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Acknowledgments
Human Physiology, 1st edition and WileyPLUS Learning Space would Space with ORION. Their thoughtful execution of resources and as-
not be possible without the help of many, particularly the numerous sessment has raised the bar for excellence. Thank you, so very much.
academic colleagues that collaborated with us along the way. First to Brian Antonsen Marshall University
thank are three contributors whose work informed and enhanced our Heidi Bustamante University of Colorado, Boulder
focus on critical thinking and connections to real life situations and
Patrick Cafferty Emory University
activities:
Janet Casagrand University of Colorado, Boulder
David Mallory, Marshall University, was instrumental in the design
and execution of the Critical Thinking Boxes and step-by-step ques- Lynn Diener Mount Mary University
tions found in each chapter. His choice of topics for these boxes John Erickson Ivy Tech State College-SE
adds great interest and really highlights the connections to the Gregory Loftin Metropolitan Community College
concepts at hand. Thank you so much, David. of Kansas City
Lynn Diener, Mount Mary University, researched and chose most Geoff Goellner Minnesota State University
of the primary research papers used in the Research to Reality Michael Griffin Angelo State University
activities. She used her expertise and experience in implementing Jill Tall Youngstown State University
this kind of investigation in her own classes to bring this unique
Tracy Wagner Washburn University of Topeka
feature to this project. I am very grateful, Lynn.
Paul Wagner Washburn University of Topeka
Heidi Bustamante, University of Colorado, Boulder, excels at writ-
ing thoughtful questions that challenge students to critically think Chad Wayne University of Houston
about what they have learned. I am so happy that she chose to en- The development of a first edition text and media for a course as
hance this text by writing the end-of-chapter Ponder This questions. complex as human physiology is a long process and would not be pos-
I really appreciate your work, Heidi. sible without the continued involvement of those “in the trenches”
I am also very grateful that Wiley commissioned a board of aca- teaching the course, who guided and informed our choices all along
demic advisors with expertise in teaching physiology to help inform the way. I am very grateful to my colleagues who have reviewed the
not only individual resources integrated with this text, but also as a drafts of manuscript, participated in focus groups and workshops, or
sounding board for all of us as we progressed through development. offered suggestions for improvement.
Ateegh Al-Arabi Johnson County Community College
WILEY ADVISORY BOARD FOR ANATOMY Brenda Alston-Mills Michigan State University
AND PHYSIOLOGY Gwen Bachman University of Nebraska, Lincoln
Heidi Bustamante University of Colorado, Boulder Ari Berowitz University of Oklahoma, Norman
Patrick Cafferty Emory University Eric Blough Marshall University
Janet Casagrand University of Colorado, Boulder Sunny K. Boyd University of Notre Dame
Lynn Diener Mount Mary University Carol Britson University of Mississippi
John Erickson Ivy Tech State College-SE Jackie Brittingham Simpson College
Melaney Farr Salt Lake Community College Brent Bruot Kent State University
Geoff Goellner Minnesota State University Heidi Bustamante University of Colorado, Boulder
Michael Griffin Angelo State University Sherell Byrd Fort Lewis College
Wanda Hargroder Louisiana State University Martin Burg Grand Valley State University
DJ Hennager Kirkwood Community College Phyllis Callahan Miami University
Sandra Hutchinson Santa Monica College Jackie Carnegie University of Ottawa
Heather Labbe University of Montana - Missoula Robert Carroll East Carolina University
Tom Lancraft St. Petersburg College Janet Casagrand University of Colorado, Boulder
David Mallory Marshall University Daniel Castellanos Florida International University
Russell Nolan Baton Rouge Community College Chris Dewitt University of South Carolina Aiken
Terry Thompson Wor-Wic Community College Debora Christensen Drake University
Tracy Wagner Washburn University of Topeka Ruth Clark Washington University, St. Louis
Paul Wagner Washburn University of Topeka Josefa Cubina New York Institute of Technology
I thank the following group of academics who have contributed Maria de Bellard California State University, Northridge
to the creation and integration of this text with WileyPLUS Learning Michael Deschenes College of William and Mary

vi • Acknowledgments
Lynn Diener Mount Mary University David Nutting University of Tennessee, Memphis
Stephen Dodd University of Florida Linda Ogren University of California - Santa Cruz
John Erickson Ivy Tech Ok-Kyong Park-Sarge University of Kentucky
Max G. Ervin Middle Tennessee State University William Percy University of South Dakota
Carol Fassbinder-Orth Creighton University David Pistole Indiana University of Pennsylvania
Ralph Ferges Palomar College Robert Preston Illinois State University
James Ferraro Southern Illinois School of Medicine David Petzel Creighton University
Michael Finkler Indiana University - Kokomo Steve Price Virginia Commonwealth University
John Fishback Ozarks Tech Community College Peter Reiser Ohio State University
David Flory University of Central Florida Nick A. Ritucci Wright State University
Victor Fomin University of Delaware Laurel Roberts University of Pittsburgh
Michelle French University of Toronto Sonia Rocah-Sanchez Creighton University
Geoff Goellner Minnesota State University Dean Scherer Oklahoma State University
Barbara Goodman University of South Dakota David Schulz University of Missouri
Mike Griffin Angelo State University Virginia Shea University of North Carolina, Chapel Hill
Bryan Hamilton Waynesburg University Allison Shearer Grossmont College
Janet Haynes Long Island University Rachel Smetanka Southern Utah University
Susan Heaphy The Ohio State University at Lima James Strauss Pennsylvania State University
Steven Heidemann Michigan State University Erik Swenson University of Washington
Stephen Henderson California State University, Chico Jill Tall Youngstown State University
William Higgins University of Maryland Bonnie Tarricone Ivy Tech Community College at Indianapolis
Steven Hobbs University of Colorado Mark Thomas University of Northern Colorado
James Hoffman Diablo Valley College Maureen Tubbiola St. Cloud State University
Kelly Johnson University of Kansas Paul Wagner Washburn University of Topeka
Steven King Oregon Health & Science University Tracy Wagner Washburn University of Topeka
Jennifer Kneafsey Tulsa Community College - Curt Walker Dixie State College of Utah
Northeast Campus Richard Walker University of Calgary
Megan Knoch Indiana University of Pennsylvania R. Douglass Watson University of Alabama, Birmingham
Joan Lafuze Indiana University East Chad Wayne University of Houston
Dean Lauritzen City College of San Francisco
Last, but certainly not least, I owe tremendous gratitude to the
John Lepri University of North Carolina, Greensboro team at Wiley. This team of collaborative publishing professionals is
Paul Lonquich California State University, Northridge dedicated, enthusiastic, and talented and each brings a level of skill to
Jennifer Lundmark California State University - Sacramento the job that, I believe, is unparalleled. My hat is off to the entire team:
David Mallory Marshall University Bonnie Roesch, Executive Editor; Karen Trost, Developmental Editor;
Theresa Martin San Mateo Community College Brittany Cheetham, Associate Editor; Maria Guarascio, Senior Market-
ing Manager; Linda Muriello, Senior Product Designer; Lindsey Myers,
Michael Masson Santa Barbara Community College
Market Solutions Assistant; Trish McFadden, Senior Production
Tamara Mau University of California, Berkeley Editor; Mary Ann Price, Senior Photo Editor; Claudia Volano and Anna
Eric McElroy College of Charleston Melhorn, Illustration Coordinators; Thomas Nery, Senior Designer. I
Thomas McNeilis Dixie State College of Utah also want to acknowledge Kevin Witt, Director, for his support of this
John McReynolds University of Michigan project. My heartfelt thanks to you all!
Jamie Melling Western University Bryan Derrickson
Jeanne Mitchell Truman State University Department of Science, PO Box 3028
Diane Morel University of the Sciences Valencia College
Tim Mullican Dakota Wesleyan University Orlando, FL 32802
[email protected]
Cheryl Neudauer Minneapolis Community
and Technical College
Acknowledgments • vii
 Brief Contents
1 An Introduction to Physiology 1

