Basic Medical Endocrinology Fourth Edition ii Companion Web Site: http://books.elsevier.com/companions/9780123739759 Basic Medical Endocrinology, Fourth Edition by H. Maurice Goodman Resources for Professors: • All figures from the book available as PowerPoint slides • Links to web sites carefully chosen to supplement the content of the textbook T O O L S ALL TEACHING NE EDS FOR YOUR textbooks.elsevier.com ACADEMIC PRESS To adopt this book for course use, visit http://textbooks.elsevier.com. Basic Medical Endocrinology Fourth Edition H. Maurice Goodman Department of Physiology University of Massachusetts Medical School AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Cover Credits: Background: FIGURE 3-2 Histology of the human thyroid. Simple cuboidal cells (arrows) make up the follicles. C _ thyroid colloid (thyroglobulin), which fi lls the follicles. (From Borysenko, M. and Beringer, T. (1979) Functional Histology, 312. Little, Brown, Boston by permission of Lippincott, Williams and Wilkins, Philadelphia.) Black/green: FIGURE 7-11 Confocal fl uorescent microscope images of cultured mouse adipocytes that were transfected with a GLUT4-enhanced green fl uorescent protein fusion construct and then incubated in the absence (A) or presence (B) of insulin for 30 min. Insulin stimulation results in the translocation of GLUT4 from intracellular storage sites to the plasma membrane. (From Watson, R.T., Kanzaki, M., and Pessin, J. (2004) Regulated membrane traffi cking of the insulin-responsive glucose transporter 4 in adipocytes. Endocr. Revs. 25: 177–204, by permission of Th e Endocrine Society.) Blue fi gure: FIGURE 10-14 Low-power photomicrograph of a portion of the thyroid gland of a normal dog. Parafollicular (C) cells are indicated in the walls of the follicles. (From Ham, A.W. and Cormack, D. H. (1979) Histology, 8th Edition, 802, by permission of Lippincott, Williams and Wilkins, Philadelphia.) Red, white and blue: FIGURE 11-3 Schematic representation of the tibial epiphyseal growth plate. (Modifi ed from Nilsson, O., Marino, R., De Luca, F., Phillip, M., and Baron, J. (2005) Endocrine regulation of the growth plate. Hormone Research 64: 157–165 by permission of S. Karger AG, Basel.) Pink, yellow, white: FIGURE 12-1 Histological section of human testis. Th e transected tubules show various stages of spermatogenesis. (From di Fiore, M.S.H. (1981) Atlas of Human Histology, 5th Edition, 209. Lea & Febiger, by permission of Lippincott, Williams and Wilkins, Philadelphia.) Academic Press is an imprint of Elsevier 30 Corporate Drive, Suite 400, Burlington, MA 01803, USA 525 B Street, Suite 1900, San Diego, California 92101-4495, USA 84 Th eobald’s Road, London WC1X 8RR, UK Copyright © 2009, Elsevier Inc. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permissions may be sought directly from Elsevier’s Science & Technology Rights (cid:2) (cid:2) Department in Oxford, UK: phone: ( 44) 1865 843830, fax: ( 44) 1865 853333, E-mail: [email protected]. You may also complete your request online via the Elsevier homepage (h ttp://elsevier.com ), by selecting “ Support & Contact” then “ Copyright and Permission ” and then “ Obtaining Permissions. ” Library of Congress Cataloging-in-Publication Data Application Submitted British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library. ISBN: 