Basic Medical Endocrinology Fourth Edition
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Basic Medical Endocrinology Fourth Edition
i
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
FOR
A
L
LYOUR TEACHING N E textbooks.elsevier.com
E
D
S
ACADEMIC PRESS
To adopt this book for course use, visit http://textbooks.elsevier.com.
Basic Medical Endocrinology
Fourth Edition
H. Maurice Goodman Department of Psychology 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 iii
Cover Credits:
Background: FIGURE 3-2 Histology of the human thyroid. Simple cuboidal cells (arrows) make up the follicles. C _ thyroid colloid (thyroglobulin), which fills 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 fluorescent microscope images of cultured mouse adipocytes that were transfected with a GLUT4-enhanced green fluorescent 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 trafficking of the insulin-responsive glucose transporter 4 in adipocytes. Endocr. Revs. 25: 177–204, by permission of The Endocrine Society.)
Blue figure: 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. (Modified 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. The 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 Theobald’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 Department in Oxford, UK: phone: (ϩ44) 1865 843830, fax: (ϩ44) 1865 853333, E-mail: [email protected] You may also complete your request online via the Elsevier homepage (http://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 08 09 10 9 8 7 6 5 4 3 2 1
This volume is dedicated to my children’s children: Dylan, Adam, Rebecca, and Joshua
Contents
List of Figures
xiii
Preface to the Fourth Edition xxvii
Preface to the First Edition xxix
Preface to the Second Edition xxxi
Preface to the Third Edition xxxiii
CHAPTER 1 INTRODUCTION
1
Biosynthesis of Hormones
4
Storage and Secretion
8
Hormones in Blood
8
Hormone Degradation
10
Mechanisms of Hormone
Action
10
SPECIFICITY
10
CHARACTERISTICS OF RECEPTORS
11
HORMONAL ACTIONS MEDIATED
BY INTRACELLULAR
RECEPTORS
12
HORMONAL ACTIONS MEDIATED BY
SURFACE RECEPTORS
14
The G-protein Coupled Receptors
14
Receptors that Signal through
Tyrosine Kinase
19
REGULATION OF HORMONE
SECRETION
21
Negative Feedback
21
Positive Feedback
23
Feed Forward
23
MEASUREMENT OF HORMONES
23
Immunoassays
24
Hormone Levels in Blood
26
Suggested Reading
26
CHAPTER 2 PITUITARY GLAND 29
Morphology
29
Physiology of the Anterior
Pituitary Gland
31
GLYCOPROTEIN HORMONES
31
GROWTH HORMONE AND
PROLACTIN
33
ADRENOCORTICOTROPIN FAMILY
33
DEVELOPMENT OF THE ANTERIOR
PITUITARY GLAND
34
REGULATION OF ANTERIOR
PITUITARY FUNCTION
35
HYPOPHYSIOTROPIC HORMONES
35
Thyrotropin-releasing hormone (TRH) 36
Gonadotropin Releasing Hormone
37
Control of GH Secretion
37
Corticotropin Releasing Hormone
37
Dopamine and control of Prolactin
Secretion
38
