Standard Atomic Weights

Transcription

1

2 Standard Atomic Weights Based on the assigned relative mass of 12 C = 12. For the sake of completeness, all known elements are included in the list. Several of those more recently discovered are represented only by the unstable isotopes. In each case, the values in parentheses in the atomic weight column are the mass numbers of the most stable isotopes. Atomic Atomic Atomic Atomic Name Symbol No. Weight Valence Name Symbol No. Weight Valence Actinium Ac Mercury Hg ,2 Aluminum Al (hydrargyrum) Americium Am 95 (243) 3,4,5,6 Molybdenum Mo ,4,6 Antimony Sb ,5 Neodymium Nd (stibium) Neon Ne Argon Ar Neptunium Np ,5,6 Arsenic As ,5 Nickel Ni ,3 Astatine At 85 (210) 1,3,5,7 Niobium Nb ,5 Barium Ba (columbium) Berkelium Bk 97 (247) 3,4 Nitrogen N ,5 Beryllium Be Nobelium No 102 (259)... Bismuth Bi ,5 Osmium Os ,3,4,8 Boron B Oxygen O Bromine Br ,3,5,7 Palladium Pd ,4,6 Cadmium Cd Phosphorus P ,5 Calcium Ca Platinum Pt ,4 Californium Cf 98 (251)... Plutonium Pu 94 (244) 3,4,5,6 Carbon C ,4 Polonium Po 84 (209)... Cerium Ce ,4 Potassium K Cesium Cs (kalium) Chlorine Cl ,3,5,7 Praseodymium Pr Chromium Cr ,3,6 Promethium Pm 61 (145) 3 Cobalt Co ,3 Protactinium Pa Columbium Radium Ra (see Niobium) Radon Rn 86 (222) 0 Copper Cu ,2 Rhenium Re Curium Cm 96 (247) 3 Rhodium Rh Dysprosium Dy Rubidium Rb Einsteinium Es 99 (252)... Ruthenium Ru ,4,6,8 Erbium Er Samarium Sm ,3 Europium Eu ,3 Scandium Sc Fermium Fm 100 (257)... Selenium Se ,4,6 Fluorine F Silicon Si Francium Fr 87 (223) 1 Silver Ag Gadolinium Gd (argentum) Gallium Ga ,3 Sodium Na Germanium Ge (natrium) Gold Au ,3 Strontium Sr (aurum) Sulfur S ,4,6 Hafnium Hf Tantalum Ta Helium He Technetium Tc 43 (98) 6,7 Holmium Ho Tellurium Te ,4,6 Hydrogen H Terbium Tb Indium In Thallium Tl ,3 Iodine I ,3,5,7 Thorium Th Iridium Ir ,4 Thulium Tm Iron Fe ,3 Tin Sn ,4 (ferrum) (stannum) Krypton Kr Titanium Ti ,4 Lanthanum La Tungsten W Lawrencium Lr 103 (260)... (wolfram) Lead Pb ,4 Uranium U ,6 (plumbum) Vanadium V ,5 Lithium Li Xenon Xe Lutetium Lu Ytterbium Yb ,3 Magnesium Mg Yttrium Y Manganese Mn ,3,4,6,7 Zinc Zn Mendelevium Md 101 (258)... Zirconium Zr Modified and reproduced, with permission from Lide DR (editor-in-chief): CRC Handbook of Chemistry and Physics, 83rd ed. CRC Press,

3 a LANGE medical book Review of Medical Physiology twenty-second edition William F. Ganong, MD Jack and DeLoris Lange Professor of Physiology Emeritus University of California San Francisco Lange Medical Books/McGraw-Hill Medical Publishing Division New York Chicago San Francisco Lisbon London Madrid Mexico City Milan New Deli San Juan Seoul Singapore Sydney Toronto

