LATERAL LOBE OF THYROID GLAND
ISTHMUS OF THYROID GLAND

Trachea

Brain

Optic nerve
Right lung Ethmoidal cells Periorbital fat

Arch of aorta Superior nasal concha

Superior nasal meatus

Nasal septum: Middle nasal concha

Anterior view
Perpendicular
Middle nasal meatus
plate of ethmoid
Maxillary sinus

Vomer

Inferior nasal concha

Inferior nasal meatus
Hard palate

Tongue

Frontal section showing conchae and meatuses

SEM x8000 SEM x4000 SEM x2700

Extension
Hyperextension
Flexion
Flexion

Extension
Flexion
Flexion

Hyperextension
Extension
Extension

Hyperextension

Atlanto-occipital and cervical Shoulder joint Elbow joint Wrist joint

intervertebral joints

Lateral
flexion
Extension

Flexion
Extension
Hyperextension Flexion

Hip joint Knee joint

Intervertebral joints

v
 c21TheCardiovascularSystemBloodVesselsAndHemodynamics.indd Page 747 9/16/13 8:35 AM f-481

Enhancing our emphasis on the importance of homeostasis and the mechanisms that support it, we have re-
designed the illustrations describing feedback diagrams throughout the text. Introduced in the first chapter, the
distinctive design helps students recognize the key components of a feedback cycle, whether studying the control
manBody.indd Page 10 7/11/13 11:08 AM f-481 /204/WB00924/9781118345009/ch01/text_s
of blood pressure, regulation of breathing, regulation of glomerular filtration
Figure 21.14 Negative feedback regulation of blood
rate, or a host of other functions involving negative or positive feedback. To pressure via baroreceptor reflexes.
aid visual learners, color is used consistentlygreen for a controlled condition,
When blood pressure decreases, heart rate increases.
blue for receptors, purple for the control center, and red for effectors.

Figure 1.3 Homeostatic regulation of blood pressure by STIMULUS

a negative feedback system. The broken return arrow with a
negative sign surrounded by a circle symbolizes negative feedback.
Disrupts homeostasis
If the response reverses the stimulus, a system is by decreasing
operating by negative feedback.
CONTROLLED CONDITION
STIMULUS Blood pressure

Disrupts homeostasis
by increasing

CONTROLLED CONDITION RECEPTORS

Blood pressure Baroreceptors
in carotid sinus
and arch of aorta

RECEPTORS
Baroreceptors
in certain Input Stretch less, which decreases
blood vessels rate of nerve impulses

CONTROL CENTERS
Input Nerve impulses CV center in Adrenal
medulla oblongata medulla
Return to homeostasis
when increased
CONTROL CENTER cardiac output and
Brain increased vascular
resistance bring
Output blood pressure
Increased Increased secretion back to normal
Return to sympathetic, of epinephrine and
homeostasis when decreased para- norepinephrine
the response brings sympathetic from adrenal medulla
blood pressure stimulation
back to normal
Output Nerve impulses EFFECTORS

Heart
Blood
EFFECTORS vessels

Heart

Blood Increased stroke
vessels volume and heart rate
lead to increased
cardiac output (CO)
Constriction of blood
vessels increases
systemic vascular
resistance (SVR)

RESPONSE
Increased blood pressure

RESPONSE
A decrease in heart rate
and the dilation (widening) Does this negative feedback cycle represent the changes
of blood vessels cause that occur when you lie down or when you stand up?
blood pressure to decrease

What would happen to heart rate if some stimulus

caused blood pressure to decrease? Would this occur by
way of positive or negative feedback?

vi
In addition, following the
chapter or chapters covering
each body system, a page
is devoted to fostering
understanding of how each
system contributes to overall
homeostasis through its
interaction with other body
FOCUS on HOMEOSTASIS
LYMPHATIC SYSTEM
and IMMUNITY
INTEGUMENTARY
Glucocorticoids such as cortisol depress
SYSTEM inflammation and immune responses
Androgens stimulate growth of axillary Thymic hormones promote maturation of
and pubic hair and activation of
sebaceous glands
T cells (a type of white blood cell)

systems. These Focus on Excess melanocyte-stimulating hormone

Homeostasis pages have
been redesigned for a more
effective presentation of this
summary material.
(MSH) causes darkening of skin
SKELETAL
SYSTEM
Human growth hormone (hGH) and
insulinlike growth factors (IGFs) stimulate
bone growth
Estrogens cause closure of the epiphyseal
plates at the end of puberty and help
maintain bone mass in adults
Parathyroid hormone (PTH) and calcitonin
regulate levels of calcium and other
minerals in bone matrix and blood
Thyroid hormones are needed for normal
development and growth of the skeleton
MUSCULAR
SYSTEM
RESPIRATORY
SYSTEM
Epinephrine and norepinephrine dilate
(widen) airways during exercise and other
stresses
Erythropoietin regulates amount of
oxygen carried in blood by adjusting
number of red blood cells
DIGESTIVE
SYSTEM
Epinephrine and norepinephrine depress
activity of the digestive system
Gastrin, cholecystokinin, secretin, and
glucose-dependent insulinotropic
peptide (GIP) help regulate digestion
CONTRIBUTIONS OF Calcitriol promotes absorption of dietary
calcium