Secretion and Actions of
Hypophysiotropic Hormones
38
FEEDBACK CONTROL OF ANTERIOR
PITUITARY FUNCTION
38
Physiology of the Posterior
Pituitary
39
REGULATION OF POSTERIOR
PITUITARY FUNCTION
40
Suggested Reading
41
CHAPTER 3 THYROID GLAND 43
Morphology
43
vii
Contents
Thyroid Hormones
BIOSYNTHESIS Iodine Trapping Thyroglobulin Synthesis Incorporation of Iodine HORMONE STORAGE SECRETION
Control of Thyroid Function
EFFECTS OF THYROID-STIMULATING HORMONE
EFFECTS OF THE THYROID-STIMULATING IMMUNOGLOBULINS
AUTOREGULATION OF THYROID HORMONE SYNTHESIS
Thyroid Hormones in Blood Metabolism of Thyroid Hormones Physiological Effects of Thyroid
Hormones
GROWTH AND MATURATION Skeletal System Central Nervous System AUTONOMIC NERVOUS SYSTEM CARDIOVASCULAR SYSTEM METABOLISM Oxidative Metabolism and Thermogenesis Carbohydrate Metaboliism Lipid Metabolism Nitrogen Metabolism
Regulation of Thyroid Hormone Secretion
Mechanism of Thyroid Hormone Action
Suggested Reading
44 Glucocorticoids and Responses to Injury
77
45 Adrenal Cortical Function During Stress
81
45 Mechanism of Action of Glucocorticoids
82
45 Regulation of Glucocorticoid Secretion
82
46 Adrenal Medulla
86
46 BIOSYNTHESIS OF MEDULLARY
46
CATECHOLAMINES
86
48 STORAGE, RELEASE, AND METABOLISM
OF MEDULLARY HORMONES
87
48 PHYSIOLOGICAL ACTIONS OF MEDULLARY
HORMONES
88
49 REGULATION OF ADRENAL MEDULLARY
FUNCTION
89
49 Suggested Reading
89
49
50
CHAPTER 5 PRINCIPLES OF HORMONAL 5522 INTEGRATION 91
52
52 Integration of Hormonal Signals at the
52 Cellular and Molecular Level
91
53 AUGMENTATION, ANTAGONISM,
53
AND SYNERGY
91
53 PERMISSIVENESS
93
55 MAINTAINING SIGNAL FIDELITY
93
55 Modulation of Responding Systems
93
56 SENSITIVITY AND CAPACITY
94
SPARE RECEPTORS
96
56 A Hormonal Integration at the
Whole Animal Level
97
57 REDUNDANCY
97
59 REINFORCEMENT
98
PUSH–PULL MECHANISMS
99
CHAPTER 4 ADRENAL GLANDS
Morphology Adrenal Cortex
ADRENOCORTICAL HORMONES Control of Adrenal Cortical Hormone Synthesis Adrenal Steroid Hormones in Blood Postsecretory Metabolism of Adrenal Cortical
Hormones
Physiology of the Mineralocorticoids
Effects of Aldosterone on the Kidney Regulation of Aldosterone Secretion PHYSIOLOGY OF THE GLUCOCORTICOIDS Effects on Energy Metabolism Effects on Water Balance Effects on Lung Development
viii
61 CHAPTER 6 HORMONES OF THE
61 GASTROINTESTINAL TRACT 101
62
62 Functional Anatomy of the GI Tract
101
66 Overview of Digestion and Absorption 104
69 Hormones of the Gastrointestinal Tract 104
THE GASTRIN/CHOLECYSTOKININ FAMILY
105
69 Physiological Actions of Gastrin
106
71 Regulation of Gastrin Secretion
108
71 Cholecystokinin (CCK)
109
74 THE SECRETIN/GLUCAGON SUPERFAMILY
112
75 Secretin
114
75 GIP (Glucose-dependent Insulinotropic Polypeptide/
76
Gastric Inhibitory Peptide)
116
76 Glucagon-like Peptides 1 and 2 (GLP-1 and GLP-2) 118
The Pancreatic Polypeptide Family