4 Review of Medical Physiology, Twenty-Second Edition Copyright 2005 by The McGraw-Hill Companies, Inc. All rights reserved. Printed in the United States of America. Except as permitted under the United States Copyright Act of 1976, no part of this publication may be reproduced or distributed in any form or by any means, or stored in a data base or retrieval system, without the prior written permission of the publisher. Previous editions copyright 2003, 2001 by The McGraw-Hill Companies, Inc.; copyright 1999, 1997, 1995, 1993, 1991, by Appleton & Lange; copyright 1963 through 1989 by Lange Medical Publications DOC/DOC ISBN ISSN Notice Medicine is an ever-changing science. As new research and clinical experience broaden our knowledge, changes in treatment and drug therapy are required. The author and the publisher of this work have checked with sources believed to be reliable in their efforts to provide information that is complete and generally in accord with the standards accepted at the time of publication. However, in view of the possibility of human error or changes in medical sciences, neither the author nor the publisher nor any other party who has been involved in the preparation or publication of this work warrants that the information contained herein is in every respect accurate or complete, and they disclaim all responsibility for any errors or omissions or for the results obtained from use of the information contained in this work. Readers are encouraged to confirm the information contained herein with other sources. For example and in particular, readers are advised to check the product information sheet included in the package of each drug they plan to administer to be certain that the information contained in this work is accurate and that changes have not been made in the recommended dose or in the contraindications for administration. This recommendation is of particular importance in connection with new or infrequently used drugs. The book was set in Adobe Garamond by Rainbow Graphics. The editors were Janet Foltin, Harriet Lebowitz, and Regina Y. Brown. The production supervisor was Catherine H. Saggese. The cover designer was Mary McKeon. The art manager was Charissa Baker. The index was prepared by Katherine Pitcoff. RR Donnelley was printer and binder. This book is printed on acid-free paper.

5 Contents Preface xi SECTION I. INTRODUCTION The General & Cellular Basis of Medical Physiology Introduction 1 Transport Across Cell Membranes 28 General Principles 1 The Capillary Wall 35 Functional Morphology of the Cell 8 Intercellular Communication 36 Structure & Function of Homeostasis 48 DNA & RNA 18 Aging 48 Section I References 49 SECTION II. PHYSIOLOGY OF NERVE & MUSCLE CELLS Excitable Tissue: Nerve Introduction 51 Properties of Mixed Nerves 60 Nerve Cells 51 Nerve Fiber Types & Function 60 Excitation & Conduction 54 Neurotrophins 61 Ionic Basis of Excitation Neuroglia 63 & Conduction Excitable Tissue: Muscle Introduction 65 Cardiac Muscle 78 Skeletal Muscle 65 Morphology 78 Morphology 65 Electrical Properties 78 Electrical Phenomena Mechanical Properties 78 & Ionic Fluxes 68 Metabolism 81 Contractile Responses 68 Pacemaker Tissue 81 Energy Sources & Metabolism 74 Smooth Muscle 82 Properties of Skeletal Muscles Morphology 82 in the Intact Organism 75 Visceral Smooth Muscle 82 Multi-Unit Smooth Muscle Synaptic & Junctional Transmission Introduction 85 Principal Neurotransmitter Systems 94 Synaptic Transmission 85 Synaptic Plasticity & Learning 116 Functional Anatomy 85 Neuromuscular Transmission 116 Electrical Events in Postsynaptic Neuromuscular Junction 116 Neurons 88 Nerve Endings in Smooth & Cardiac Inhibition & Facilitation Muscle 118 at Synapses 91 Denervation Hypersensitivity 119 Chemical Transmission of Synaptic Activity 94 iii

6 iv / CONTENTS 5. Initiation of Impulses in Sense Organs Introduction 121 Generation of Impulses in Different Nerves 123 Sense Organs & Receptors 121 Coding of Sensory Information 124 The Senses 121 Section II References 127 SECTION III. FUNCTIONS OF THE NERVOUS SYSTEM Reflexes Introduction 129 Polysynaptic Reflexes: The Withdrawal Reflex 134 Monosynaptic Reflexes: General Properties of Reflexes 137 The Stretch Reflex Cutaneous, Deep, & Visceral Sensation Introduction 138 Temperature 142 Pathways 138 Pain 142 Touch 141 Other Sensations 147 Proprioception Vision Introduction 148 Responses in the Visual Pathways & Cortex 160 Anatomic Considerations 148 Color Vision 163 The Image-Forming Mechanism 152 Other Aspects of Visual Function 166 The Photoreceptor Mechanism 156 Eye Movements Hearing & Equilibrium Introduction 171 Hearing 176 Anatomic Considerations 171 Vestibular Function 183 Hair Cells Smell & Taste Introduction 185 Taste 188 Smell 185 Receptor Organs & Pathways Alert Behavior, Sleep, & the Electrical Activity of the Brain Introduction 192 Evoked Cortical Potentials 193 The Thalamus & the Cerebral The Electroencephalogram 194 Cortex 192 Physiologic Basis of the EEG, Consciousness, The Reticular Formation & the Reticular & Sleep 196 Activating System Control of Posture & Movement Introduction 202 Spinal Integration 207 General Principles 202 Medullary Components 210 Corticospinal & Corticobulbar Midbrain Components 211 System 203 Cortical Components 212 Anatomy & Function 203 Basal Ganglia 213 Posture-Regulating Systems 206 Cerebellum 217