Epinephrine and norepinephrine help THE ENDOCRINE SYSTEM Leptin suppresses appetite

increase blood flow to exercising muscle

PTH maintains proper level of Ca2+,
needed for muscle contraction
Glucagon, insulin, and other hormones
regulate metabolism in muscle fibers
hGH, IGFs, and thyroid hormones help
maintain muscle mass
NERVOUS
SYSTEM
Several hormones, especially thyroid
hormones, insulin, and growth hormone,
influence growth and development of the
nervous system
PTH maintains proper level of Ca2+,
needed for generation and conduction of
nerve impulses
CARDIOVASCULAR
SYSTEM
Erythropoietin (EPO) promotes formation
of red blood cells
Aldosterone and antidiuretic hormone
(ADH) increase blood volume
Epinephrine and norepinephrine increase
heart rate and force of contraction
Several hormones elevate blood pressure
during exercise and other stresses
FOR ALL BODY SYSTEMS
Together with the nervous system, circulating
and local hormones of the endocrine system
regulate activity and growth of target cells
throughout the body
Several hormones regulate metabolism,
uptake of glucose, and molecules used for
ATP production by body cells
URINARY
SYSTEM
ADH, aldosterone, and atrial natriuretic
peptide (ANP) adjust the rate of loss of
water and ions in the urine, thereby
regulating blood volume and ion content
of the blood
REPRODUCTIVE
SYSTEMS
Hypothalamic releasing and inhibiting
hormones, follicle-stimulating hormone
(FSH), and luteinizing hormone (LH)
regulate development, growth, and
secretions of the gonads (ovaries and
testes)
Estrogens and testosterone contribute to
development of oocytes and sperm and
stimulate development of secondary sex
characteristics
Prolactin promotes milk secretion in
mammary glands
Oxytocin causes contraction of the uterus
and ejection of milk from the mammary
glands

We are most excited about the enhanced digital experience now available with the 14th edition of this
text. WileyPLUS now includes a powerful new adaptive learning component called ORION that allows
students to take charge of their study time in ways they have not previously experienced and prepares
them for more meaningful classroom and laboratory interactions. WileyPLUS itself has been refreshed
with a new design that allows easier discoverability and access to the rich resources including new 3-D
animations, Interactions, Muscles in Motion, Real Anatomy, Anatomy Drill and Practice, and PowerPhys.
New for the 14th edition is a digital alternative called All Access Pack for Principles of Anatomy
and Physiology,14th edition. This choice offers you a full e-text to download and keep, full access to
WileyPLUS, and a Study Resource Guide to use as a basis for taking notes in class and studying. It provides
you with everything you need for your course, anytime, anywhere, on any device.

vii
 BRIEF CONTENTS
1AN INTRODUCTION TO THE HUMAN BODY 1

2THE CHEMICAL LEVEL OF ORGANIZATION 27

3THE CELLULAR LEVEL OF ORGANIZATION 59

4THE TISSUE LEVEL OF ORGANIZATION 106

5THE INTEGUMENTARY SYSTEM 142

6THE SKELETAL SYSTEM: BONE TISSUE 169

7THE SKELETAL SYSTEM: THE AXIAL SKELETON 192

8THE SKELETAL SYSTEM: THE APPENDICULAR SKELETON 231

9JOINTS 258

10MUSCULAR TISSUE 291

11THE MUSCULAR SYSTEM 328

12NERVOUS TISSUE 399

13THE SPINAL CORD AND SPINAL NERVES 442

14THE BRAIN AND CRANIAL NERVES 473

15THE AUTONOMIC NERVOUS SYSTEM 523

16SENSORY, MOTOR, AND INTEGRATIVE SYSTEMS 546

17THE SPECIAL SENSES 572

18THE ENDOCRINE SYSTEM 615

19THE CARDIOVASCULAR SYSTEM: THE BLOOD 661

20THE CARDIOVASCULAR SYSTEM: THE HEART 688

21THE CARDIOVASCULAR SYSTEM: BLOOD VESSELS AND HEMODYNAMICS 729

22THE LYMPHATIC SYSTEM AND IMMUNITY 799

23THE RESPIRATORY SYSTEM 840

24THE DIGESTIVE SYSTEM 886

25METABOLISM AND NUTRITION 940

26THE URINARY SYSTEM 979

27FLUID, ELECTROLYTE, AND ACIDBASE HOMEOSTASIS 1023

28THE REPRODUCTIVE SYSTEMS 1041

29DEVELOPMENT AND INHERITANCE 1089

APPENDIX A:MEASUREMENTS A-1 APPENDIX B:PERIODIC TABLE B-3 APPENDIX C:NORMAL

VALUES FOR SELECTED BLOOD TESTS C-4 APPENDIX D:NORMAL VALUES FOR SELECTED URINE
TESTS D-6 APPENDIX E:ANSWERS E-8 GLOSSARY G-1 CREDITS C-1 INDEX I-1

xiv
 An Introduction
to the Human Body
The human body and homeostasis
Humans have many ways to maintain homeostasis, the state of relative stability of the bodys
internal environment. Disruptions to homeostasis often set in motion corrective cycles, called
feedback systems, that help restore the conditions needed for health and life.