2 Chemical Composition of the Body 19

3 Cells 56

4 Metabolism 95

5 Transport Across the Plasma Membrane 128

6 Cell Signaling 156

7 The Nervous System and Neuronal Excitability 187

8 The Central Nervous System 238

9 Sensory Systems 275

10 Autonomic and Somatic Nervous Systems 341

11 Muscle 364

12 Control of Body Movement 411

13 The Endocrine System 432

14 The Cardiovascular System: The Heart 484

15 The Cardiovascular System: Blood Vessels and Hemodynamics 524

16 The Cardiovascular System: The Blood 565

17 The Immune System 593

18 The Respiratory System 625

19 The Urinary System 664

20 Fluid, Electrolyte, and Acid–Base Homeostasis 702

21 The Digestive System 721

22 Metabolic Adaptation, Energy Balance, and Temperature

Regulation 764

23 The Reproductive Systems 781

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
Glossary G-1 Index I-1

viii • Brief Contents

Contents
1 An Introduction to Physiology 1 Van der Waals Interactions Involve Transient
Fluctuations in Electron Distribution 27
1.1 Physiology Defined 2
2.3 Chemical Reactions 28
1.2 Levels of Organization in the Body 2
2.4 Inorganic Compounds and Solutions 28
1.3 Life Processes 5
Water Is Vital to Life 29
1.4 Homeostasis 5
Solutions, Colloids, and Suspensions Are Types of
Maintenance of Body Fluid Volume and Composition is Mixtures 30
Essential to Homeostasis 6
Inorganic Acids, Bases, and Salts Dissociate When
Homeostasis Is Regulated via Feedback Systems and Dissolved in Water 31
Feedforward Control 6
The Body’s Chemical Reactions Are Sensitive to
Homeostatic Imbalances Can Lead to Disorders, Changes in pH 31
Diseases, or Even Death 11
Buffer Systems Convert Strong Acids or
1.5 Physiology as a Science 11 Bases into Weak Acids or Bases 31
The History of Physiology Spans Thousands of Years 11 2.5 Organic Compounds 33
The Scientific Method Is a Systematic Way of Acquiring Carbon and Its Functional Groups Provide Useful
Knowledge About the Natural World 12 Properties to Living Organisms 33
Scientific Literature Helps Physiologists Conduct Carbohydrates Are Important Sources of Chemical
Research 13 Energy 34
Physiologists Use the Mechanistic Approach to Explain Lipids Contribute to Energy Storage, Membrane
How the Body Functions 13 Structure, and Hormone Production 35
Concept Mapping Allows Physiologists to Illustrate Proteins Are Chains of Amino Acids That Have
the Relationships Between Ideas 14 Diverse Roles 41
1.6 Key Themes of Physiology 14 Nucleic Acids Store and Express Genetic
Critical Thinking 10 / From Research to Reality 15 Information 48
ATP Serves as the Energy Currency of Living
Systems 49
Critical Thinking 40 / From Research to Reality 51
2 Chemical Composition of
the Body 19
2.1 How Matter Is Organized 20 3 Cells 56
All Forms of Matter Are Composed of Atoms 20
3.1 Components of a Cell 57
An Atom Contains Protons, Neutrons, and
Electrons 20 3.2 The Plasma Membrane 58
Each Atom Has an Atomic Number and a The Plasma Membrane Consists of a Lipid Bilayer
Mass Number 21 and a Variety of Proteins 58
Atomic Mass Is Measured In Daltons 22 Membrane Proteins Have Many Functions 59
Atoms Can Give Rise to Ions, Molecules, and Membrane Fluidity Allows Membrane Components to
Compounds 23 Interact and Move Around 60

2.2 Chemical Bonds 23 3.3 Cytoplasm 60

Ionic Bonds Involve the Loss or Gain of Electrons 24 The Cytosol Is the Site of Many Chemical Reactions 60
Covalent Bonds Involve Sharing of Electrons 25 Organelles Function in Cellular Growth, Maintenance,
and Reproduction 60
Hydrogen Bonds Result from Attraction of Oppositely
Charged Regions of Molecules 26 3.4 Nucleus 68
Contents • ix
 3.5 Gene Expression 70 NAD+ and FAD Help Generate ATP by Carrying
Transcription Conveys Genetic Information Hydrogen Atoms to the Electron Transport Chain 105
from DNA to RNA 71 Cellular Respiration Produces ATP by Breaking Down a
Translation Uses Genetic Information Carried Nutrient Molecule in the Presence of Oxygen 106
by mRNA to Synthesize a Protein 72 4.5 Carbohydrate Metabolism 109
3.6 Cell Division 74 Glucose Catabolism Can Generate 30 or 32 ATP 109
Somatic Cell Division Produces Two Identical Cells 74 Glucose Anabolism Includes Glycogenesis
Cell Destiny Is Controlled by Many Factors 76 and Gluconeogenesis 115