121
Neurotensin
122
The Motilin/Ghrelin Family
122
Concluding Comments
124
Suggested Reading
127
CHAPTER 7 THE PANCREATIC ISLETS 129
Morphology of the Endocrine Pancreas Glucagon
BIOSYNTHESIS, SECRETION, AND METABOLISM
PHYSIOLOGICAL ACTIONS OF GLUCAGON Glucose Production Glycogenolysis Gluconeogenesis Lipogenesis and Ketogenesis Ureogenesis REGULATION OF GLUCAGON SECRETION
Insulin
BIOSYNTHESIS, SECRETION, AND METABOLISM
PHYSIOLOGICAL ACTIONS OF INSULIN Effects of Insulin Deficiency Effects on Adipose Tissue Effects on Muscle Effects on Liver MECHANISM OF INSULIN ACTION REGULATION OF INSULIN SECRETION Glucose Other Circulating Metabolites Hormonal and Neural Control Cellular Events
Somatostatin Suggested Reading
129 130
130 130 131 131 132 133 133 134
134
134 135 135 137 138 140 143 144 144 145 146 146
149 149
CHAPTER 8 HORMONAL REGULATION OF FUEL METABOLISM
General Features of Energy Metabolism
BODY FUELS Glucose Glycogen Protein Fat Problems Inherent in the Use of Glucose and
Fat as Metabolic Fuels
151
151
151 151 152 152 152
152
Contents
FUEL CONSUMPTION
153
The Glucose Fatty Acid Cycle
153
AMP Activated Kinase (AMPK)
154
Overall Regulation of Blood Glucose
Concentration
155
Short-term Regulation
156
Long-term Regulation
156
Integrated Actions of Metabolic
Hormones
157
ADIPOSE TISSUE
157
MUSCLE
158
LIVER
158
PANCREATIC ISLETS
159
Regulation of Metabolism During
Feeding and Fasting
159
POSTPRANDIAL PERIOD
159
POSTABSORPTIVE PERIOD
160
FASTING
160
Hormonal Interactions During Exercise 162
SHORT-TERM MAXIMAL EFFORT
163
SUSTAINED AEROBIC EXERCISE
163
Long-term Regulation of Fuel Storage 164
ADIPOGENESIS
165
HYPOTHALAMIC CONTROL OF APPETITE AND
FOOD INTAKE
166
Peripheral Input to Hypothalamic
Feeding and Satiety Neurons
168
ADIPOSITY SIGNALS
168
Leptin
168
Insulin as an Adiposity Signal
170
Adipokines
171
Satiety Signals
172
PERSPECTIVE
173
Suggested Reading
174
CHAPTER 9 REGULATION OF SALT AND WATER BALANCE
General Considerations Salt and Water Balance Antidiuretic Hormone (ADH)
ANTIDIURETIC EFFECT EFFECTS ON BLOOD PRESSURE REGULATION OF ADH SECRETION Plasma Osmolality Blood Volume Dysfunctional States
The Renin-Angiotensin-Aldosterone System
ACTIONS OF ANGIOTENSIN II
175
175 176 178
179 180 181 181 181 182
183
184
ix
Contents
Actions on the Adrenal Cortex
184
Actions on the Kidney
185
Cardiovascular Effects
185
Central Nervous System Effects
185
Regulation of the Renin-Angiotensin-
Aldosterone System
186
Atrial Natriuretic Factor (ANF)
186
PHYSIOLOGICAL ACTIONS
188
Cardiovascular Actions
188
Renal Actions
189
Effects on Aldosterone Secretion
189
Other Effects
190
REGULATION OF ANF SECRETION
190
Integrated Compensatory Responses to
Changes in Salt and Water Balance 190
HEMORRHAGE
192
Response of the Renin Angiotensin System
192
Response of the ADH System
193
Response of Aldosterone
193