Our fascinating journey through the human body begins with an overview of the meanings of anatomy and physiology,
followed by a discussion of the organization of the human body and the properties that it shares with all living things.
Next, you will discover how the body regulates its own internal environment; this unceasing process, called homeostasis,
is a major theme in every chapter of this book. Finally, we introduce the basic vocabulary that will help you speak about
the body in a way that is understood by scientists and health-care professionals alike.
ers
arch e
Res
oto h
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br
Li
to

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Ph
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nc
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Ferm
Mauro

Did you ever wonder why an autopsy is performed

1
2 CHAPTER 1 AN INTRODUCTION TO THE HUMAN BODY

1.1 Anatomy and Physiology
Defined
OBJECTIVE
Define anatomy and physiology, and name several
branches of these sciences.
Two branches of scienceanatomy and physiologyprovide the
foundation for understanding the bodys parts and functions.
Anatomy (a-NAT-o-me; ana- up; -tomy process of cutting) is
the science of body structures and the relationships among them.
It was first studied by dissection (dis-SEK-shun; dis- apart;
-section act of cutting), the careful cutting apart of body struc-
tures to study their relationships. Today, a variety of imaging
techniques (see Table 1.3) also contribute to the advancement of
anatomical knowledge. Whereas anatomy deals with structures of
the body, physiology (fiz-e-OL-o-je; physio- nature; -logy
study of) is the science of body functionshow the body parts work.
Table 1.1 describes several branches of anatomy and physiology.
Because structure and function are so closely related, you will
learn about the human body by studying its anatomy and physiol-
ogy together. The structure of a part of the body often reflects its
functions. For example, the bones of the skull join tightly to form
a rigid case that protects the brain. The bones of the fingers are
more loosely joined to allow a variety of movements. The walls of
the air sacs in the lungs are very thin, permitting rapid movement
of inhaled oxygen into the blood.
CHECKPOINT
1. What body function might a respiratory therapist strive
to improve? What structures are involved?
2. Give your own example of how the structure of a part of
the body is related to its function.
1.2 Levels of Structural
Organization and
Body Systems
OBJECTIVES
Describe the bodys six levels of structural organization.
List the 11 systems of the human body, representative
organs present in each, and their general functions.

TABLE
TABLE 1.1
1.3
Selected Branches of Anatomy and Physiology
BRANCH OF ANATOMY STUDY OF BRANCH OF PHYSIOLOGY STUDY OF

Embryology The first eight weeks of Neurophysiology Functional properties of nerve

(em-bre-OL-o-je; development after fertilization
embry- embryo; of a human egg.
-logy study of)
Developmental biology The complete development of an
individual from fertilization to
death.
Cell biology Cellular structure and functions.
Histology Microscopic structure of tissues.
(his-TOL-o -je; hist- tissue)
Gross anatomy Structures that can be examined
without a microscope.
Systemic anatomy Structure of specific systems of
the body such as the nervous or
respiratory systems.
Regional anatomy Specific regions of the body such
as the head or chest.
Surface anatomy Surface markings of the body
to understand internal anatomy
through visualization and
palpation (gentle touch).
Imaging anatomy Body structures that can be
visualized with techniques such
as x-rays, MRI, and CT scans.
Pathological anatomy Structural changes (gross to
(path-o-LOJ-i-kal; microscopic) associated with
path- disease) disease.
(NOOR-o-fiz-e-ol-o-je; cells.
neuro- nerve)
Endocrinology
(en-do-kri-NOL-o-je;
endo- within; -crin secretion)
Cardiovascular physiology
(kar-de-o-VAS-ku-lar;
cardi- heart;
vascular blood vessels)
Immunology
(im-u-NOL-o-je;
immun- not susceptible)
Respiratory physiology
(RES-pi-ra-tor-e;
respira- to breathe)
Renal physiology
(RE-nal; ren- kidney)
Exercise physiology
Pathophysiology
(Path-o-fiz-e-ol-o-je)
Hormones (chemical regulators in
the blood) and how they control
body functions.
Functions of the heart and blood
vessels.
The bodys defenses against
disease-causing agents.
Functions of the air passageways
and lungs.
Functions of the kidneys.
Changes in cell and organ functions
due to muscular activity.
Functional changes associated
with disease and aging.
 1.2 LEVELS OF STRUCTURAL ORGANIZATION AND BODY SYSTEMS 3
The levels of organization of a languageletters, words, sentences, of matter that participate in chemical reactions, and molecules,
paragraphs, and so oncan be compared to the levels of organiza- two or more atoms joined together. Certain atoms, such as
carbon (C), hydrogen (H), oxygen (O), nitrogen (N), phos-

1
tion of the human body. Your exploration of the human body will
extend from atoms and molecules to the whole person. From the phorus (P), calcium (Ca), and sulfur (S), are essential for

smallest to the largest, six levels of organization will help you to
understand anatomy and physiology: the chemical, cellular, tissue,
organ, system, and organismal levels of organization (Figure 1.1).
1 Chemical level. This very basic level can be compared to the
letters of the alphabet and includes atoms, the smallest units
C H A P T E R
maintaining life. Two familiar molecules found in the body
are deoxyribonucleic acid (DNA), the genetic material passed
from one generation to the next, and glucose, commonly
known as blood sugar. Chapters 2 and 25 focus on the chemi-
cal level of organization.