3.7 Cellular Diversity 78 4.6 Lipid Metabolism 117

Lipid Catabolism Involves Lipolysis and Beta
3.8 Organization of Cells into Tissues 79
Oxidation 117
Epithelial Tissue Serves as a Barrier, Secretes
Lipid Anabolism Occurs via Lipogenesis 118
Substances, and Absorbs Materials 79
Connective Tissue Supports, Insulates, and 4.7 Protein Metabolism 118
Protects the Organs of the Body 81 Protein Catabolism Leads to Deamination and
Muscle Tissue Specializes in Contraction 85 Keto Acid Breakdown 118
Nervous Tissue Detects and Responds to Changes in Protein Anabolism Results in the Synthesis of
the Environment 85 New Proteins 118
Cell Junctions Connect Adjacent Cells 86 4.8 Nutrition and Metabolism 120
Critical Thinking 88 / From Research to Reality 89 Many Minerals Have Known Functions in the Body 121
Vitamins Help Maintain Growth and Normal
Metabolism 121
4 Metabolism 95 Critical Thinking 107 / From Research to Reality 123

4.1 An Overview of Metabolism 96

4.2 Energy and Metabolism 96
5 Transport Across the Plasma
Energy Exists in Different Forms 96
Membrane 128
Chemical Reactions Release or Absorb Energy 97
5.1 Selective Permeability of the Plasma
Activation Energy Is Needed to Start a Chemical
Reaction 97
Membrane 129
Catalysts Lower the Activation Energy of Chemical 5.2 Gradients Across the Plasma
Reactions 98 Membrane 130
4.3 Enzymes 98 5.3 Classification of Membrane Transport
Enzymes Have Important Properties 99 Processes as Passive or Active 131
Enzymes Catalyze Reactions by Helping 5.4 Passive Transport 132
Molecules Interact 99 Diffusion Relies on the Kinetic Energy That
Many Enzymes Require a Cofactor in Order to Is Intrinsic to All Particles 132
Function 100 Simple Diffusion Occurs When a Solute Moves down Its
Various Factors Influence the Rate of Gradient Without Any Help 133
an Enzyme-Catalyzed Reaction 100 Facilitated Diffusion Uses a Protein to Move a Solute
A Sequence of Enzymatic Reactions Constitutes a down Its Gradient 134
Metabolic Pathway 101 Osmosis Results in Net Movement of Water Across a
Metabolic Pathways Are Shut down by Feedback Selectively Permeable Membrane 137
Inhibition 102
5.5 Active Transport 140
4.4 Role of ATP in Metabolism 103 Primary Active Transport Uses Energy from ATP to Move
Catabolism and Anabolism Are Coupled by ATP 103 a Solute Against Its Gradient 140
ATP Is Generated via Substrate-Level Phosphorylation Secondary Active Transport Uses Energy from an Ionic
and Oxidative Phosphorylation 103 Gradient to Move a Solute Against Its Gradient 141

x • Contents
 5.6 Vesicular Transport 145 7 The Nervous System and Neuronal
Endocytosis Allows Ligands, Large Solid Particles, and Excitability 187
Droplets of Extracellular Fluid to Enter Cells 145 7.1 Overview of the Nervous System 188
Exocytosis Is Used to Release Digestive Enzymes, The Nervous System Is Organized into the Central
Hormones, and Neurotransmitters from Certain Nervous System and the Peripheral Nervous
Cells 147 System 188
5.7 Transepithelial Transport 148 The Nervous System Performs Sensory, Integrative, and
Critical Thinking 144 / From Research to Reality 151 Motor Functions 188
7.2 Cells of the Nervous System 190
Neurons Are Responsible for the Main Functions of the
6 Cell Signaling 156 Nervous System 190
6.1 Methods of Cell-to-Cell Neuroglia Provide Physical, Nutritional, and Metabolic
Communication 157 Support to Neurons 193
Gap Junctions Electrically Couple Cells Together 157 Myelination Increases the Speed of Action Potential
Conduction 194
Cell-to-Cell Binding Is Important for Development and
Defense 157 Damaged Neurons Have a Limited Ability
to Repair Themselves 195
Communication Through Extracellular Chemical
Messengers Permits a Wide Variety of Responses 158 7.3 Electrical Signals in Neurons 196
6.2 Extracellular Chemical Messengers 158 Ion Channels Permit Certain Ions to Move Across the
Different Types of Extracellular Chemical Messengers Plasma Membrane 198
Exist 158 Resting Membrane Potential Is the Voltage That Exists
Extracellular Messengers Are Chemically Classified as Across the Plasma Membrane in an Excitable Cell at
Water-Soluble or Lipid-Soluble 161 Rest 198

Extracellular Messengers Travel Through Interstitial Graded Potentials Can Be Depolarizing or

Fluid and/or Blood to Reach Their Target Cells 161 Hyperpolarizing 207
Action Potentials Are Generated When the Axon
6.3 Receptors 162 Membrane Reaches Threshold 210
Messenger–Receptor Binding Exhibits Action Potentials Undergo Propagation 212
Several Properties 162
The Extracellular Concentrations of Several Ions
Receptors Are Located in the Target Cell’s Plasma Influence Neuronal Excitability 217
Membrane or Inside the Target Cell 164
Receptors Are Subject to Down-Regulation and 7.4 Signal Transmission at Synapses 218
Up-Regulation 164 Electrical Synapses Involve Gap Junctions 218
6.4 Signal Transduction Pathways 166 Chemical Synapses Involve the Release of
Neurotransmitter into a Synaptic Cleft 219
Signal Transduction Begins with Binding of Messenger
and Ends with a Cellular Response 166 Neurotransmitters Excite or Inhibit the
Postsynaptic Cell 221
Lipid-Soluble Extracellular Messengers Activate
Signaling Pathways by Binding to Intracellular There Are Two Main Types of Neurotransmitter
Receptors 168 Receptors: Ionotropic and Metabotropic 221
Water-Soluble Extracellular Messengers Activate Neurotransmitter Can Be Removed in Different
Signaling Pathways by Binding to Plasma Membrane Ways 223
Receptors 169 Postsynaptic Potentials Are Summated 224
Signal Amplification Promotes a Greater Cellular Presynaptic Modulation Regulates Neurotransmitter
Response 180 Release 225
Signal Termination Prevents Overstimulation of the
7.5 Neurotransmitters 226
Target Cell 180
There Are Several Types of Small-Molecule
6.5 Comparison of the Nervous and Endocrine Neurotransmitters 228
Systems 182 Neuropeptides Are Composed of Amino Acids Linked
Critical Thinking 171 / From Research to Reality 183 by Peptide Bonds 230