Response of ANF
193
DEHYDRATION
193
SALT LOADING AND DEPLETION
194
Suggested Reading
195
CHAPTER 10 HORMONAL REGULATION
OF CALCIUM BALANCE
197
General Features of Calcium Balance
DISTRIBUTION OF CALCIUM IN THE BODY CALCIUM BALANCE Intestinal Absorption Bone Kidney Phosphorus Balance
Parathyroid Glands and Parathyroid Hormone (PTH)
BIOSYNTHESIS, STORAGE, AND SECRETION OF PTH
MECHANISMS OF PARATHYROID HORMONE ACTIONS
PHYSIOLOGICAL ACTIONS OF PTH Actions on Bone Actions on Kidney Effects on Intestinal Absorption PTH FRAGMENTS PARATHYROID HORMONE-RELATED
PEPTIDE (PTHRP) REGULATION OF PTH SECRETION
Calcitonin
197
198 198 199 199 201 202
202
203
204 204 204 205 207 207
207 208
209
CELLS OF ORIGIN
209
BIOSYNTHESIS, SECRETION, AND
METABOLISM
209
PHYSIOLOGICAL ACTIONS OF CALCITONIN
210
Actions on Bone
210
Actions on Kidney
210
REGULATION OF SECRETION
210
The Vitamin D-Endocrine System
211
SYNTHESIS AND METABOLISM
211
PHYSIOLOGICAL ACTIONS OF 1,25(OH)2D3
211
Actions on Intestine
212
Actions on Bone
213
Actions on Kidney
213
Actions on Parathyroid Glands
213
REGULATION OF 1,25(OH)2D3 PRODUCTION
214
Calcium Regulation of Plasma Calcium
Concentrations
214
Integrated Actions of Calcitropic
Hormones
215
RESPONSE TO A HYPOCALCEMIC
CHALLENGE
215
RESPONSE TO A HYPERCALCEMIC
CHALLENGE
216
Other Hormones That Influence Calcium
Balance
216
Suggested Reading
217
CHAPTER 11 HORMONAL CONTROL OF GROWTH
Growth Hormone
SYNTHESIS, SECRETION, AND METABOLISM MODE OF ACTION PHYSIOLOGICAL ACTIONS OF GROWTH
HORMONE Effects on Skeletal Growth Effects of GH/IGF-I on Body Composition
Regulation of GH Secretion
Effects of Age Regulators of GH Secretion ACTIONS OF GHRH, SOMATOSTATIN, IGF-I,
AND GHRELIN ON THE SOMATOTROPE
Thyroid Hormones
DEPENDENCE OF GH SYNTHESIS AND SECRETION ON T3
IMPORTANCE OF T3 FOR EXPRESSION OF GH ACTIONS
Insulin Gonadal Hormones
EFFECTS OF ESTROGENS ON EPIPHYSEAL GROWTH PLATES
219
219
221 221
222 222 226
227
228 229
231
231
233
233
233 234
235
x
i
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
FOR
A
L
LYOUR TEACHING N E textbooks.elsevier.com
E
D
S
ACADEMIC PRESS
To adopt this book for course use, visit http://textbooks.elsevier.com.
Basic Medical Endocrinology
Fourth Edition
H. Maurice Goodman Department of Psychology 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 iii
Cover Credits:
Background: FIGURE 3-2 Histology of the human thyroid. Simple cuboidal cells (arrows) make up the follicles. C _ thyroid colloid (thyroglobulin), which fills 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 fluorescent microscope images of cultured mouse adipocytes that were transfected with a GLUT4-enhanced green fluorescent 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 trafficking of the insulin-responsive glucose transporter 4 in adipocytes. Endocr. Revs. 25: 177–204, by permission of The Endocrine Society.)