7 CONTENTS / v 13. The Autonomic Nervous System Introduction 223 Chemical Transmission at Autonomic Anatomic Organization of Autonomic Junctions 223 Outflow 223 Responses of Effector Organs to Autonomic Nerve Impulses Central Regulation of Visceral Function Introduction 232 Relation to Cyclic Phenomena 235 Medulla Oblongata 232 Hunger 235 Hypothalamus 233 Thirst 240 Anatomic Considerations 233 Control of Posterior Pituitary Secretion 242 Hypothalamic Function 234 Control of Anterior Pituitary Secretion 248 Relation to Autonomic Function 234 Temperature Regulation 251 Relation to Sleep Neural Basis of Instinctual Behavior & Emotions Introduction 256 Other Emotions 259 Anatomic Considerations 256 Motivation & Addiction 260 Limbic Functions 256 Brain Chemistry & Behavior 261 Sexual Behavior Higher Functions of the Nervous System : Conditioned Reflexes, Learning, & Related Phenomena Introduction 266 Learning & Memory 266 Methods 266 Functions of the Neocortex 272 Section III References 276 SECTION IV. ENDOCRINOLOGY, METABOLISM, & REPRODUCTIVE FUNCTION Energy Balance, Metabolism, & Nutrition Introduction 279 Protein Metabolism 292 Energy Metabolism 279 Fat Metabolism 298 Intermediary Metabolism 282 Nutrition 311 Carbohydrate Metabolism The Thyroid Gland Introduction 317 Effects of Thyroid Hormones 323 Anatomic Considerations 317 Regulation of Thyroid Secretion 326 Formation & Secretion Clinical Correlates 328 of Thyroid Hormones 317 Transport & Metabolism of Thyroid Hormones Endocrine Functions of the Pancreas & Regulation of Carbohydrate Metabolism Introduction 333 Fate of Secreted Insulin 335 Islet Cell Structure 333 Effects of Insulin 336 Structure, Biosynthesis, & Secretion Mechanism of Action 338 of Insulin 334 Consequences of Insulin Deficiency 340

8 vi / CONTENTS Insulin Excess 344 Effects of Other Hormones & Exercise Regulation of Insulin Secretion 345 on Carbohydrate Metabolism 351 Glucagon 348 Hypoglycemia & Diabetes Mellitus in Humans 353 Other Islet Cell Hormones The Adrenal Medulla & Adrenal Cortex Introduction 356 Physiologic Effects of Adrenal Morphology 356 Glucocorticoids 369 Adrenal Medulla 358 Pharmacologic & Pathologic Effects Structure & Function of Medullary of Glucocorticoids 370 Hormones 358 Regulation of Glucocorticoid Regulation of Adrenal Medullary Secretion 372 Secretion 361 Effects of Mineralocorticoids 375 Adrenal Cortex 361 Regulation of Aldosterone Secretion 377 Structure & Biosynthesis of Role of Mineralocorticoids in the Adrenocortical Hormones 361 Regulation of Salt Balance 380 Transport, Metabolism, & Excretion Summary of the Effects of of Adrenocortical Hormones 366 Adrenocortical Hyper- Effects of Adrenal Androgens & Hypofunction in Humans 380 & Estrogens Hormonal Control of Calcium Metabolism & the Physiology of Bone Introduction 382 The Parathyroid Glands 390 Calcium & Phosphorus Metabolism 382 Calcitonin 393 Bone Physiology 383 Effects of Other Hormones & Humoral Agents on Vitamin D & the Calcium Metabolism 395 Hydroxycholecalciferols The Pituitary Gland Introduction 396 Physiology of Growth 404 Morphology 396 Pituitary Insufficiency 408 Intermediate-Lobe Hormones 397 Pituitary Hyperfunction in Humans 409 Growth Hormone The Gonads: Development & Function of the Reproductive System Introduction 411 Gametogenesis & Ejaculation 424 Sex Differentiation & Development 411 Endocrine Function of the Testes 428 Chromosomal Sex 411 Control of Testicular Function 431 Embryology of the Human Abnormalities of Testicular Function 433 Reproductive System 413 The Female Reproductive System 433 Aberrant Sexual Differentiation 414 The Menstrual Cycle 433 Puberty 418 Ovarian Hormones 438 Precocious & Delayed Puberty 420 Control of Ovarian Function 444 Menopause 421 Abnormalities of Ovarian Function 447 Pituitary Gonadotropins & Prolactin 421 Pregnancy 448 The Male Reproductive System 424 Lactation 451 Structure 424