Contents • xi
 7.6 Neural Circuits 231 Thermal Sensations Provide Information About
How Hot or Cold the Skin Is 289
Critical Thinking 216 / From Research to Reality 233
Pain Sensations Protect the Body from Stimuli That Can
Cause Tissue Damage 290
Proprioceptive Sensations Provide Information About
8 The Central Nervous System 238
Muscle and Joint Position 294
8.1 Spinal Cord 239 Two Major Pathways Convey Somatic Sensory Input to
The Spinal Cord Is Protected by Vertebrae and the Primary Somatosensory Cortex 296
Meninges 239 The Primary Somatosensory Cortex Allows Precise
Spinal Nerves Link the Spinal Cord to Sensory Localization of Somatic Sensory Stimuli 297
Receptors and Effectors 239 The Somatosensory Association Area Permits
The Internal Organization of the Spinal Cord Allows Recognition of Somatic Sensory Stimuli 297
Processing of Sensory Input and Motor Output 239 Visceral Sensations Provide Input About Internal
The Spinal Cord Propagates Signals Along Sensory Conditions 297
and Motor Tracts and Coordinates Reflexes 242
9.3 The Olfactory System 298
8.2 Brain 244
The Olfactory Epithelium Contains the Receptors
The Brain Is Protected in Many Ways 244 for Smell 298
The Brain Depends on a Continuous Supply of Oxygen Olfactory Transduction Converts an Olfactory
and Glucose by the Blood 247 Stimulus into a Receptor Potential 299
Cranial Nerves Link the Brain to Sensory Receptors and The Olfactory Pathway Extends from Olfactory
Effectors 248 Receptors to the Olfactory Areas of the Brain 300
The Different Parts of the Brain Perform a Variety of The Threshold for Detecting Odors Is Low 300
Functions 249
The Olfactory System Rapidly Adapts 301
8.3 Integrative Functions of the Cerebrum 260
Wakefulness and Sleep Involve Multiple Areas 9.4 The Gustatory System 301
of the Brain 260 Taste Buds Contain the Receptors for Taste 301
Language Is an Important Form Taste Transduction Converts a Gustatory Stimulus
of Communication 263 into a Receptor Potential 301
Emotions Can Trigger Autonomic and The Gustatory Pathway Extends from Taste Receptors
Somatic Motor Responses 264 to the Gustatory Cortex 303
Motivation Is Promoted by the Mesolimbic Dopamine The Threshold for Detecting Taste Can Vary 304
Pathway 264
Taste Adaptation Occurs at Many Levels 304
Learning and Memory Allow Us to Acquire, Store, and
Recall Information 267 9.5 The Visual System 304
Critical Thinking 259 / From Research to Reality 271 Visible Light Is the Part of the Electromagnetic
Spectrum That the Eye Can Detect 304
Accessory Structures Protect, Lubricate,
9 Sensory Systems 275 and Move the Eye 305
The Eye Consists of Several Functional
9.1 Overview of Sensation 276 Components 306
The Process of Sensation Involves Four Events 276
The Eye Forms Images of Objects on the Retina 308
There Are Different Types of Sensory Receptors 276
Convergence of the Eyes Maintains Binocular Vision 311
Sensory Neurons Have Receptive Fields 277
Two Types of Photoreceptors Are Required for Normal
Sensory Coding Distinguishes the Attributes Vision 312
of a Stimulus 279
Both Eyes Receive Input from the Left and Right Visual
A Sensory Pathway Conveys Sensory Information 286 Fields 320
9.2 The Somatic Sensory System 286 The Visual Pathway Extends from Photoreceptors to the
Visual Areas of the Brain 320
Tactile Sensations Allow Us to Feel Touch, Pressure,
Vibration, Itch, and Tickle 286 The Primary Visual Cortex Perceives Light 320