978-0-12-373975-9 For information on all Academic Press publications visit our Web site at www.elsevierdirect.com Printed in China 09 10 9 8 7 6 5 4 3 2 Th is volume is dedicated to my children’s children: Dylan, Adam, Rebecca, and Joshua This page intentionally left blank Contents List of Figures xiii CHAPTER 2 PITUITARY GLAND 29 Preface to the Fourth Edition xxvii Preface to the First Edition xxix Morphology 29 Preface to the Second Edition xxxi Physiology of the Anterior Preface to the Third Edition xxxiii Pituitary Gland 31 G LYCOPROTEIN HORMONES 31 G ROWTH HORMONE AND CHAPTER 1 INTRODUCTION 1 PROLACTIN 33 ADRENOCORTICOTROPIN FAMILY 33 DEVELOPMENT OF THE ANTERIOR Biosynthesis of Hormones 4 PITUITARY GLAND 34 Storage and Secretion 8 REGULATION OF ANTERIOR Hormones in Blood 8 PITUITARY FUNCTION 35 Hormone Degradation 10 H YPOPHYSIOTROPIC HORMONES 35 Mechanisms of Hormone Th yrotropin Releasing Hormone (TRH) 36 Action 10 Gonadotropin Releasing Hormone 37 SPECIFICITY 10 Control of GH Secretion 37 CHARACTERISTICS OF RECEPTORS 11 Corticotropin Releasing Hormone 37 HORMONAL ACTIONS MEDIATED Dopamine and control of Prolactin BY INTRACELLULAR Secretion 38 RECEPTORS 12 Secretion and Actions of HORMONAL ACTIONS MEDIATED BY Hypophysiotropic Hormones 38 SURFACE RECEPTORS 14 FEEDBACK CONTROL OF ANTERIOR Th e G-protein Coupled Receptors 14 PITUITARY FUNCTION 38 Receptors that Signal through Physiology of the Posterior Tyrosine Kinase 19 Pituitary 39 REGULATION OF HORMONE REGULATION OF POSTERIOR SECRETION 21 PITUITARY FUNCTION 40 Negative Feedback 21 Suggested Reading 41 Positive Feedback 23 Feed Forward 23 M EASUREMENT OF HORMONES 23 CHAPTER 3 THYROID GLAND 43 Immunoassays 23 Hormone Levels in Blood 26 Suggested Reading 26 Morphology 43 vviiii Contents Thyroid Hormones 44 Glucocorticoids and Responses to Injury 77 BIOSYNTHESIS 45 Adrenocortical Function During Stress 81 Iodine Trapping 45 Mechanism of Action of Glucocorticoids 82 Th yroglobulin Synthesis 45 Regulation of Glucocorticoid Secretion 82 Incorporation of Iodine 46 Adrenal Medulla 86 HORMONE STORAGE 46 BIOSYNTHESIS OF THE MEDULLARY SECRETION 46 CATECHOLAMINES 86 Control of Thyroid Function 48 STORAGE, RELEASE, AND METABOLISM EFFECTS OF THYROID-STIMULATING OF MEDULLARY HORMONES 87 HORMONE 48 PHYSIOLOGICAL ACTIONS OF MEDULLARY EFFECTS OF THE THYROID-STIMULATING HORMONES 88 IMMUNOGLOBULINS 49 R EGULATION OF ADRENAL MEDULLARY AUTOREGULATION OF THYROID HORMONE FUNCTION 89 SYNTHESIS 49 Suggested Reading 89 Thyroid Hormones in Blood 49 Metabolism of Thyroid Hormones 50 Physiological Eff ects of Thyroid C HAPTER 5 PRINCIPLES OF HORMONAL Hormones 52 INTEGRATION 91 GROWTH AND MATURATION 52 Skeletal System 52 Central Nervous System 52 Integration of Hormonal Signals at the AUTONOMIC NERVOUS SYSTEM 52 Cellular and Molecular Level 91 C ARDIOVASCULAR SYSTEM 53 AUGMENTATION, ANTAGONISM, M ETABOLISM 53 AND SYNERGY 91 Oxidative Metabolism and Th ermogenesis 53 PERMISSIVENESS 93 C arbohydrate Metaboliism 55 MAINTAINING SIGNAL FIDELITY 93 Lipid Metabolism 55 Modulation of Responding Systems 93 N itrogen Metabolism 56 SENSITIVITY AND CAPACITY 94 Regulation of Thyroid Hormone SPARE RECEPTORS 96 Secretion 56 Hormonal Integration at the Mechanism of Thyroid Hormone Whole Animal Level 97 Action 57 REDUNDANCY 97 REINFORCEMENT 98 Suggested