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Medical Physiology

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Medical Physiology

THIRD EDITION

WALTER F. BORON, MD, PhD


Professor
David N. and Inez Myers/Antonio Scarpa Chairman
Department of Physiology and Biophysics
Case Western Reserve University
Cleveland, Ohio

EMILE L. BOULPAEP, MD
Professor
Department of Cellular and Molecular Physiology
Yale University School of Medicine
New Haven, Connecticut
Table of Contents

Instructions for online access

Cover image

Title Page

Copyright

Contributors

Video Table of Contents

Preface to the Third Edition


The eBook

Acknowledgments

Preface to the First Edition


Target Audience

Content of the Textbook

Emphasis of the Textbook

Creating the Textbook

Special Features

Acknowledgments

Section I Introduction
Chapter 1 Foundations of Physiology
References

Section II Physiology of Cells and Molecules

Chapter 2 Functional Organization of the Cell


Structure of Biological Membranes

Function of Membrane Proteins

Cellular Organelles and the Cytoskeleton

Synthesis and Recycling of Membrane Proteins

Specialized Cell Types

References

Chapter 3 Signal Transduction


Mechanisms of Cellular Communication

Receptors That are Ion Channels

Receptors Coupled to G Proteins

Receptors That are Catalytic

Nuclear Receptors

References

Chapter 4 Regulation of Gene Expression


From Genes to Proteins

The Promoter and Regulatory Elements

Transcription Factors

Regulation of Inducible Gene Expression by Signal-Transduction Pathways

Epigenetic Regulation of Gene Expression

Post-Translational Regulation of Gene Expression

References

Glossary

Chapter 5 Transport of Solutes and Water


The Intracellular and Extracellular Fluids

Solute Transport Across Cell Membranes

Regulation of Intracellular Ion Concentrations

Water Transport and the Regulation of Cell Volume

Transport of Solutes and Water Across Epithelia

References

Chapter 6 Electrophysiology of the Cell Membrane


Ionic Basis of Membrane Potentials

Electrical Model of a Cell Membrane

Molecular Physiology of Ion Channels

References

Chapter 7 Electrical Excitability and Action Potentials


Mechanisms of Nerve and Muscle Action Potentials

Physiology of Voltage-Gated Channels and Their Relatives

Propagation of Action Potentials

References

Chapter 8 Synaptic Transmission and the Neuromuscular Junction


Mechanisms of Synaptic Transmission

Synaptic Transmission at the Neuromuscular Junction

Toxins and Drugs Affecting Synaptic Transmission

References

Chapter 9 Cellular Physiology of Skeletal, Cardiac, and Smooth Muscle


Skeletal Muscle

Cardiac Muscle

Smooth Muscle

Diversity among Muscles

References
Section III The Nervous System

Chapter 10 Organization of the Nervous System


Cells of the Nervous System

Development of Neurons and Glial Cells

Subdivisions of the Nervous System

References

Chapter 11 The Neuronal Microenvironment


Cerebrospinal Fluid

Brain Extracellular Space

The Blood-Brain Barrier

Glial Cells

References

Chapter 12 Physiology of Neurons


Signal Conduction in Dendrites

Control of Spiking Patterns in the Soma

Axonal Conduction

References

Chapter 13 Synaptic Transmission in the Nervous System


Neuronal Synapses

Neurotransmitter Systems of the Brain

Fast Amino Acid–Mediated Synapses in the CNS

Plasticity of Central Synapses

References

Chapter 14 The Autonomic Nervous System


Organization of the Visceral Control System

Synaptic Physiology of the Autonomic Nervous System

Central Nervous System Control of the Viscera


References

Chapter 15 Sensory Transduction


Chemoreception

Visual Transduction

Vestibular and Auditory Transduction: Hair Cells

Somatic Sensory Receptors, Proprioception, and Pain

References

Chapter 16 Circuits of the Central Nervous System


Elements of Neural Circuits

Simple, Stereotyped Responses: Spinal Reflex Circuits

Rhythmic Activity: Central Pattern Generators

Spatial Representations: Sensory and Motor Maps in the Brain

Temporal Representations: Time-Measuring Circuits

References

Section IV The Cardiovascular System

Chapter 17 Organization of the Cardiovascular System


Elements of the Cardiovascular System

Hemodynamics

How Blood Flows

Origins of Pressure in the Circulation

How to Measure Blood Pressure, Blood Flow, and Cardiac Volumes

References

Chapter 18 Blood
Blood Composition

Blood Viscosity

Hemostasis and Fibrinolysis

References
Chapter 19 Arteries and Veins
Arterial Distribution and Venous Collection Systems