Figure 1.1 Levels of structural organization in the human body.

The levels of structural organization are chemical, cellular, tissue, organ, system, and organismal.

2 CELLULAR LEVEL

1 CHEMICAL LEVEL

3 TISSUE LEVEL

Smooth muscle cell

Atoms (C, H, O, N, P)
Smooth muscle tissue
Molecule (DNA)
5 SYSTEM LEVEL
Epithelial
and
connective
tissues
4 ORGAN LEVEL

Salivary glands
Mouth
Pharynx

Smooth muscle
Esophagus tissue layers

Epithelial tissue
Stomach

Stomach
Liver
Pancreas
Gallbladder (behind stomach)

Large intestine Small intestine

6 ORGANISMAL LEVEL
Digestive system

Which level of structural organization is composed of two or more different types of tissues that work together to perform a
specific function?
4 CHAPTER 1 AN INTRODUCTION TO THE HUMAN BODY
2 Cellular level. Molecules combine to form cells, the basic
structural and functional units of an organism that are com-
posed of chemicals. Just as words are the smallest elements
of language that make sense, cells are the smallest living
units in the human body. Among the many kinds of cells in
your body are muscle cells, nerve cells, and epithelial cells.
Figure 1.1 shows a smooth muscle cell, one of the three types
of muscle cells in the body. The cellular level of organization
is the focus of Chapter 3.
3 Tissue level. Tissues are groups of cells and the materials
surrounding them that work together to perform a particular
function, similar to the way words are put together to form
sentences. There are just four basic types of tissues in your
body: epithelial tissue, connective tissue, muscular tissue, and
nervous tissue. Epithelial tissue covers body surfaces, lines
hollow organs and cavities, and forms glands. Connective tis-
sue connects, supports, and protects body organs while distrib-
uting blood vessels to other tissues. Muscular tissue contracts
to make body parts move and generates heat. Nervous tissue
carries information from one part of the body to another through
nerve impulses. Chapter 4 describes the tissue level of organi-
zation in greater detail. Shown in Figure 1.1 is smooth muscle
tissue, which consists of tightly packed smooth muscle cells.
4 Organ level. At the organ level different types of tissues are
joined together. Similar to the relationship between sentences
and paragraphs, organs are structures that are composed of
two or more different types of tissues; they have specific func-
tions and usually have recognizable shapes. Examples of
organs are the stomach, skin, bones, heart, liver, lungs, and
TABLE 1.2
The Eleven Systems of the Human Body
INTEGUMENTARY SYSTEM (CHAPTER 5)
Components: Skin and associated Hair
structures, such as hair,
fingernails and toenails, sweat
glands, and oil glands.
Functions: Protects body; helps
regulate body temperature;
eliminates some wastes; helps
make vitamin D; detects Skin and
sensations such as touch, pain, associated
warmth, and cold; stores fat and glands
provides insulation.
Fingernails
T
Toenails
brain. Figure 1.1 shows how several tissues make up the stom-
ach. The stomachs outer covering is a layer of epithelial
tissue and connective tissue that reduces friction when the
stomach moves and rubs against other organs. Underneath are
three layers of a type of muscular tissue called smooth muscle
tissue, which contracts to churn and mix food and then push it
into the next digestive organ, the small intestine. The inner-
most lining is an epithelial tissue layer that produces fluid and
chemicals responsible for digestion in the stomach.
5 System level. A system (or chapter in our language analogy)
consists of related organs (paragraphs) with a common func-
tion. An example of the system level, also called the organ-
system level, is the digestive system, which breaks down and
absorbs food. Its organs include the mouth, salivary glands,
pharynx (throat), esophagus (food tube), stomach, small in-
testine, large intestine, liver, gallbladder, and pancreas.
Sometimes an organ is part of more than one system. The
pancreas, for example, is part of both the digestive system
and the hormone-producing endocrine system.
6 Organismal level. An organism (OR-ga-nizm), any living
individual, can be compared to a book in our analogy. All the
parts of the human body functioning together constitute the
total organism.
In the chapters that follow, you will study the anatomy and
physiology of the body systems. Table 1.2 lists the components
and introduces the functions of these systems. You will also dis-
cover that all body systems influence one another. As you study
each of the body systems in more detail, you will discover how
SKELETAL SYSTEM (CHAPTERS 69)
Components: Bones and joints
of the body and their associated
cartilages.
Functions: Supports and
protects body; provides surface
area for muscle attachments; Bone
aids body movements; houses Cartilage
cells that produce blood cells;
stores minerals and lipids (fats).
Joint
 1.3 CHARACTERISTICS OF THE LIVING HUMAN ORGANISM 5
they work together to maintain health, provide protection from CHECKPOINT
disease, and allow for reproduction of the human species. 3. Define the following terms: atom, molecule, cell, tissue,

1
organ, system, and organism.
4. At what levels of organization would an exercise

