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10th edition
Pulmonary
Physiology
Michael G. Levitzky
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Pulmonary
Physiology
Tenth Edition
MichaelG.Levitzky,PhD
EmeritusProfessorofPhysiology
LouisianaSfofeUniversityHealthSciencesCenter
NewOrleans,Louisiana
Now York Chicago San Francisco Athens Ijondon Madrid
MexicoCity Milan NowDelhi Singapore Sydney Toronto
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Contents
Preface
Chapter 1 Function and Structure of the Respiratory System
Functions of the Respiratory System
Structure of the Respiratory System
Key Concepts
Suggested Readings
Chapter 2 Mechanics of Breathing
Generation of a Pressure Difference Between Atmosphere and Alveoli
Pressure-Volume Relationships in the Respiratory System
Interaction of Lung and Chest Wall: The Static Pressure-Volume Curve
Airways Resistance
The Work of Breathing
Key Concepts
Clinical Problems
Suggested Readings
Chapter 3 Alveolar Ventilation
The Lung Volumes
Measurement of the Lung Volumes
Anatomic Dead Space and Alveolar Ventilation
Measurement of Alveolar Ventilation
Alveolar Ventilation and Alveolar Oxygen and Carbon Dioxide Levels
Regional Distribution of Alveolar Ventilation
The Closing Volume
The Effects of Aging
Key Concepts
Clinical Problems
Suggested Readings
Chapter 4 Blood Flow to the Lung
The Bronchial Circulation
The Functional Anatomy of the Pulmonary Circulation
Pulmonary Vascular Resistance
The Regional Distribution of Pulmonary Blood Flow: The Zones of the Lung
Hypoxic Pulmonary Vasoconstriction
Effects of Mechanical Ventilation on Pulmonary Blood Flow
Pulmonary Edema
Key Concepts
Clinical Problems
Suggested Readings
Chapter 5 Ventilation-Perfusion Relationships
The Concept of Matching Ventilation and Perfusion
Consequences of High and Low
Testing for Nonuniform Distribution of Inspired Gas and Pulmonary Blood Flow
Regional Differences and Their Consequences in the Lung
Key Concepts
Clinical Problems
Suggested Readings
Chapter 6 Diffusion of Gases and Interpretation of Pulmonary Function Tests
Fick’s Law for Diffusion
Limitations of Gas Transfer
Diffusion of Oxygen
Diffusion of Carbon Dioxide
Measurement of Diffusing Capacity
Interpretation of Pulmonary Function Tests
Key Concepts
Clinical Problems
Suggested Readings
Chapter 7 Transport of Oxygen and Carbon Dioxide in the Blood
Transport of Oxygen by the Blood
Hemoglobin and the Physiologic Implications of the Oxyhemoglobin Dissociation Curve
Influences on the Oxyhemoglobin Dissociation Curve
Transport of Carbon Dioxide by the Blood
The Carbon Dioxide Dissociation Curve
The Bohr and Haldane Effects Explained
Key Concepts
Clinical Problems
Suggested Readings
Chapter 8 Acid-Base Balance
The Chemistry of Acids, Bases, and Buffers
Buffer Systems of the Human Body
Acidosis and Alkalosis
Respiratory and Renal Compensatory Mechanisms
Clinical Interpretation of Blood Gases and Acid-Base Status
The Causes of Hypoxia
Key Concepts
Clinical Problems
Suggested Readings
Chapter 9 Control of Breathing
The Generation of Spontaneous Rhythmicity
The Medullary Respiratory Center
The Pontine Respiratory Groups
Spinal Pathways
Reflex Mechanisms of Respiratory Control
Influences of Higher Centers
The Response to Carbon Dioxide
The Response to Hydrogen Ions
The Response to Hypoxia
The Response to Exercise
Altered Breathing Patterns
Key Concepts
Clinical Problems
Suggested Readings
Chapter 10 Nonrespiratory Functions of the Lung
Pulmonary Defense Mechanisms
Nonrespiratory Functions of the Pulmonary Circulation
Metabolic Functions of the Lung
Key Concepts
Suggested Readings
Chapter 11 The Respiratory System Under Stress
Exercise and the Respiratory System
Altitude and Acclimatization
Diving and the Respiratory System
Sleep and the Respiratory System
Key Concepts
Clinical Problems
Suggested Readings
Clinical Problem Answers
Appendix
I. Symbols Used in Respiratory Physiology
II. The Laws Governing the Behavior of Gases
III. Frequently Used Equations
IV. Table of Normal Respiratory and Circulatory Values
Index
Preface
The tenth edition of Pulmonary Physiology has been thoroughly updated. New figures and tables have
been added and others revised to help students visualize concepts more clearly and learn the material
more effectively. New references and clinical correlations have been added to the end of each chapter. It
has been more than 40 years since Pulmonary Physiology was first published. The book is now older
than most of its readers!