xii • Contents
 The Visual Association Area Performs Several Autonomic Reflexes Help Maintain Homeostasis 353
Functions 322 Autonomic Control Centers Are Present in the Brain
The Visual System Can Undergo Light and and Spinal Cord 354
Dark Adaptation 322 Some Autonomic Responses Can Be Voluntarily
9.6 The Auditory and Vestibular Systems 322 Controlled via Biofeedback 355
The Ear Has Different Functional Components 322 10.2 Somatic Nervous System 356
Sound Waves Are Generated from a Vibrating The Somatic Nervous System Regulates the Activity
Object 325 of Skeletal Muscle 356
Transmission of Sound Waves Through the Ear Involves A Somatic Motor Pathway Is Comprised of a Somatic
Several Steps 326 Motor Neuron and Skeletal Muscle 356
Inner Hair Cells Are Responsible for Sound The Neuromuscular Junction Is the Site Where a
Transduction 327 Somatic Motor Neuron Communicates with a Skeletal
Outer Hair Cells Enhance the Sensitivity of Inner Hair Muscle Fiber 356
Cells 328 The Events at the NMJ Can Be Altered by
Pitch Discrimination Depends on Which Region of the Chemicals 358
Basilar Membrane Vibrates 328 Critical Thinking 350 / From Research to Reality 361
Loudness Discrimination Depends on How Much the
Basilar Membrane Vibrates 329
The Auditory Pathway Conveys Sound Input to the
Auditory Cortex 329 11 Muscle 364
The Primary Auditory Cortex Perceives Sound 330 11.1 Overview of Muscle 365
The Auditory Association Area Allows You to Recognize Three Types of Muscle Exist in the Body 365
a Sound 330
Muscle Performs a Variety of Functions 366
Deafness Results from Defects in the Conductive or
Muscle Has Several Important Properties 366
Neural Pathways Associated with the Ear 330
Equilibrium Is the Sense of Balance 330 11.2 Organization of Skeletal Muscle 367
The Otolithic Organs Detect Linear Acceleration or The Components of a Skeletal Muscle Fiber Allow
Deceleration and Head Tilt 330 Contraction to Occur 367
The Semicircular Ducts Detect Rotational Acceleration There Are Three Types of Muscle Proteins 372
or Deceleration 332 11.3 Contraction and Relaxation of Skeletal
Equilibrium Pathways Convey Vestibular Input to Muscle Fibers 375
Different Parts of the Brain 334
Muscle Contraction Occurs by the Sliding Filament
Critical Thinking 288 / From Research to Reality 335 Mechanism 375
The Contraction Cycle Involves Four Major Steps 375
The Neuromuscular Junction Is the Synapse Between a
10 Autonomic and Somatic Nervous Somatic Motor Neuron and a Skeletal Muscle Fiber 377
Systems 341 The Skeletal Muscle Action Potential Has Two Main
10.1 Autonomic Nervous System 342 Phases: A Depolarizing Phase and a Repolarizing
Phase 377
The Autonomic Nervous System Regulates the Activity
of Smooth Muscle, Cardiac Muscle, and Glands 342 Excitation–Contraction Coupling in Skeletal Muscle
Links the Muscle Action Potential to Muscle
An Autonomic Motor Pathway Is Comprised of Two
Contraction 378
Autonomic Motor Neurons and a Visceral Effector 342
Skeletal Muscle Relaxes in Response to a Decrease
The Neuroeffector Junction Is the Site Where an
in the Sarcoplasmic Ca2+ Concentration 379
Autonomic Postganglionic Neuron Communicates
with a Visceral Effector 345 11.4 ATP Production in Skeletal Muscle 383
The Autonomic Nervous System Uses Different Types Creatine Phosphate Is the First Source of ATP During
of Neurotransmitters and Receptors 346 Muscle Contraction 383
The Autonomic Nervous System Performs Anaerobic Glycolysis Produces ATP When Oxygen
a Variety of Functions 351 Levels Are Low 384

Contents • xiii
 Aerobic Respiration Generates ATP When Sufficient Several Factors Can Regulate Smooth Muscle
Oxygen Is Available 384 Activity 402
Several Factors Contribute to Muscle Fatigue 384 The Stress–Relaxation Response Allows Changes in
Oxygen Consumption Increases for a While After Smooth Muscle Length Without Affecting the Ability
Exercise 385 to Contract 402
Smooth Muscle Produces ATP by Aerobic Respiration
11.5 Skeletal Muscle Mechanics 385 and Anaerobic Glycolysis 402
A Motor Unit Is Comprised of a Somatic Motor Neuron 11.9 Regeneration of Muscle 402
and Its Muscle Fibers 385
Critical Thinking 382 / From Research to Reality 405
A Muscle Twitch Consists of Latent, Contraction, and
Relaxation Periods 385
Graded Contractions Can Occur in Skeletal Muscle 386
Numerous Factors Determine Muscle Tension 386 12 Control of Body Movement 411
Muscle Tone Is Established by Different Motor Units 12.1 Overview of Motor Control 412
That Are Alternately Active and Inactive 388
Lower Motor Neurons Provide Output from
Movement Involves the Interaction of Skeletal Muscles, the CNS to Skeletal Muscle Fibers 412
Tendons, and Bones 388
There Are Four Sources of Input to Lower Motor
There Are Two Major Categories of Muscle Neurons 412
Contractions: Isotonic and Isometric 391
12.2 Local Level of Motor Control 413
11.6 Types of Skeletal Muscle Fibers 392 Somatic Reflexes Allow Fast, Involuntary Contractions
Slow Oxidative Fibers Have a High Resistance to of Skeletal Muscle 413
Fatigue 392
Central Pattern Generators Are Responsible for
Fast Oxidative–Glycolytic Fibers Have a Moderate Locomotion 419
Resistance to Fatigue 392
12.3 Control of Movement by the Cerebral
Fast Glycolytic Fibers Have a Low Resistance to
Fatigue 392 Cortex 419
The Three Types of Skeletal Muscle Fibers Are The Premotor Cortex Creates a Motor Plan 419
Distributed Differently and Recruited in a The Primary Motor Cortex Controls the Execution of
Certain Order 392 Voluntary Movements 420
Exercise Can Induce Changes in the Different Types of The Primary Motor Cortex Gives Rise to the Direct
Skeletal Muscle Fibers 393 Motor Pathways 420

11.7 Cardiac Muscle 394 12.4 Control of Movement by the Brain

Stem 422
11.8 Smooth Muscle 396
Brain Stem Motor Centers Give Rise to the Indirect
The Components of a Smooth Muscle Fiber Allow Motor Pathways 422
Considerable Tension to Develop During Contraction 396
The Vestibular Nuclei Help Control Posture
Contraction and Relaxation Occur More Slowly in in Response to Changes in Balance 422
Smooth Muscle Than in Striated Muscle 396
The Reticular Formation Helps Regulate Posture
Smooth Muscle Tone Allows Maintenance of Steady and Muscle Tone During Ongoing Movements 423
Pressure 398
The Superior Colliculus Promotes Reflexive
The Autonomic Nervous System Regulates Smooth Movements of the Head and Trunk and Saccadic
Muscle 398 Eye Movements 423
Two Forms of Smooth Muscle Exist: Single-Unit and The Red Nucleus Helps Control Voluntary Movements
Multi-Unit 398 of the Upper Limbs 423
Smooth Muscle Can Exhibit Autorhythmicity 399
12.5 The Basal Nuclei and Motor Control 424
Contractile Smooth Muscle Fibers Can Produce Action
Potentials When Excited by Autorhythmic Signals or 12.6 Modulation of Movement by the
Other Stimuli 399 Cerebellum 426
Excitation–Contraction Coupling in Smooth Muscle Critical Thinking 427 / From Research to
Involves Several Mechanisms 400 Reality 428