Noninvasive
C L INIC A L C ON N E C T ION |
Diagnostic Techniques
Health-care professionals and students of anatomy and physiol-
ogy commonly use several noninvasive diagnostic techniques to
assess certain aspects of body structure and function. A noninvasive
diagnostic technique is one that does not involve insertion of an
instrument or device through the skin or a body opening. In inspec-
tion, the examiner observes the body for any changes that deviate
from normal. For example, a physician may examine the mouth cavity
for evidence of disease. Following inspection, one or more additional
techniques may be employed. In palpation (pal-PA-shun; palp- gently
touching) the examiner feels body surfaces with the hands. An exam-
ple is palpating the abdomen to detect enlarged or tender internal
organs or abnormal masses. In auscultation (aws-kul-TA-shun; auscult-
listening) the examiner listens to body sounds to evaluate the func-
tioning of certain organs, often using a stethoscope to amplify the
sounds. An example is auscultation of the lungs during breathing to
check for crackling sounds associated with abnormal fluid accumula-
tion. In percussion (pur-KUSH-un; percus- beat through) the exam-
iner taps on the body surface with the fingertips and listens to the
resulting sound. Hollow cavities or spaces produce a different sound
than solid organs. For example, percussion may reveal the abnormal
presence of fluid in the lungs or air in the intestines. It may also pro-
vide information about the size, consistency, and position of an under-
lying structure. An understanding of anatomy is important for the
effective application of most of these diagnostic techniques.
C H A P T E R
physiologist study the human body? (Hint: Refer to
Table 1.1.)
5. Referring to Table 1.2, which body systems help eliminate
wastes?
1.3 Characteristics of the Living
Human Organism
OBJECTIVES
Define the important life processes of the human body.
Basic Life Processes
Certain processes distinguish organisms, or living things, from
nonliving things. Following are the six most important life pro-
cesses of the human body:
1. Metabolism (me-TAB-o-lizm) is the sum of all chemical
processes that occur in the body. One phase of metabolism is
catabolism (ka-TAB-o-lizm; catabol- throwing down; -ism
a condition), the breakdown of complex chemical substances
into simpler components. The other phase of metabolism is
anabolism (a-NAB-o-lizm; anabol- a raising up), the build-
ing up of complex chemical substances from smaller, simpler
components. For example, digestive processes catabolize (split)

MUSCULAR SYSTEM (CHAPTERS 10, 11) NERVOUS SYSTEM (CHAPTERS 1217)

Components: Specifically, skeletal Components: Brain, spinal cord, Brain

muscle tissuemuscle usually nerves, and special sense organs, such
attached to bones (other muscle tissues as eyes and ears.
include smooth and cardiac). Functions: Generates action potentials
Functions: Participates in body Skeletal (nerve impulses) to regulate body
muscle
movements, such as walking; activities; detects changes in bodys Spinal
maintains posture; produces heat. T
Tendon internal and external environments, cord
interprets changes, and responds by
causing muscular contractions or
glandular secretions.

Nerve

TA B L E 1. 2 CONTINUES
 Development and
Inheritance
Development, inheritance, and homeostasis
Both the genetic material inherited from parents (heredity) and normal development in the
uterus (environment) play important roles in determining the homeostasis of a developing
embryo and fetus and the subsequent birth of a healthy child.

Developmental biology is the study of the sequence of events from the fertilization of a secondary oocyte by a sperm cell to
the formation of an adult organism. Pregnancy is a sequence of events that begins with fertilization; proceeds to implantation,
embryonic development, and fetal development; and ideally ends with birth about 38 weeks later, or 40 weeks after the last
menstrual period.

Obstetrics (ob-STET-riks; obstetrix midwife) deals with the management of pregnancy, labor, and the neonatal period
-tal; pre- before; -natal birth) is the time from
-tal), the first 28 days after birth. Prenatal development (pre-NA
(ne-o-NA
ffertilization
ertilization tto and is divided into three periods of three calendar months each,
o birth an
called
calle trimesters.
ed trimeste

1. The ffirst trimester is the most critical stage of development, during

which the rudiments of all the major organ systems appear, and also
during which the developing organism is the most vulnerable to the
dur
effects of drugs, radiation, and microbes.
ef

22. The second trimester is characterized by the nearly complete

development of organ systems. By the end of this stage, the
fetus assumes distinctively human features.

33. The third trimester represents a period of rapid fetal growth.

During the early stages of this period, most of the organ
systems are becoming fully functional.

this chapter, we focus on the developmental sequence from

In thi
fertilization through implantation, embryonic and fetal development,
fertilizat
s
rbi
labor, birth, aand the principles of inheritance (the passage of hereditary traits
o
n/C
cea
O
from one generation to another).

Did you ever wonder why the heart, blood vessels, and blood
begin to form so early in the developmental process