Blue figure: 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. (Modified 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. The 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 Theobald’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 Department in Oxford, UK: phone: (ϩ44) 1865 843830, fax: (ϩ44) 1865 853333, E-mail: [email protected] You may also complete your request online via the Elsevier homepage (http://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 08 09 10 9 8 7 6 5 4 3 2 1
This volume is dedicated to my children’s children: Dylan, Adam, Rebecca, and Joshua
Contents
List of Figures
xiii
Preface to the Fourth Edition xxvii
Preface to the First Edition xxix
Preface to the Second Edition xxxi
Preface to the Third Edition xxxiii
CHAPTER 1 INTRODUCTION
1
Biosynthesis of Hormones
4
Storage and Secretion
8
Hormones in Blood
8
Hormone Degradation
10
Mechanisms of Hormone
Action
10
SPECIFICITY
10
CHARACTERISTICS OF RECEPTORS
11
HORMONAL ACTIONS MEDIATED
BY INTRACELLULAR
RECEPTORS
12
HORMONAL ACTIONS MEDIATED BY
SURFACE RECEPTORS
14
The G-protein Coupled Receptors
14
Receptors that Signal through
Tyrosine Kinase
19
REGULATION OF HORMONE
SECRETION
21
Negative Feedback
21
Positive Feedback
23
Feed Forward
23
MEASUREMENT OF HORMONES
23
Immunoassays
24
Hormone Levels in Blood
26
Suggested Reading
26
CHAPTER 2 PITUITARY GLAND 29
Morphology
29
Physiology of the Anterior
Pituitary Gland
31
GLYCOPROTEIN HORMONES
31
GROWTH HORMONE AND
PROLACTIN
33
ADRENOCORTICOTROPIN FAMILY
33
DEVELOPMENT OF THE ANTERIOR
PITUITARY GLAND
34
REGULATION OF ANTERIOR
PITUITARY FUNCTION
35
HYPOPHYSIOTROPIC HORMONES
35
Thyrotropin-releasing hormone (TRH) 36
Gonadotropin Releasing Hormone
37
Control of GH Secretion
37
Corticotropin Releasing Hormone
37
Dopamine and control of Prolactin
Secretion
38
Secretion and Actions of
Hypophysiotropic Hormones
38
FEEDBACK CONTROL OF ANTERIOR
PITUITARY FUNCTION
38
Physiology of the Posterior
Pituitary
39
REGULATION OF POSTERIOR
PITUITARY FUNCTION
40
Suggested Reading
41
CHAPTER 3 THYROID GLAND 43
Morphology
43
vii
Contents
Thyroid Hormones
BIOSYNTHESIS Iodine Trapping Thyroglobulin Synthesis Incorporation of Iodine HORMONE STORAGE SECRETION
Control of Thyroid Function
EFFECTS OF THYROID-STIMULATING HORMONE
EFFECTS OF THE THYROID-STIMULATING IMMUNOGLOBULINS
AUTOREGULATION OF THYROID HORMONE SYNTHESIS
Thyroid Hormones in Blood Metabolism of Thyroid Hormones Physiological Effects of Thyroid
Hormones
GROWTH AND MATURATION Skeletal System Central Nervous System AUTONOMIC NERVOUS SYSTEM CARDIOVASCULAR SYSTEM METABOLISM Oxidative Metabolism and Thermogenesis Carbohydrate Metaboliism Lipid Metabolism Nitrogen Metabolism
Regulation of Thyroid Hormone Secretion
Mechanism of Thyroid Hormone Action
Suggested Reading
44 Glucocorticoids and Responses to Injury
77
45 Adrenal Cortical Function During Stress
81
45 Mechanism of Action of Glucocorticoids
82
45 Regulation of Glucocorticoid Secretion
82
46 Adrenal Medulla
86
46 BIOSYNTHESIS OF MEDULLARY
46
CATECHOLAMINES
86
48 STORAGE, RELEASE, AND METABOLISM
OF MEDULLARY HORMONES
87
48 PHYSIOLOGICAL ACTIONS OF MEDULLARY
HORMONES
88