9 CONTENTS / vii 24. Endocrine Functions of the Kidneys, Heart, & Pineal Gland Introduction 454 Hormones of the Heart & Other Natriuretic The Renin-Angiotensin System 454 Factors 460 Erythropoietin 459 Pineal Gland 462 Section IV References 465 SECTION V. GASTROINTESTINAL FUNCTION Digestion & Absorption Introduction 467 Lipids 473 Carbohydrates 467 Absorption of Water & Electrolytes 475 Proteins & Nucleic Acids 471 Absorption of Vitamins & Minerals Regulation of Gastrointestinal Function Introduction 479 Exocrine Portion of the Pancreas 497 General Considerations 479 Liver & Biliary System 498 Gastrointestinal Hormones 482 Small Intestine 504 Mouth & Esophagus 488 Colon 508 Stomach 491 Section V References 512 SECTION VI. CIRCULATION Circulating Body Fluids Introduction 515 Red Blood Cells 532 Blood 515 Blood Types 537 Bone Marrow 515 Plasma 539 White Blood Cells 516 Hemostasis 540 Immunity 520 Lymph 546 Platelets Origin of the Heartbeat & the Electrical Activity of the Heart Introduction 547 Cardiac Arrhythmias 554 Origin & Spread of Cardiac Electrocardiographic Findings in Other Cardiac Excitation 547 & Systemic Diseases 561 The Electrocardiogram The Heart as a Pump Introduction 565 Cardiac Output 570 Mechanical Events of the Cardiac Cycle Dynamics of Blood & Lymph Flow Introduction 577 Capillary Circulation 590 Functional Morphology 577 Lymphatic Circulation & Interstitial Fluid Biophysical Considerations 581 Volume 593 Arterial & Arteriolar Circulation 587 Venous Circulation Cardiovascular Regulatory Mechanisms Introduction 597 Systemic Regulation by Hormones 600 Local Regulation 597 Systemic Regulation by the Nervous System 602 Substances Secreted by the Endothelium 598

10 viii / CONTENTS 32. Circulation Through Special Regions Introduction 611 Brain Metabolism & Oxygen Cerebral Circulation 611 Requirements 619 Anatomic Considerations 611 Coronary Circulation 620 Cerebrospinal Fluid 612 Splanchnic Circulation 623 The Blood-Brain Barrier 614 Cutaneous Circulation 625 Cerebral Blood Flow & Placental & Fetal Circulation 627 Its Regulation Cardiovascular Homeostasis in Health & Disease Introduction 630 Inflammation & Wound Healing 635 Compensations for Gravitational Shock 636 Effects 630 Hypertension 641 Exercise 632 Heart Failure 643 Section VI References 644 SECTION VII. RESPIRATION Pulmonary Function Introduction 647 Gas Exchange in the Lungs 660 Properties of Gases 647 Pulmonary Circulation 661 Anatomy of the Lungs 649 Other Functions of the Respiratory System 664 Mechanics of Respiration Gas Transport Between the Lungs & the Tissues Introduction 666 Carbon Dioxide Transport 669 Oxygen Transport Regulation of Respiration Introduction 671 Chemical Control of Breathing 672 Neural Control of Breathing 671 Nonchemical Influences on Respiration 678 Regulation of Respiratory Activity Respiratory Adjustments in Health & Disease Introduction 681 Hypercapnia & Hypocapnia 692 Effects of Exercise 681 Other Respiratory Abnormalities 692 Hypoxia 683 Diseases Affecting the Pulmonary Circulation 694 Hypoxic Hypoxia 684 Effects of Increased Barometric Pressure 694 Other Forms of Hypoxia 690 Artificial Respiration 695 Oxygen Treatment 691 Section VII References 697 SECTION VIII. FORMATION & EXCRETION OF URINE Renal Function & Micturition Introduction 699 Tubular Function 708 Functional Anatomy 699 Water Excretion 713 Renal Circulation 702 Acidification of the Urine Glomerular Filtration 705 & Bicarbonate Excretion 720