1089
1090 CHAPTER 29 DEVELOPMENT AND INHERITANCE

lucida, called ZP3, acts as a sperm receptor. Its binding to specific

29.1 Embryonic Period
OBJECTIVE
Explain the major developmental events that occur during
the embryonic period.
First Week of Development
The embryonic period extends from fertilization through the
eighth week. The first week of development is characterized by
several significant events including fertilization, cleavage of the
zygote, blastocyst formation, and implantation.
Fertilization
During fertilization (fer-ti-li-ZA -shun; fertil- fruitful), the ge-
netic material from a haploid sperm cell (spermatozoon) and a
haploid secondary oocyte merges into a single diploid nucleus. Of
the 200 million sperm introduced into the vagina, fewer than
2million (1%) reach the cervix of the uterus and only about 200
reach the secondary oocyte. Fertilization normally occurs in the
uterine (fallopian) tube within 12 to 24 hours after ovulation.
Sperm can remain viable for about 48 hours after deposition in
the vagina, although a secondary oocyte is viable for only about
24 hours after ovulation. Thus, pregnancy is most likely to occur
if intercourse takes place during a 3-day windowfrom 2 days
before ovulation to 1 day after ovulation.
Sperm swim from the vagina into the cervical canal by the whip-
like movements of their tails (flagella). The passage of sperm
through the rest of the uterus and then into the uterine tube results
mainly from contractions of the walls of these organs. Prostaglan-
dins in semen are believed to stimulate uterine motility at the time
of intercourse and to aid in the movement of sperm through the
uterus and into the uterine tube. Sperm that reach the vicinity of the
oocyte within minutes after ejaculation are not capable of fertiliz-
ing it until about seven hours later. During this time in the female
reproductive tract, mostly in the uterine tube, sperm undergo
capacitation (ka-pas-i-TA -shun; capacit- capable of), a series of
functional changes that cause the sperms tail to beat even more
membrane proteins in the sperm head triggers the acrosomal re-
action, the release of the contents of the acrosome. The acroso-
mal enzymes digest a path through the zona pellucida as the lash-
ing sperm tail pushes the sperm cell onward. Although many
sperm bind to ZP3 molecules and undergo acrosomal reactions,
only the first sperm cell to penetrate the entire zona pellucida and
reach the oocytes plasma membrane fuses with the oocyte.
The fusion of a sperm cell with a secondary oocyte sets in motion
events that block polyspermy (POL-e-sper-me), fertilization by
more than one sperm cell. Within a few seconds, the cell membrane
of the oocyte depolarizes, which acts as a fast block to polyspermy
the inability of a depolarized oocyte to fuse with another sperm.
Depolarization also triggers the intracellular release of calcium ions,
which stimulate exocytosis of secretory vesicles from the oocyte.
Figure 29.1 Selected structures and events in fertilization.
During fertilization, genetic material from a sperm cell
and a secondary oocyte merge to form a single diploid
nucleus.
Sperm cell
PATH OF SPERM
CELL:
Corona radiata
Zona pellucida
Plasma membrane
of secondary
oocyte
First polar Cytoplasm of
body secondary oocyte

vigorously and prepare its plasma membrane to fuse with the

oocytes plasma membrane. During capacitation, sperm are acted (a) Sperm cell penetrating secondary oocyte
on by secretions in the female reproductive tract that result in the
removal of cholesterol, glycoproteins, and proteins from the plasma Head of Secondary Pronuclei
membrane around the head of the sperm cell. Only capacitated sperm cell oocyte
sperm are capable of being attracted by and responding to chemical
factors produced by the surrounding cells of the ovulated oocyte.
For fertilization to occur, a sperm cell first must penetrate two
layers: the corona radiata (ko-RO-na crown; ra-de-A-ta to
shine), the granulosa cells that surround the secondary oocyte,
and the zona pellucida (ZO-na zone; pe-LOO-si-da allow-
ing passage of light), the clear glycoprotein layer between the
corona radiata and the oocytes plasma membrane (Figure 29.1a).
The acrosome (AK-ro-som), a helmetlike structure that covers SEM 1100x LM 250x
the head of a sperm (see Figure 28.6), contains several enzymes. (b) Sperm cell in contact with (c) Male and female pronuclei
Acrosomal enzymes and strong tail movements by the sperm help secondary oocyte
it penetrate the cells of the corona radiata and come in contact
with the zona pellucida. One of the glycoproteins in the zona pel- What is capacitation?

The molecules released by exocytosis inactivate ZP3 and harden the
entire zona pellucida, events called the slow block to polyspermy.
Once a sperm cell enters a secondary oocyte, the oocyte first
must complete meiosis II. It divides into a larger ovum (mature
egg) and a smaller second polar body that fragments and disinte-
grates (see Figure 28.15). The nucleus in the head of the sperm
develops into the male pronucleus, and the nucleus of the fertil-
ized ovum develops into the female pronucleus (Figure 29.1c).
After the male and female pronuclei form, they fuse, producing a
single diploid nucleus, a process known as syngamy (SIN-ga-
me). Thus, the fusion of the haploid (n) pronuclei restores the
diploid number (2n) of 46 chromosomes. The fertilized ovum
now is called a zygote (zygon yolk).
Dizygotic (fraternal) twins are produced from the independent
release of two secondary oocytes and the subsequent fertilization of
each by different sperm. They are the same age and in the uterus at
the same time, but genetically they are as dissimilar as any other
siblings. Dizygotic twins may or may not be the same sex. Because
monozygotic (identical) twins develop from a single fertilized
ovum, they contain exactly the same genetic material and are al-
ways the same sex. Monozygotic twins arise from separation of the
29.1 EMBRYONIC PERIOD 1091
During the formation of the blastocyst two distinct cell popula-
tions arise: the embryoblast and trophoblast (Figure 29.2e). The
embryoblast (EM-bre-o-blast), or inner cell mass, is located in-
ternally and eventually develops into the embryo. The trophoblast
(TRO F-o-blast; tropho- develop or nourish) is the outer superfi-
cial layer of cells that forms the spherelike wall of the blastocyst. It
will ultimately develop into the outer chorionic sac that surrounds
the fetus and the fetal portion of the placenta, the site of exchange
of nutrients and wastes between the mother and fetus. Around the
fifth day after fertilization, the blastocyst hatches from the zona
pellucida by digesting a hole in it with an enzyme, and then
squeezing through the hole. This shedding of the zona pellucida is
Figure 29.2 Cleavage and the formation of the
morula and blastocyst.
Cleavage refers to the early, rapid mitotic divisions of a
zygote.
Polar bodies
Blastomeres