Elastic Properties of Blood Vessels

References

Chapter 20 The Microcirculation


Capillary Exchange of Solutes

Capillary Exchange of Water

Lymphatics

Regulation of the Microcirculation

References

Chapter 21 Cardiac Electrophysiology and the Electrocardiogram


Electrophysiology of Cardiac Cells

The Electrocardiogram

Cardiac Arrhythmias

References

Chapter 22 The Heart as a Pump


The Cardiac Cycle

Cardiac Dynamics

From Contractile Filaments to a Regulated Pump

References

Chapter 23 Regulation of Arterial Pressure and Cardiac Output


Short-Term Regulation of Arterial Pressure

Regulation of Cardiac Output

Matching of Venous Return and Cardiac Output

Intermediate- and Long-Term Control of the Circulation

References

Chapter 24 Special Circulations


The Brain
The Heart

The Skeletal Muscle

The Splanchnic Organs

The Skin

References

Chapter 25 Integrated Control of the Cardiovascular System


Interaction among the Different Cardiovascular Control Systems

Response to Erect Posture

Responses to Acute Emotional Stress

Response to Exercise

Response to Hemorrhage

References

Section V The Respiratory System

Chapter 26 Organization of the Respiratory System


Comparative Physiology of Respiration

Organization of the Respiratory System in Humans

Lung Volumes and Capacities

References

Chapter 27 Mechanics of Ventilation


Static Properties of the Lung

Dynamic Properties of the Lung

References

Chapter 28 Acid-Base Physiology


pH and Buffers

Acid-Base Chemistry When Is the Only Buffer

Acid-Base Chemistry in the Presence of and Buffers—The Davenport Diagram

pH Regulation of Intracellular Fluid


References

Chapter 29 Transport of Oxygen and Carbon Dioxide in the Blood


Carriage of O2

Carriage of CO2

References

Chapter 30 Gas Exchange in the Lungs


Diffusion of Gases

Diffusion and Perfusion Limitations on Gas Transport

References

Chapter 31 Ventilation and Perfusion of the Lungs


Ventilation

Perfusion of the Lung

Matching Ventilation and Perfusion

References

Chapter 32 Control of Ventilation


Overview of the Respiratory Control System

Neurons That Control Ventilation

Generation of the Respiratory Rhythm

Chemical Control of Ventilation

Modulation of Ventilatory Control

References

Section VI The Urinary System

Chapter 33 Organization of the Urinary System


Functional Anatomy of the Kidney

Main Elements of Renal Function


Measuring Renal Clearance and Transport

The Ureters and Bladder

References

Chapter 34 Glomerular Filtration and Renal Blood Flow


Glomerular Filtration

Renal Blood Flow

Control of Renal Blood Flow and Glomerular Filtration

References

Chapter 35 Transport of Sodium and Chloride


Na+ and Cl− Transport by Different Segments of The Nephron

Na+ and Cl−, and Water Transport at the Cellular and Molecular Level

Regulation of Na+ and Cl− Transport

References

Chapter 36 Transport of Urea, Glucose, Phosphate, Calcium, Magnesium, and Organic Solutes
Urea

Glucose

Other Organic Solutes

Phosphate

Calcium

Magnesium

References

Chapter 37 Transport of Potassium


Potassium Balance and the Overall Renal Handling of Potassium

Potassium Transport by Different Segments of the Nephron

Potassium Transport at the Cellular and Molecular Levels

Regulation of Renal Potassium Excretion

References

Chapter 38 Urine Concentration and Dilution


Water Balance and the Overall Renal Handling of Water

Water Transport by Different Segments of the Nephron

Generation of a Hyperosmotic Medulla and Urine

Regulation by Arginine Vasopressin

References

Chapter 39 Transport of Acids and Bases


Acid-Base Balance and the Overall Renal Handling of Acid

Acid-Base Transport by Different Segments of the Nephron

Acid-Base Transport at the Cellular and Molecular Levels

Regulation of Renal Acid Secretion

References

Chapter 40 Integration of Salt and Water Balance


Control of Extracellular Fluid Volume

Control of Water Content (Extracellular Osmolality)

References

Section VII The Gastrointestinal System

Chapter 41 Organization of the Gastrointestinal System


Overview of Digestive Processes

Regulation of Gastrointestinal Function

Gastrointestinal Motility

References

Chapter 42 Gastric Function


Functional Anatomy of the Stomach

Acid Secretion

Pepsinogen Secretion

Protection of the Gastric Surface Epithelium and Neutralization of Acid in the Duodenum