Perhaps the greatest pleasure associated with the first nine editions has been receiving many
favorable comments and helpful suggestions from students, readers, and colleagues, and I have used many
of these suggestions in the preparation of this new edition.
This book is intended to be used both as an introductory text for beginning students and as a review
for residents and fellows in such fields as internal medicine, anesthesiology, pediatrics, and pulmonary
medicine. Students who have studied the text should be able to review for licensure and certification
examinations by concentrating on the figures, key concepts, tables, and appendices, which summarize
nearly all of the material in the book.
My goal in writing this book is to provide first-year medical students (as well as respiratory therapy,
nursing, nurse-anesthesia, and other students) with a solid background on the aspects of pulmonary
physiology essential for an understanding of clinical medicine. My approach is to encourage self-
sufficiency not only in studying pulmonary physiology for the first time, but also in understanding the basic
concepts of pulmonary physiology well enough to apply them with confidence to future patients.
I believe that the ways to accomplish this are to inform the reader of the goals of each chapter with
clearly stated learning objectives, to give detailed and complete explanations of physiologic mechanisms
and demonstrate how they apply to pathologic states, and to give the reader a means of self-testing by
providing clinical correlations, problems, and pulmonary function test data to interpret.
The challenge is to write a book that students can read without difficulty in the limited amount of time
allocated to pulmonary physiology in the typical curriculum. The material must be presented in a way that
discourages memorization without real comprehension, because only those students who understand the
basic mechanisms are able to apply them to new situations. The result of this approach should be a book
that covers the essentials of the respiratory system as concisely as possible yet raises no questions in
students’ minds without answering them. I hope that I have achieved these goals in writing this book.
I would like to thank the many people whose comments have helped me revise the various editions of
the book, including my colleagues in respiratory physiology and pulmonary medicine, my colleagues at
LSU Health Sciences Center and Tulane University School of Medicine, and my students and readers. I
give special thanks to Drs. Andy Pellett and Lisa Harrison-Bernard of Louisiana State University Health
Sciences Center and Hershel Raff of the Medical College of Wisconsin. I would also like to thank
everyone who has helped me prepare the manuscripts, illustrations, and proofs, including my many editors
at McGraw Hill, Betsy Giaimo, and my wife Elizabeth. This book is dedicated in memory of Robert S.
Alexander and Jim Miller.
Michael G. Levitzky
1
Function and Structure of the Respiratory System
OBJECTIVES
The reader states the functions of the respiratory system and relates the structural organization of
the system to its functions.
Describes the exchange of oxygen and carbon dioxide with the atmosphere and relates gas
exchange to the metabolism of the tissues of the body.
Defines the role of the respiratory system in acid-base balance.
Lists the nonrespiratory functions of the lungs.
Defines and describes the alveolar-capillary unit, the site of gas exchange in the lungs.
Describes the transport of gas through the conducting airways to and from the alveoli.
Describes the structural characteristics of the airways.
Lists the components of the chest wall and relates the functions of the muscles of respiration to
the movement of air into and out of the alveoli.
Describes the central nervous system initiation of breathing and the innervation of the
respiratory muscles.
The main functions of the respiratory system are to obtain oxygen from the external environment
and supply it to the cells and to remove from the body the carbon dioxide produced by cellular
metabolism.
The respiratory system is composed of the lungs, the conducting airways, the parts of the central
nervous system concerned with the control of the muscles of respiration, and the chest wall. The chest
wall consists of the muscles of respiration—such as the diaphragm, the intercostal muscles, and the
abdominal muscles—and the rib cage.
FUNCTIONS OF THE RESPIRATORY SYSTEM
The functions of the respiratory system include gas exchange, acid-base balance, phonation,
pulmonary defense and metabolism, and the handling of bioactive materials.