developing cells into two embryos, which in 99% of the cases occurs
Zona pellucida
before 8 days have passed. Separations that occur later than 8days (a) CLEAVAGE OF
ZYGOTE, TWO-CELL
are likely to produce conjoined twins, a situation in which the STAGE (day 1)
twins are joined together and share some body structures.
Cleavage of the Zygote
After fertilization, rapid mitotic cell divisions of the zygote called Nucleus
cleavage (KLEV-ij) take place (Figure 29.2). The first division of Cytoplasm
the zygote begins about 24 hours after fertilization and is completed (b) CLEAVAGE OF
about 6 hours later. Each succeeding division takes slightly less ZYGOTE, FOUR-CELL
time. By the second day after fertilization, the second cleavage is STAGE (day 2)
completed and there are four cells (Figure 29.2b). By the end of the
third day, there are 16 cells. The progressively smaller cells pro-
duced by cleavage are called blastomeres (BLAS-to-merz; blasto-
germ or sprout; -meres parts). Successive cleavages eventu-
ally produce a solid sphere of cells called the morula (MOR-u-la
mulberry). The morula is still surrounded by the zona pellucida and (c) MORULA
is about the same size as the original zygote (Figure 29.2c). (day 4)

Blastocyst Formation
By the end of the fourth day, the number of cells in the morula in-
creases as it continues to move through the uterine tube toward the
uterine cavity. When the morula enters the uterine cavity on day 4
(d) BLASTOCYST,
or 5, a glycogen-rich secretion from the glands of the endometrium EXTERNAL VIEW
of the uterus passes into the uterine cavity and enters the morula (day 5)
through the zona pellucida. This fluid, called uterine milk, along

29
with nutrients stored in the cytoplasm of the blastomeres of the
morula, provides nourishment for the developing morula. At the C H A P T E R
32-cell stage, the fluid enters the morula, collects between the blas- Embryoblast
tomeres, and reorganizes them around a large fluid-filled cavity (inner cell mass)
(e) BLASTOCYST
called the blastocyst cavity (BLAS-to-sist; blasto- germ or (sectioned), Blastocyst cavity
sprout; -cyst bag), also called the blastocoel (BLAS-to-sel) (Fig- INTERNAL VIEW
Trophoblast
(day 5)
ure 29.2e). Once the blastocyst cavity is formed, the developing
mass is called the blastocyst. Though it now has hundreds of cells, What is the histological difference between a morula
the blastocyst is still about the same size as the original zygote. and a blastocyst?
1092 CHAPTER 29 DEVELOPMENT AND INHERITANCE