Reading 59 PUSH–PULL MECHANISMS 99 CHAPTER 4 ADRENAL GLANDS 61 CHAPTER 6 HORMONES OF THE GASTROINTESTINAL TRACT 101 Morphology 61 Adrenal Cortex 62 Functional Anatomy of the GI Tract 101 A DRENOCORTICAL HORMONES 62 Overview of Digestion and Absorption 104 Control of Adrenocortical Hormone Synthesis 66 Hormones of the Gastrointestinal Tract 104 Adrenal Steroid Hormones in Blood 69 Postsecretory Metabolism of Adrenal Cortical THE GASTRIN/CHOLECYSTOKININ FAMILY 105 Hormones 69 Physiological Actions of Gastrin 106 Physiology of the Mineralocorticoids 71 Regulation of Gastrin Secretion 108 Eff ects of Aldosterone on the Kidney 71 Cholecystokinin (CCK) 109 THE SECRETIN/GLUCAGON SUPERFAMILY 112 Regulation of Aldosterone Secretion 74 PHYSIOLOGY OF THE GLUCOCORTICOIDS 75 Secretin 114 Eff ects on Energy Metabolism 75 GIP (Glucose-dependent Insulinotropic Polypeptide/ Eff ects on Water Balance 76 Gastric Inhibitory Peptide) 116 Eff ects on Lung Development 76 Glucagon-like Peptides 1 and 2 (GLP-1 and GLP-2) 118 viii Contents Th e Pancreatic Polypeptide Family 121 F UEL CONSUMPTION 153 N eurotensin 122 Th e Glucose Fatty Acid Cycle 153 Th e Motilin/Ghrelin Family 122 AMP Activated Kinase (AMPK) 154 Concluding Comments 124 Overall Regulation of Blood Glucose Suggested Reading 127 Concentration 155 S hort-term Regulation 156 Long-term Regulation 156 CHAPTER 7 THE PANCREATIC ISLETS 129 Integrated Actions of Metabolic Hormones 157 ADIPOSE ISSUE 157 Morphology of the Endocrine Pancreas 129 M USCLE 159 Glucagon 130 LIVER 159 BIOSYNTHESIS, SECRETION, AND PANCREATIC ISLETS 159 METABOLISM 130 Regulation of Metabolism During P HYSIOLOGICAL ACTIONS OF GLUCAGON 130 Feeding and Fasting 160 Glucose Production 131 POSTPRANDIAL PERIOD 160 Glycogenolysis 131 P OSTABSORPTIVE PERIOD 162 G luconeogenesis 132 F ASTING 162 Lipogenesis and Ketogenesis 133 Hormonal Interactions During Exercise 163 U reogenesis 133 S HORT-TERM MAXIMAL EFFORT 163 R EGULATION OF GLUCAGON SECRETION 134 S USTAINED AEROBIC EXERCISE 163 Insulin 134 Long-term Regulation of Fuel Storage 164 BIOSYNTHESIS, SECRETION, ADIPOGENESIS 165 AND METABOLISM 134 H YPOTHALAMIC CONTROL OF APPETITE AND P HYSIOLOGICAL ACTIONS OF INSULIN 135 FOOD INTAKE 166 Eff ects of Insulin Defi ciency 135 Peripheral Input to Hypothalamic Eff ects on Adipose Tissue 137 Feeding and Satiety Neurons 168 Eff ects on Muscle 138 ADIPOSITY SIGNALS 168 Eff ects on Liver 140 Leptin 168 M ECHANISM OF INSULIN ACTION 143 Insulin as an Adiposity Signal 170 R EGULATION OF INSULIN SECRETION 144 Adipokines 171 Glucose 144 Satiety Signals 172 Other Circulating Metabolites 145 PERSPECTIVE 173 Hormonal and Neural Control 146 Suggested Reading 174 C ellular Events 146 Somatostatin 149 Suggested Reading 149 C HAPTER 9 REGULATION OF SALT AND WATER BALANCE 175 CHAPTER 8 HORMONAL REGULATION General Considerations 175 OF FUEL METABOLISM 151 Salt and Water Balance 176 Antidiuretic Hormone (ADH) 178 General Features of Energy ANTIDIURETIC EFFECT 179 Metabolism 151 EFFECTS ON BLOOD PRESSURE 180 B ODY FUELS 151` R EGULATION OF ADH SECRETION 181 Glucose 151 P lasma Osmolality 181 Glycogen 152 Blood Volume 181 Protein 152 Dysfunctional States 182 The Renin-Angiotensin-Aldosterone Fat 152 Problems Inherent in the Use of Glucose and System 183 Fat as Metabolic Fuels 152 ACTIONS OF ANGIOTENSIN II 184 ix