11 CONTENTS / ix Regulation of Na + & Cl Excretion 723 Effects of Disordered Renal Function 725 Regulation of K + Excretion 724 The Bladder 726 Diuretics Regulation of Extracellular Fluid Composition & Volume Introduction 729 Defense of Specific Ionic Composition 730 Defense of Tonicity 729 Defense of H + Concentration 730 Defense of Volume 729 Section VIII References 738 Self-Study: Objectives, Essay Questions, & Multiple-Choice Questions (black edges) Answers to Quantitative & Multiple-Choice Questions (black edges) Appendix General References 811 Some Standard Respiratory Symbols 821 Normal Values & the Statistical Equivalents of Metric, United States, Evaluation of Data 811 & English Measures 821 Abbreviations & Symbols Commonly Greek Alphabet 822 Used in Physiology 814 Index Standard Atomic Weights Inside Front Cover Ranges of Normal Values in Human Whole Blood, Plasma, or Serum Inside Back Cover

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13 Preface This book is designed to provide a concise summary of mammalian and, particularly, of human physiology that medical students and others can use by itself or can supplement with readings in other texts, monographs, and reviews. Pertinent aspects of general and comparative physiology are also included. Summaries of relevant anatomic considerations will be found in each section, but this book is written primarily for those who have some knowledge of anatomy, chemistry, and biochemistry. Examples from clinical medicine are given where pertinent to illustrate physiologic points. In many of the chapters, physicians desiring to use this book as a review will find short discussions of important symptoms produced by disordered function. Review of Medical Physiology also includes a self-study section to help students review for Board and other examinations and an appendix that contains general references, a discussion of statistical methods, a glossary of abbreviations, acronyms, and symbols commonly used in physiology, and several useful tables. The index is comprehensive and specifically designed for ease in locating important terms, topics, and concepts. In writing this book, the author has not been able to be complete and concise without also being dogmatic. I believe, however, that the conclusions presented without detailed discussion of the experimental data on which they are based are supported by the bulk of the current evidence. Much of this evidence can be found in the papers cited in the credit lines accompanying the illustrations. Further discussions of particular subjects and information on subjects not considered in detail can be found in the references listed at the end of each section. Information about serial review publications that provide up-to-date discussion of various physiologic subjects is included in the note on general references in the appendix. In the interest of brevity and clarity, I have in most instances omitted the names of the many investigators whose work made possible the view of physiology presented here. This omission is in no way intended to slight their contributions, but including their names and specific references to original papers would greatly increase the length of the book. In this twenty-second edition, as in previous editions, the entire book has been revised, with a view to eliminating errors, incorporating suggestions of readers, updating concepts, and discarding material that is no longer relevant. In this way, the book has been kept concise while remaining as up-to-date and accurate as possible. Since the last edition, research on the regulation of food intake has continued at a rapid pace, and this topic has been expanded in the current edition. So has consideration of mitochondria and molecular motors, with emphasis on the ubiquity of the latter. Chapter 38 on renal function has been reorganized as well as updated. The section on estrogen receptors has been revised in terms of the complexity of the receptor and the way this relates to tailor-made estrogens used in the treatment of disease. Other topics on which there is new information include melanopsin, pheromones related to lactation, von Willebrand factor, and the complexity of connexons. The self-study section has been updated, with emphasis placed on physiology in relation to disease, in keeping with the current trend in the United States Medical Licensing Examinations (USMLE). I am greatly indebted to the many individuals who helped with the preparation of this book. Those who were especially helpful in the preparation of the twenty-second edition include Drs. Stephen McPhee, Dan Stites, David Gardner, Igor Mitrovic, Michael Jobin, Krishna Rao, and Johannes Werzowa. Andrea Chase provided invaluable secretarial assistance, and, as always, my wife made important contributions.special thanks are due to Jim Ransom, who edited the first edition of this book over 42 years ago and now has come back to make helpful and worthwhile comments on the two most recent editions. Many associates and friends provided unpublished illustrative materials, and numerous authors and publishers generously granted permission to reproduce illustrations from other books and journals. I also thank all the students and others who took the time to write to me offering helpful criticisms and suggestions. Such comments are always welcome, and I solicit additional corrections and criticisms, which may be addressed to me at Department of Physiology University of California San Francisco, CA USA Since this book was first published in 1963, the following translations have been published: Bulgarian, Chinese (2 independent translations), Czech (2 editions), French (2 independent translations), German (4 editions), Greek (2 editions), Hungarian, Indonesian (4 editions), Italian (9 editions), Japanese (17 editions), Korean, Malaysian, xi