Filling and Emptying of the Stomach


References

Chapter 43 Pancreatic and Salivary Glands


Overview of Exocrine Gland Physiology

Pancreatic Acinar Cell

Pancreatic Duct Cell

Composition, Function, and Control of Pancreatic Secretion

Salivary Acinar Cell

Salivary Duct Cell

Composition, Function, and Control of Salivary Secretion

References

Chapter 44 Intestinal Fluid and Electrolyte Movement


Functional Anatomy

Overview of Fluid and Electrolyte Movement in the Intestines

Cellular Mechanisms of Na+ Absorption

Cellular Mechanisms of Cl− Absorption and Secretion

Cellular Mechanisms of K+ Absorption and Secretion

Regulation of Intestinal Ion Transport

References

Chapter 45 Nutrient Digestion and Absorption


Carbohydrate Digestion

Carbohydrate Absorption

Protein Digestion

Protein, Peptide, and Amino-Acid Absorption

Lipid Digestion

Lipid Absorption

Digestion and Absorption of Vitamins and Minerals

Nutritional Requirements

References

Chapter 46 Hepatobiliary Function


Overview of Liver Physiology

Functional Anatomy of the Liver and Biliary Tree

Uptake, Processing, and Secretion of Compounds by Hepatocytes

Bile Formation

Enterohepatic Circulation of Bile Acids

The Liver as a Metabolic Organ

References

Section VIII The Endocrine System

Chapter 47 Organization of Endocrine Control


Principles of Endocrine Function

Peptide Hormones

Amine Hormones

Steroid and Thyroid Hormones

References

Chapter 48 Endocrine Regulation of Growth and Body Mass


Growth Hormone

Growth-Promoting Hormones

Regulation of Body Mass

References

Chapter 49 The Thyroid Gland


Synthesis of Thyroid Hormones

Action of Thyroid Hormones

Hypothalamic-Pituitary-Thyroid Axis

References

Chapter 50 The Adrenal Gland


The Adrenal Cortex: Cortisol

The Adrenal Cortex: Aldosterone


The Adrenal Medulla

References

Chapter 51 The Endocrine Pancreas


Insulin

Glucagon

Somatostatin

References

Chapter 52 The Parathyroid Glands and Vitamin D


Calcium and Phosphate Balance

Physiology of Bone

Parathyroid Hormone

Vitamin D

Calcitonin and Other Hormones

References

Section IX The Reproductive System

Chapter 53 Sexual Differentiation


Genetic Aspects of Sexual Differentiation

Differentiation of the Gonads

Development of the Accessory Sex Organs

Differentiation of the External Genitalia

Endocrine and Paracrine Control of Sexual Differentiation

Puberty

References

Chapter 54 The Male Reproductive System


Hypothalamic-Pituitary-Gonadal Axis

Testosterone

Biology of Spermatogenesis and Semen


Male Sex Act

References

Chapter 55 The Female Reproductive System


Hypothalamic-Pituitary-Gonadal Axis and Control of the Menstrual Cycle

Ovarian Steroids

The Ovarian Cycle: Folliculogenesis, Ovulation, and Formation of the Corpus Luteum

The Endometrial Cycle

Female Sex Act

Menopause

References

Chapter 56 Fertilization, Pregnancy, and Lactation


Transport of Gametes and Fertilization

Implantation of the Developing Embryo

Physiology of the Placenta

The Maternal-Placental-Fetal Unit

Response of the Mother to Pregnancy

Parturition

Lactation

References

Chapter 57 Fetal and Neonatal Physiology


Biology of Fetal Growth

Development and Maturation of the Cardiopulmonary System

Cardiopulmonary Adjustments at Birth

Neonatal Physiology

References

Section X Physiology of Everyday Life

Chapter 58 Metabolism
Forms of Energy

Energy Balance

Energy Interconversion From Cycling between 6-Carbon and 3-Carbon Molecules

Energy Capture (Anabolism)

Energy Liberation (Catabolism)

Integrative Metabolism During Fasting

References

Chapter 59 Regulation of Body Temperature


Heat and Temperature: Advantages of Homeothermy

Modes of Heat Transfer

Active Regulation of Body Temperature by the Central Nervous System

Hypothermia, Hyperthermia, and Fever

References

Chapter 60 Exercise Physiology and Sports Science


Motor Units and Muscle Function

Conversion of Chemical Energy to Mechanical Work

Muscle Fatigue

Determinants of Maximal O2 Uptake and Consumption

Sweating

Endurance (Aerobic) Training

References

Chapter 61 Environmental Physiology


The Environment

Diving Physiology

High-Altitude Physiology

Flight and Space Physiology

References

Chapter 62 The Physiology of Aging


Concepts in Aging
Cellular and Molecular Mechanisms of Aging

Aging of the Human Physiological Systems

Aging Slowly

References

Index
Copyright

1600 John F. Kennedy Blvd.


Ste 1800
Philadelphia, PA 19103-2899

MEDICAL PHYSIOLOGY, THIRD EDITION


ISBN: 978-1-4557-4377-3
INTERNATIONAL EDITION
ISBN: 978-0-323-42796-8

Copyright © 2017 by Elsevier, Inc. All rights reserved.