xiv • Contents
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13 The Endocrine System 432 13.9 Other Endocrine Organs and Tissues 469
13.1 Overview of the Endocrine System 433 The Skin Secretes Cholecalciferol 469
The Endocrine System Consists of All Glands, Organs, The Thymus Secretes Hormones That Promote Immune
and Tissues That Contain Hormone-Secreting Cells 433 Function 471
Hormones Influence Target Cells by Binding The Heart Secretes Atrial Natriuretic Peptide 471
to Receptors 433 The Liver Produces Two Hormones 471
Hormones Are Chemically Classified as Lipid-Soluble or The Stomach and Small Intestine Secrete Hormones
Water-Soluble 434 That Regulate Digestive Activities 471
Hormones Circulate Through the Blood in Free Form or The Kidneys Secrete Calcitriol and Erythropoietin 471
Bound to Transport Proteins 435
Adipose Tissue Secretes Leptin 472
A Hormone’s Mechanism of Action Depends on the
The Placenta Secretes Several Hormones That Help
Signaling Pathway That It Activates 435
Maintain Pregnancy and Prepare the Maternal Body
Hormones Can Have Permissive, Synergistic, for Birth 472
or Antagonistic Effects 439
13.10 Endocrine Control of Growth 472
Hormone Secretion Is Controlled in a Variety of
Ways 440 Bones Grow in Length and in Thickness 472
Bones Undergo Remodeling 472
13.2 Pituitary Gland 440
Several Hormones Affect Bone Growth 474
The Anterior Pituitary Secretes Many Hormones 442
Factors Other Than Hormones Also Influence
The Posterior Pituitary Releases Oxytocin Bone Growth 474
and Antidiuretic Hormone 448
13.11 The Stress Response 475
13.3 Thyroid Gland 452
The Fight-or-Flight Response Allows the Body to
Thyroid Follicles Form, Store, and Release Thyroid Handle Stress Quickly 475
Hormones 452
The Resistance Reaction Provides a Longer-lasting
Thyroid Hormones Increase Basal Metabolic Rate and Response to Stress 475
Have Other Effects 453
After the Resistance Reaction Is Exhaustion 475
Secretion of Thyroid Hormones Is Regulated by the
Hypothalamus and Anterior Pituitary 454 Stress Can Lead to Disease 475

Calcitonin Lowers the Blood Calcium Level 454 Critical Thinking 470 / From Research to Reality 478

13.4 Parathyroid Glands 455

13.5 Adrenal Glands 457 14 The Cardiovascular System:
The Adrenal Cortex Consists of Three Zones That The Heart 484
Secrete Hormones 457
14.1 Basic Design of the Cardiovascular
The Adrenal Medulla Is an Extension of the Sympathetic System 485
Nervous System That Secretes Hormones 461
The Heart Pumps Blood Through the Pulmonary and
13.6 Pineal Gland 461 Systemic Circulations 485
13.7 Pancreas 463 Blood Is Distributed in the Systemic Circulation Mainly
via Parallel Flow 487
The Pancreas Secretes Several Hormones, Including
Insulin and Glucagon 463 14.2 Organization of the Heart 488
Insulin Stimulates Uptake of Glucose, Fatty Acids, and The Pericardium Protects and Anchors the
Amino Acids and Synthesis of Glycogen, Triglycerides, Three-Layered Heart 488
and Proteins 463 The Heart Contains Four Chambers and Is
Glucagon Has Functions That Are Antagonistic to Those Associated with Several Great Vessels 490
of Insulin 466 Heart Valves Ensure One-Way Blood Flow 491
Insulin and Glucagon Secretion Are Regulated The Fibrous Skeleton of the Heart Prevents
by the Blood Glucose Concentration and Overstretching of Heart Valves 492
Other Factors 467
The Coronary Circulation Supplies Blood
13.8 Ovaries and Testes 469 to the Heart Wall 492

Contents • xv
 14.3 Cardiac Muscle Tissue and the Cardiac Diffusion Is the Most Important Method of Capillary
Conduction System 495 Exchange 531
Interconnected Cardiac Muscle Fibers Act as a Transcytosis Allows Transport of Large Molecules Across
Functional Syncytium 495 Capillary Walls 532
The Conduction System of the Heart Ensures Bulk Flow Regulates the Relative Volumes of Blood and
Coordinated Contraction 497 Interstitial Fluid 532
Contractile Fibers Produce Action Potentials The Lymphatic System Begins in the Tissues Close to
in Response to Autorhythmic Fibers 499 Blood Capillaries 533
Excitation–Contraction Coupling Links Cardiac Action 15.3 Hemodynamics 536
Potentials to Cardiac Contraction 500 Blood Flow Is Affected by Pressure and Resistance 536
Cardiac Muscle Fibers Have a Long Refractory Blood Flow Through Blood Vessels Can Be Laminar or
Period 501 Turbulent 539
Cardiac Muscle Produces ATP via Aerobic Blood Pressure Is Hydrostatic Pressure Exerted by Blood
Respiration 502 on Blood Vessel Walls 540
The Electrocardiogram Records Electrical Signals Vascular Compliance Is High in Veins and Low in
Generated by the Heart 502 Arteries 543
ECG Waves Predict the Timing of Atrial and Ventricular Velocity of Blood Flow Is Inversely Proportional to the
Systole and Diastole 502 Cross-Sectional Area 544
14.4 The Cardiac Cycle 506 Venous Return Brings Blood Back to the Heart 544
The Cardiac Cycle Has Five Phases 506 15.4 Control of Blood Flow 546
At Rest a Cardiac Cycle Lasts About 0.8 Intrinsic Control of Blood Flow Includes Physical
Seconds 508 Changes and Local Mediators 548
Two Major Heart Sounds Can Be Heard During Each Extrinsic Control of Blood Flow Includes Nerves and
Cardiac Cycle 508 Hormones 550
14.5 Cardiac Output 508 15.5 Regulation of Mean Arterial Pressure 552
Stroke Volume Is Regulated by Preload, Contractility, The Nervous System Is an Important Regulator of Mean
and Afterload 508 Arterial Pressure 553
Heart Rate Is Regulated Mainly by the ANS and Certain Hormones Also Regulate Mean Arterial Pressure 556
Chemicals 512
15.6 Shock and Homeostasis 556
14.6 Exercise and the Heart 517
There Are Four Types of Shock 556
Critical Thinking 514 / From Research to Reality 520
The Body’s Response to Shock Involves Several
Compensatory Mechanisms 557
Shock Has Many Signs and Symptoms 557
15 The Cardiovascular System: Blood Critical Thinking 559 / From Research to Reality 560
Vessels and Hemodynamics 524
15.1 Overview of the Vasculature 525
The Layers of a Blood Vessel Contribute to Vessel
Function 526 16 The Cardiovascular System:
Arteries Carry Blood Away from the Heart 526 The Blood 565
Arterioles Deliver Blood to Capillaries 527 16.1 Overview of Blood 566
Capillaries Are the Sites of Nutrient and Waste Blood Performs Important Functions in the Body 566
Exchange 527 Plasma and Cellular Elements Are the Two Main
Venules Drain Blood from Capillaries 528 Components of Blood 566
Veins Carry Blood back to the Heart 528 Blood Cells Are Formed in Bone Marrow 569
Systemic Veins and Venules Function as Blood 16.2 Erythrocytes 570
Reservoirs 530
Erythrocytes Transport Oxygen and Carbon
15.2 Capillary Exchange and Lymphatics 531 Dioxide 570