49 REGULATION OF ADRENAL MEDULLARY
FUNCTION
89
49 Suggested Reading
89
49
50
CHAPTER 5 PRINCIPLES OF HORMONAL 5522 INTEGRATION 91
52
52 Integration of Hormonal Signals at the
52 Cellular and Molecular Level
91
53 AUGMENTATION, ANTAGONISM,
53
AND SYNERGY
91
53 PERMISSIVENESS
93
55 MAINTAINING SIGNAL FIDELITY
93
55 Modulation of Responding Systems
93
56 SENSITIVITY AND CAPACITY
94
SPARE RECEPTORS
96
56 A Hormonal Integration at the
Whole Animal Level
97
57 REDUNDANCY
97
59 REINFORCEMENT
98
PUSH–PULL MECHANISMS
99
CHAPTER 4 ADRENAL GLANDS
Morphology Adrenal Cortex
ADRENOCORTICAL HORMONES Control of Adrenal Cortical Hormone Synthesis Adrenal Steroid Hormones in Blood Postsecretory Metabolism of Adrenal Cortical
Hormones
Physiology of the Mineralocorticoids
Effects of Aldosterone on the Kidney Regulation of Aldosterone Secretion PHYSIOLOGY OF THE GLUCOCORTICOIDS Effects on Energy Metabolism Effects on Water Balance Effects on Lung Development
viii
61 CHAPTER 6 HORMONES OF THE
61 GASTROINTESTINAL TRACT 101
62
62 Functional Anatomy of the GI Tract
101
66 Overview of Digestion and Absorption 104
69 Hormones of the Gastrointestinal Tract 104
THE GASTRIN/CHOLECYSTOKININ FAMILY
105
69 Physiological Actions of Gastrin
106
71 Regulation of Gastrin Secretion
108
71 Cholecystokinin (CCK)
109
74 THE SECRETIN/GLUCAGON SUPERFAMILY
112
75 Secretin
114
75 GIP (Glucose-dependent Insulinotropic Polypeptide/
76
Gastric Inhibitory Peptide)
116
76 Glucagon-like Peptides 1 and 2 (GLP-1 and GLP-2) 118
The Pancreatic Polypeptide Family
121
Neurotensin
122
The Motilin/Ghrelin Family
122
Concluding Comments
124
Suggested Reading
127
CHAPTER 7 THE PANCREATIC ISLETS 129
Morphology of the Endocrine Pancreas Glucagon
BIOSYNTHESIS, SECRETION, AND METABOLISM
PHYSIOLOGICAL ACTIONS OF GLUCAGON Glucose Production Glycogenolysis Gluconeogenesis Lipogenesis and Ketogenesis Ureogenesis REGULATION OF GLUCAGON SECRETION
Insulin
BIOSYNTHESIS, SECRETION, AND METABOLISM
PHYSIOLOGICAL ACTIONS OF INSULIN Effects of Insulin Deficiency Effects on Adipose Tissue Effects on Muscle Effects on Liver MECHANISM OF INSULIN ACTION REGULATION OF INSULIN SECRETION Glucose Other Circulating Metabolites Hormonal and Neural Control Cellular Events
Somatostatin Suggested Reading
129 130
130 130 131 131 132 133 133 134
134
134 135 135 137 138 140 143 144 144 145 146 146
149 149
CHAPTER 8 HORMONAL REGULATION OF FUEL METABOLISM
General Features of Energy Metabolism
BODY FUELS Glucose Glycogen Protein Fat Problems Inherent in the Use of Glucose and
Fat as Metabolic Fuels
151
151
151 151 152 152 152
152
Contents
FUEL CONSUMPTION
153
The Glucose Fatty Acid Cycle
153
AMP Activated Kinase (AMPK)
154
Overall Regulation of Blood Glucose
Concentration
155
Short-term Regulation
156
Long-term Regulation
156
Integrated Actions of Metabolic
Hormones
157
ADIPOSE TISSUE
157
MUSCLE
158
LIVER
158
PANCREATIC ISLETS
159
Regulation of Metabolism During
Feeding and Fasting
159
POSTPRANDIAL PERIOD
159
POSTABSORPTIVE PERIOD
160
FASTING
160
Hormonal Interactions During Exercise 162
SHORT-TERM MAXIMAL EFFORT
163
SUSTAINED AEROBIC EXERCISE
163
Long-term Regulation of Fuel Storage 164
ADIPOGENESIS