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Previous editions copyrighted 2012, 2009, 2005, and 2003.

Library of Congress Cataloging-in-Publication Data


Names: Boron, Walter F., editor. | Boulpaep, Emile L., editor.
Title: Medical physiology / [edited by] Walter F. Boron, Emile L. Boulpaep.
Other titles: Medical physiology (Boron)
Description: Edition 3. | Philadelphia, PA : Elsevier, [2017] | Includes bibliographical references and
index.
Identifiers: LCCN 2016005260| ISBN 9781455743773 (hardcover : alk. paper) | ISBN 9780323427968
(International ed.)
Subjects: | MESH: Physiological Phenomena | Cell Physiological Phenomena
Classification: LCC QP34.5 | NLM QT 104 | DDC 612—dc23 LC record available at
http://lccn.loc.gov/2016005260

Executive Content Strategist: Elyse O'Grady


Senior Content Development Specialist: Marybeth Thiel
Publishing Services Manager: Julie Eddy
Senior Project Manager: David Stein
Design Direction: Julia Dummitt

Printed in China

Last digit is the print number: 9 8 7 6 5 4 3 2 1


Contributors

Peter S. Aronson MD
C.N.H. Long Professor of Internal Medicine
Professor of Cellular and Molecular Physiology
Section of Nephrology
Department of Internal Medicine
Yale University School of Medicine
New Haven, Connecticut

Eugene J. Barrett MD, PhD


Professor
Departments of Medicine and Pharmacology
University of Virginia School of Medicine
Charlottesville, Virginia

Paula Q. Barrett PhD


Professor
Department of Pharmacology
University of Virginia School of Medicine
Charlottesville, Virginia

Henry J. Binder MD
Professor Emeritus of Medicine
Department of Internal Medicine—Digestive Diseases
Yale University School of Medicine
New Haven, Connecticut

Walter F. Boron MD, PhD


Professor
David N. and Inez Myers/Antonio Scarpa Chairman
Department of Physiology and Biophysics
Case Western Reserve University
Cleveland, Ohio

Emile L. Boulpaep MD
Professor
Department of Cellular and Molecular Physiology
Yale University School of Medicine
New Haven, Connecticut
Lloyd Cantley MD, FASN
Professor
Department of Internal Medicine
Department of Cellular and Molecular Physiology
Yale University School of Medicine
New Haven, Connecticut

Michael J. Caplan MD, PhD


C.N.H. Long Professor and Chair
Department of Cellular and Molecular Physiology
Yale University School of Medicine
New Haven, Connecticut

Barry W. Connors PhD


Professor and Chair
Department of Neuroscience
Alpert Medical School
Brown University
Providence, Rhode Island

Arthur DuBois MD
Professor Emeritus of Epidemiology and Public Health and Cellular and Molecular Physiology
John B. Pierce Laboratory
New Haven, Connecticut

Gerhard Giebisch MD
Professor Emeritus of Cellular and Molecular Physiology
Department of Cellular and Molecular Physiology
Yale University School of Medicine
New Haven, Connecticut

Fred S. Gorelick MD
Professor
Departments of Internal Medicine and Cell Biology
Yale University School of Medicine
New Haven, Connecticut

Peter Igarashi MD
Nesbitt Chair and Head
Department of Medicine
University of Minnesota
Minneapolis, Minnesota

Ervin E. Jones MD, PhD


Retired
Department of Obstetrics and Gynecology
Yale University School of Medicine
New Haven, Connecticut

W. Jonathan Lederer MD, PhD


Director and Professor, Center for Biomedical Engineering and Technology and Department of
Physiology
University of Maryland School of Medicine
Baltimore, Maryland

George Lister MD
Jean McLean Wallace Professor of Pediatrics
Professor of Cellular and Molecular Physiology
Yale School of Medicine
New Haven, Connecticut

Charles M. Mansbach II, MD †


Professor of Medicine and Physiology
University of Tennessee Health Science Center
Memphis, Tennessee

Christopher R. Marino MD
Professor of Medicine
University of Tennessee Health Science Center
Chief of Staff
VA Medical Center
Memphis, Tennessee

Edward J. Masoro PhD


Professor Emeritus of Physiology
University of Texas Health Science Center at San Antonio
San Antonio, Texas

Sam Mesiano PhD


Professor
Department of Reproductive Biology
Case Western Reserve University
Cleveland, Ohio

Edward G. Moczydlowski PhD


Senior Associate Dean of Academic Affairs & Professor of Physiology
College of Health Sciences
California Northstate University
Elk Grove, California

Shaun F. Morrison PhD


Professor

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