xvi • Contents
 Erythrocytes Are Produced via Erythropoiesis 571 Antigens Are Processed and Then Presented 605
The Erythrocyte Life Cycle Is About 120 Days 572 Cytokines Have Important Roles in Immunity 607
16.3 Leukocytes 576 Cell-Mediated Immunity Uses T Cells to Eliminate
Specific Antigens 607
The Five Types of Leukocytes Are Grouped into Two
Categories 576 Antibody-Mediated Immunity Uses Antibodies to
Inactivate Specific Antigens 610
Leukocytes Defend the Body Against Invading
Pathogens and Abnormal Cells 576 Immunological Memory Allows the Immune System to
Remember Previously Encountered Antigens 613
Leukocytes Are Produced via Leukopoiesis 577
There Are Four Ways to Acquire Adaptive Immunity 615
16.4 Platelets 577
Self-Recognition and Self-Tolerance Prevent
16.5 Hemostasis 577 the Immune System from Attacking the Body’s Own
Vascular Spasm Occurs Immediately After Vessel Tissues 618
Injury 578 Allergic Reactions May Be Immediate or Delayed 618
A Platelet Plug Is a Mass of Platelets That Accumulates Critical Thinking 617 / From Research to Reality 621
at the Site of Injury 579
Blood Clotting Results in the Formation of Fibrin
Threads at the Injured Area 580
16.6 Blood Groups and Blood Types 584 18 The Respiratory System 625
The ABO Blood Group Is Determined by the Presence 18.1 Overview of the Respiratory
or Absence of A and B Antigens 584
System 626
An Incompatible Transfusion Causes Agglutination 584
Respiration Supplies the Body with O2 and
The Rh Blood Group Is Based on the Presence or Removes CO2 626
Absence of Rh Antigens 586
The Respiratory System Is Comprised
Critical Thinking 575 / From Research to Reality 589 of Several Organs 626
The Respiratory Mucosa Produces Mucus That Traps
Particles and Lubricates the Airways 629
The Nose Brings Air into the Respiratory System 629
17 The Immune System 593 The Pharynx Is a Common Passageway for Air and
17.1 Components of the Immune System 594 Food 629
The Cells of the Immune System Include Leukocytes, The Larynx Routes Air and Food into the Proper
Mast Cells, and Dendritic Cells 594 Channels and Also Causes Vocalization 629
There Are Several Types of Lymphoid Organs and The Trachea Carries Air to the Primary Bronchi 629
Tissues 595 The Primary Bronchi Transport Air to the Lungs 629
17.2 Innate Immunity 596 The Lungs Contain Most of the Components of the
The First Line of Defense Consists of the External Respiratory System 631
Physical and Chemical Barriers of the Body 596 The Alveoli Are the Sites of Gas Exchange Between
The Second Line of Defense Is Comprised of Various Air and Blood 631
Internal Defenses 597 There Are Two Functional Zones of the Respiratory
System 632
17.3 Adaptive Immunity 602
The Pulmonary Circulation Has a High Rate of Flow,
B Cells and T Cells Are Important Components of a Low Resistance, and a Low Pressure 632
Adaptive Immunity 602
Adaptive Immunity Can Be Mediated by Cells or 18.2 Ventilation 632
Antibodies 602 The Breathing Cycle Has Three Phases 633
Clonal Selection Increases the Number of B Cells and Several Factors Affect Ventilation 636
T Cells that Can Respond to an Antigen 602 Ventilation and Perfusion of the Alveoli Are Matched
Antigens Trigger Immune Responses 604 for Optimal Gas Exchange 639
Major Histocompatibility Complex Proteins Are Types of There Are Different Patterns of Respiratory
Self-Antigens 605 Movements 640