14 xii / PREFACE Polish (2 editions), Portuguese (7 editions), Serbo-Croatian, Spanish (19 editions), Turkish (2 editions), and Ukranian. Various foreign English language editions have been published, and the book has been recorded in English on tape for the blind. The tape recording is available from Recording for the Blind, Inc., 20 Rozsel Road, Princeton, NJ USA. For computer users, the book is now available, along with several other titles in the Lange Medical Books series, in STAT!-Ref, a searchable Electronic Medical Library ( from Teton Data Systems, P.O. Box 4798 Jackson, WY USA. More information about this and other Lange and McGraw-Hill books, including addresses of the publisher s international offices, is available on McGraw-Hill s web site, San Francisco March 2005 William F. Ganong, MD

15 SECTION I Introduction The General & Cellular Basis of Medical Physiology 1 INTRODUCTION In unicellular organisms, all vital processes occur in a single cell. As the evolution of multicellular organisms has progressed, various cell groups have taken over particular functions. In humans and other vertebrate animals, the specialized cell groups include a gastrointestinal system to digest and absorb food; a respiratory system to take up O 2 and eliminate CO 2 ; a urinary system to remove wastes; a cardiovascular system to distribute food, O 2, and the products of metabolism; a reproductive system to perpetuate the species; and nervous and endocrine systems to coordinate and integrate the functions of the other systems. This book is concerned with the way these systems function and the way each contributes to the functions of the body as a whole. This chapter presents general concepts and principles that are basic to the function of all the systems. It also includes a short review of fundamental aspects of cell physiology. Additional aspects of cellular and molecular biology are considered in the relevant chapters on the various organs. GENERAL PRINCIPLES Organization of the Body The cells that make up the bodies of all but the simplest multicellular animals, both aquatic and terrestrial, exist in an internal sea of extracellular fluid (ECF) enclosed within the integument of the animal. From this fluid, the cells take up O 2 and nutrients; into it, they discharge metabolic waste products. The ECF is more dilute than present-day seawater, but its composition 1 closely resembles that of the primordial oceans in which, presumably, all life originated. In animals with a closed vascular system, the ECF is divided into two components: the interstitial fluid and the circulating blood plasma. The plasma and the cellular elements of the blood, principally red blood cells, fill the vascular system, and together they constitute the total blood volume. The interstitial fluid is that part of the ECF that is outside the vascular system, bathing the cells. The special fluids lumped together as transcellular fluids are discussed below. About a third of the total body water (TBW) is extracellular; the remaining two thirds is intracellular (intracellular fluid). Body Composition In the average young adult male, 18% of the body weight is protein and related substances, 7% is mineral, and 15% is fat. The remaining 60% is water. The distribution of this water is shown in Figure 1 1. The intracellular component of the body water accounts for about 40% of body weight and the extracellular component for about 20%. Approximately 25% of the extracellular component is in the vascular system (plasma = 5% of body weight) and 75% outside the blood vessels (interstitial fluid = 15% of body weight). The total blood volume is about 8% of body weight. Measurement of Body Fluid Volumes It is theoretically possible to measure the size of each of the body fluid compartments by injecting substances that will stay in only one compartment and then calculating the volume of fluid in which the test substance is