necessary in order to permit the next step, implantation (attach- (Figure 29.4). The decidua basalis is the portion of the endome-
ment) into the vascular, glandular endometrial lining of the uterus. trium between the embryo and the stratum basale of the uterus; it
provides large amounts of glycogen and lipids for the developing
Implantation embryo and fetus and later becomes the maternal part of the pla-
The blastocyst remains free within the uterine cavity for about centa. The decidua capsularis is the portion of the endometrium
2days before it attaches to the uterine wall. At this time the endo-
metrium is in its secretory phase. About 6 days after fertilization, Figure 29.3 Relationship of a blastocyst to the
the blastocyst loosely attaches to the endometrium in a process endometrium of the uterus at the time of implantation.
called implantation (im-plan-TA -shun) (Figure 29.3). As the
blastocyst implants, usually in either the posterior portion of the Implantation, the attachment of a blastocyst to the
fundus or the body of the uterus, it orients with the inner cell mass endometrium, occurs about 6 days after fertilization.
toward the endometrium (Figure 29.3b). About 7 days after fertil- BLASTOCYST
ization, the blastocyst attaches to the endometrium more firmly,
endometrial glands in the vicinity enlarge, and the endometrium
becomes more vascularized (forms new blood vessels). The blas- Opening of
tocyst eventually secretes enzymes and burrows into the endome- endometrial
trium, and becomes surrounded by it. gland
Following implantation, the endometrium is known as the de-
cidua (de-SID-u-a falling off). The decidua separates from the
endometrium after the fetus is delivered, much as it does in normal
menstruation. Different regions of the decidua are named based on ENDOMETRIUM
their positions relative to the site of the implanted blastocyst OF UTERUS
Uterine
cavity
Stem Cell Research and
C L I NI C AL C ON N E C T ION |
Therapeutic Cloning
(a) External view of blastocyst, about 6 days after fertilization
Stem cells are unspecialized cells that have the ability to divide
for indefinite periods and give rise to specialized cells. In the
context of human development, a zygote (fertilized ovum) is a stem
cell. Because it has the potential to form an entire organism, a zygote
is known as a totipotent stem cell (to-TIP-o-tent; totus- whole; Frontal
plane
-potentia power). Inner cell mass cells, called pluripotent stem cells
(ploo-RIP-o-tent; plur- several), can give rise to many (but not all) dif- Uterine
cavity
ferent types of cells. Later, pluripotent stem cells can undergo further
specialization into multipotent stem cells (mul-TIP-o-tent), stem cells
with a specific function. Examples include keratinocytes that produce Frontal section
new skin cells, myeloid and lymphoid stem cells that develop into blood through uterus ENDOMETRIUM OF UTERUS
cells, and spermatogonia that give rise to sperm. Pluripotent stem cells
currently used in research are derived from (1)the embryoblast of em- Endometrial gland
bryos in the blastocyst stage that were destined to be used for infertility
Opening of
treatments but were not needed and from (2)nonliving fetuses termi- endometrial gland
nated during the first 3 months of pregnancy.
Scientists are also investigating the potential clinical applications of
adult stem cellsstem cells that remain in the body throughout adult- TROPHOBLAST Blood
hood. Recent experiments suggest that the ovaries of adult mice con- vessels
tain stem cells that can develop into new ova (eggs). If these same
types of stem cells are found in the ovaries of adult women, scientists EMBRYOBLAST
could potentially harvest some of them from a woman about to un- (INNER CELL
MASS)
dergo a sterilizing medical treatment (such as chemotherapy), store
them, and then return the stem cells to her ovaries after the medical BLASTOCYST
treatment is completed in order to restore fertility. Studies have also CAVITY
suggested that stem cells in human adult red bone marrow have the
ability to differentiate into cells of the liver, kidney, heart, lung, skel-
etal muscle, skin, and organs of the gastrointestinal tract. In theory,
(b) Frontal section through endometrium of uterus and
adult stem cells from red bone marrow could be harvested from a blastocyst, about 6 days after fertilization
patient and then used to repair other tissues and organs in that pa-
tients body without having to use stem cells from embryos. How does the blastocyst merge with and burrow into
the endometrium?
 29.1 EMBRYONIC PERIOD 1093
located between the embryo and the uterine cavity. The decidua Figure 29.4 Regions of the decidua.
parietalis (par-ri-e-TAL-is) is the remaining modified endome-
trium that lines the noninvolved areas of the rest of the uterus. As The decidua is a modified portion of the endometrium
the embryo and later the fetus enlarges, the decidua capsularis that develops after implantation.
bulges into the uterine cavity and fuses with the decidua parietalis, Implanted DECIDUA BASALIS
thereby obliterating the uterine cavity. By about 27 weeks, the embryo
decidua capsularis degenerates and disappears.
The major events associated with the first week of develop-
ment are summarized in Figure 29.5.

CLINICAL CONNECTION | Ectopic Pregnancy

Ectopic pregnancy (ek-TOP-ik; ec- out of; -topic place) is

the development of an embryo or fetus outside the uterine cav- DECIDUA
ity. An ectopic pregnancy usually occurs when movement of the CAPSULARIS
fertilized ovum through the uterine tube is impaired by scarring due to DECIDUA
a prior tubal infection, decreased movement of the uterine tube PARIETALIS
smooth muscle, or abnormal tubal anatomy. Although the most com-
mon site of ectopic pregnancy is the uterine tube, ectopic pregnancies
may also occur in the ovary, abdominal cavity, or uterine cervix. Women Frontal section of uterus Details of decidua
who smoke are twice as likely to have an ectopic pregnancy because
nicotine in cigarette smoke paralyzes the cilia in the lining of the uter- Which part of the decidua helps form the maternal part
ine tube (as it does those in the respiratory airways). Scars from pelvic of the placenta?
inflammatory disease, previous uterine tube surgery, and previous ec-
topic pregnancy may also hinder movement of the fertilized ovum. CHECKPOINT
The signs and symptoms of ectopic pregnancy include one or two 1. Where does fertilization normally occur?
missed menstrual cycles followed by bleeding and acute abdominal and
2. How is polyspermy prevented?
pelvic pain. Unless removed, the developing embryo can rupture the
3. What is a morula, and how is it formed?
uterine tube, often resulting in death of the mother. Treatment options
4. Describe the layers of a blastocyst and their eventual
include surgery or the use of a cancer drug called methotrexate, which
fates.
causes embryonic cells to stop dividing and eventually disappear.
5. When, where, and how does implantation occur?

Figure 29.5 Summary of events associated with the first week of development.
Fertilization usually occurs in the uterine tube.

2. CLEAVAGE
(first cleavage
completed about 30
hours after fertilization)
3. MORULA
(34 days after fertilization)
1. FERTILIZATION
(occurs within
uterine tube
1224 hours
after ovulation) 4. BLASTOCYST
Frontal plane
(41/2 5 days after
fertilization)
Uterine cavity
5. IMPLANTATION
Ovulation (occurs about

29
6 days after
Ovary
fertilization)
Uterus: C H A P T E R
Endometrium
Myometrium

Frontal section through uterus, uterine tube, and ovary

In which phase of the uterine cycle does implantation occur?

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