Contents • xvii
 18.3 Lung Volumes and Capacities 640 19.5 Tubular Reabsorption and Tubular
18.4 Exchange of Oxygen and Carbon Secretion 675
Dioxide 643 There Are Two Types of Reabsorption Routes:
Paracellular and Transcellular 675
Two Gas Laws—Dalton’s Law and Henry’s Law—Are
Important for Understanding How Gas Exchange Transport of Substances Across the Tubular Wall Often
Occurs 643 Involves the Use of Transport Proteins 676
There Are Two Types of Gas Exchange: Pulmonary Water Reabsorption May Be Obligatory or
and Systemic 644 Facultative 676
Different Substances Are Reabsorbed or Secreted to
18.5 Transport of Oxygen and Carbon
Varying Degrees in Different Parts of the Renal Tubule
Dioxide 646 and Collecting Duct 676
Oxygen Is Transported Through the Blood Mainly Tubular Reabsorption and Tubular Secretion Are Subject
by Hemoglobin 646 to Hormonal Regulation 680
The PO2 Determines How Much Oxygen Binds to
19.6 Production of Dilute and Concentrated
Hemoglobin 649
Urine 684
Several Factors Affect the Affinity of Hemoglobin
for Oxygen 649 The Production of Dilute Urine Allows the Kidneys
to Get Rid of Excess Water 684
Fetal Hemoglobin and Adult Hemoglobin Have
Different Affinities for Oxygen 651 The Production of Concentrated Urine Allows
the Kidneys to Conserve Water 684
Carbon Dioxide Is Transported Through the Blood
in Three Forms 651 19.7 Evaluation of Kidney Function 688
Gas Exchange and Transport Can Be Summarized 652 A Urinalysis Examines the Various Properties of
Urine 688
18.6 Control of Ventilation 653
Blood Tests for Renal Function Include
The Respiratory Center Controls Breathing 653 the Blood Urea Nitrogen Test and Measurement
The Respiratory Center Is Subject to Regulation 654 of Plasma Creatinine 688
18.7 Exercise and the Respiratory System 657 Renal Plasma Clearance Indicates How Effectively the
Kidneys Are Removing a Substance from Blood
Critical Thinking 648 / From Research to Reality 659
Plasma 690
Renal Failure Occurs Because of Inadequate Kidney
Function 690
19 The Urinary System 664 19.8 Urine Transportation, Storage, and
19.1 Overview of Kidney Functions 665 Elimination 693
The Ureters Carry Urine to the Urinary Bladder 693
19.2 Organization of the Kidneys 665
The Urinary Bladder Stores Urine 694
Nephrons Perform the Main Functions of the
Kidneys 667 The Urethra Conveys Urine to the Outside
Environment 695
The Extensive Blood Supply of the Kidneys Contributes
to Renal Function 668 19.9 Waste Management in Other Body
The Juxtaglomerular Apparatus Consists of Part Systems 695
of the Distal Tubule and Afferent Arteriole 668 Critical Thinking 687 / From Research to Reality 697
19.3 Overview of Renal Physiology 668
19.4 Glomerular Filtration 669
20 Fluid, Electrolyte, and Acid–Base
The Renal Corpuscle Contains a Filtration Membrane 669
Homeostasis 702
Glomerular Filtration Is Determined by the Balance of
Four Pressures 672 20.1 Fluid Compartments and
The Glomerular Filtration Rate Is an Important Aspect Fluid Balance 703
of Kidney Function 672 The Body Can Gain or Lose Water 704
The Glomerular Filtration Rate Is Regulated in Different Body Water Gain Is Regulated Mainly by the Volume of
Ways 673 Water Intake 704

xviii • Contents
 Body Water or Solute Loss Is Regulated Mainly by 21.4 Stomach 732
Control of Their Loss in Urine 705
Mechanical Digestion in the Stomach Involves
Water Can Move Between Body Fluid Propulsion and Retropulsion 732
Compartments 706
Chemical Digestion in the Stomach Occurs as Food
20.2 Electrolytes in Body Fluids 708 Mixes with Gastric Juice 733
The Electrolyte Concentrations in Body Fluids Can Be Vomiting Rapidly Expels the Contents
Measured in Milliequivalents per Liter 708 of the GI Tract 735
Electrolytes Perform Many Functions in the Body 709 21.5 Pancreas, Liver, and Gallbladder 736
20.3 Acid–Base Balance 710 The Pancreas Secretes Pancreatic Juice 736
Buffer Systems Convert Strong Acids and Bases into The Liver Secretes Bile and Performs Many Other
Weak Acids and Bases 711 Functions 737
Exhalation of Carbon Dioxide Helps Eliminate H+ Ions 712 The Gallbladder Stores and Concentrates Bile 740
The Kidneys Remove H+ Ions by Excreting Them into 21.6 Small Intestine 740
Urine 712
The Small Intestine Secretes Intestinal Juice 742
There Are Different Types of Acid–Base Imbalances 714
Brush-Border Enzymes Are Attached to the Microvilli
Critical Thinking 715 / From Research to Reality 717 of Small Intestinal Absorptive Cells 742
Mechanical Digestion in the Small Intestine Involves
Segmentation and the Migrating Motility
21 The Digestive System 721 Complex 742
Chemical Digestion in the Small Intestine Occurs as
21.1 Overview of the Digestive System 722 Chyme Mixes with Intestinal Juice, Pancreatic Juice,
The Digestive System Consists of the Gastrointestinal and Bile 742
Tract and the Accessory Digestive Organs 722 Most Nutrients and Water Are Absorbed in the Small
There Are Six Basic Digestive Processes 722 Intestine 744
The Wall of the GI Tract Is Comprised of Four Functional 21.7 Large Intestine 750
Layers 723
Mechanical Digestion in the Large Intestine Involves
The GI Tract Has Neural Innervation 724 Haustral Churning, Peristalsis, and Mass Movement 750
GI Smooth Muscle Is Autorhythmic and Promotes Two Chemical Digestion in the Large Intestine Occurs via
Major Patterns of Motility 725 Bacteria 750
21.2 Mouth 727 Feces Are Formed in the Large Intestine 750
The Cheeks and Lips Keep Food in the Mouth During The Defecation Reflex Expels Feces from the Body 751
Chewing 728
21.8 Phases of Digestion 753
The Palate Prevents Food from Entering the Nasal
Cavity 728 The Cephalic Phase Prepares the Mouth and Stomach
for Food That Is About to Be Eaten 753
The Tongue Moves Food Toward the Pharynx and
Produces Lingual Lipase 728 The Gastric Phase Promotes Gastric Juice Secretion and
Gastric Motility 753
The Salivary Glands Secrete Saliva 728
The Intestinal Phase Promotes Digestion in the Small
The Teeth Physically Break Down Food 729 Intestine and Slows Digestion in the Stomach 754
Mechanical Digestion in the Mouth Involves There Are Many Hormones of the Digestive System 754
Chewing 729
21.9 Transport of Lipids by Lipoproteins 755
Chemical Digestion in the Mouth Occurs as Food Mixes
with Saliva 729 Critical Thinking 749 / From Research to Reality 758

21.3 Pharynx and Esophagus 729

The Pharynx Conveys Food from the Mouth to the
Esophagus 729 22 Metabolic Adaptations, Energy
The Esophagus Transports Food to the Stomach 729 Balance, and Temperature
Regulation 764
Deglutition Is the Process by Which Food Is
Swallowed 730 22.1 Metabolic Adaptations 765
Contents • xix
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