165
HYPOTHALAMIC CONTROL OF APPETITE AND
FOOD INTAKE
166
Peripheral Input to Hypothalamic
Feeding and Satiety Neurons
168
ADIPOSITY SIGNALS
168
Leptin
168
Insulin as an Adiposity Signal
170
Adipokines
171
Satiety Signals
172
PERSPECTIVE
173
Suggested Reading
174
CHAPTER 9 REGULATION OF SALT AND WATER BALANCE
General Considerations Salt and Water Balance Antidiuretic Hormone (ADH)
ANTIDIURETIC EFFECT EFFECTS ON BLOOD PRESSURE REGULATION OF ADH SECRETION Plasma Osmolality Blood Volume Dysfunctional States
The Renin-Angiotensin-Aldosterone System
ACTIONS OF ANGIOTENSIN II
175
175 176 178
179 180 181 181 181 182
183
184
ix
Contents
Actions on the Adrenal Cortex
184
Actions on the Kidney
185
Cardiovascular Effects
185
Central Nervous System Effects
185
Regulation of the Renin-Angiotensin-
Aldosterone System
186
Atrial Natriuretic Factor (ANF)
186
PHYSIOLOGICAL ACTIONS
188
Cardiovascular Actions
188
Renal Actions
189
Effects on Aldosterone Secretion
189
Other Effects
190
REGULATION OF ANF SECRETION
190
Integrated Compensatory Responses to
Changes in Salt and Water Balance 190
HEMORRHAGE
192
Response of the Renin Angiotensin System
192
Response of the ADH System
193
Response of Aldosterone
193
Response of ANF
193
DEHYDRATION
193
SALT LOADING AND DEPLETION
194
Suggested Reading
195
CHAPTER 10 HORMONAL REGULATION
OF CALCIUM BALANCE
197
General Features of Calcium Balance
DISTRIBUTION OF CALCIUM IN THE BODY CALCIUM BALANCE Intestinal Absorption Bone Kidney Phosphorus Balance
Parathyroid Glands and Parathyroid Hormone (PTH)
BIOSYNTHESIS, STORAGE, AND SECRETION OF PTH
MECHANISMS OF PARATHYROID HORMONE ACTIONS
PHYSIOLOGICAL ACTIONS OF PTH Actions on Bone Actions on Kidney Effects on Intestinal Absorption PTH FRAGMENTS PARATHYROID HORMONE-RELATED
PEPTIDE (PTHRP) REGULATION OF PTH SECRETION
Calcitonin
197
198 198 199 199 201 202
202
203
204 204 204 205 207 207
207 208
209
CELLS OF ORIGIN
209
BIOSYNTHESIS, SECRETION, AND
METABOLISM
209
PHYSIOLOGICAL ACTIONS OF CALCITONIN
210
Actions on Bone
210
Actions on Kidney
210
REGULATION OF SECRETION
210
The Vitamin D-Endocrine System
211
SYNTHESIS AND METABOLISM
211
PHYSIOLOGICAL ACTIONS OF 1,25(OH)2D3
211
Actions on Intestine
212
Actions on Bone
213
Actions on Kidney
213
Actions on Parathyroid Glands
213
REGULATION OF 1,25(OH)2D3 PRODUCTION
214
Calcium Regulation of Plasma Calcium
Concentrations
214
Integrated Actions of Calcitropic
Hormones
215
RESPONSE TO A HYPOCALCEMIC
CHALLENGE
215
RESPONSE TO A HYPERCALCEMIC
CHALLENGE
216
Other Hormones That Influence Calcium
Balance
216
Suggested Reading
217
CHAPTER 11 HORMONAL CONTROL OF GROWTH
Growth Hormone
SYNTHESIS, SECRETION, AND METABOLISM MODE OF ACTION PHYSIOLOGICAL ACTIONS OF GROWTH
HORMONE Effects on Skeletal Growth Effects of GH/IGF-I on Body Composition
Regulation of GH Secretion
Effects of Age Regulators of GH Secretion ACTIONS OF GHRH, SOMATOSTATIN, IGF-I,
AND GHRELIN ON THE SOMATOTROPE
Thyroid Hormones
DEPENDENCE OF GH SYNTHESIS AND SECRETION ON T3
IMPORTANCE OF T3 FOR EXPRESSION OF GH ACTIONS
Insulin Gonadal Hormones
EFFECTS OF ESTROGENS ON EPIPHYSEAL GROWTH PLATES
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