Contributors HUGO A. ARMELIN GERALD LITWACK BARBARA BOSS G. STANLEY MCKNIGHT MAX H. CAKE LEON L. MILLER CARLO M. CROCE BERT W. O'MALLEY P. CUATRECASAS JACK H. OPPENHEIMER JACK GORSKI JEFFREY M. ROSEN GEOFFREY GRANT GORDON H. SATO EDMOND E. GRIFFIN ROBERT T. SCHIMKE M. D. HOLLENBERG DAVID J. SHAPIRO BENITA S. KATZENELLENBOGEN MARTIN I. SURKS WYLIE VALE Biochemical Actions of Hormones Edited by GERALD LITWACK Fels Research Institute and Department of Biochemistry Temple University, School of Medicine Philadelphia, Pennsylvania VOLUME III ACADEMIC PRESS New York San Francisco London 1975 A Subsidiary of Harcourt Brace Jovanovich, Publishers COPYRIGHT © 1975, BY ACADEMIC PRESS, 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. ACADEMIC PRESS, INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 Library of Congress Cataloging in Publication Data Main entry under title: Biochemical actions of hormones. Includes bibliographies. 1. Hormones. I. Litwack, Gerald, ed. II. Axelrod, Julius, (date) [DNLM: 1. Hormones. 2. Physiology. WK102B615] QP571.B56 574.Γ927 70-107567 ISBN 0-12-452803-1 (v.3) PRINTED IN THE UNITED STATES OF AMERICA To the memory of three pioneers in the field of hormone action: Leonard D. Garren, Asher Körner, and Earl W. Sutherland List of Contributors Numbers in parentheses indicate the pages on which the authors' contributions begin. Hugo A. Armelin (1), Departmento de Bioquimica, Instituto de Quimica, Universidade de Säo Paulo, Cidade Universitaria, Sao Paulo, Brazil Barbara Boss (87), The Salk Institute, San Diego, California Max H. Cake (317), Fels Research Institute, Temple University School of Medicine, Philadelphia, Pennsylvania Carlo M. Croce (23), The Wistar Institute of Anatomy and Biology, Philadelphia, Pennsylvania P. Cuatrecasas (41), Department of Pharmacology and Experimental Therapeutics and Department of Medicine, The Johns Hopkins University School of Medicine, Baltimore, Maryland Jack Gorski (187), Departments of Biochemistry and Animal Science, University of Wisconsin, Madison, Wisconsin Geoffrey Grant (87), The Salk Institute, San Diego, California Edmond E. Griffin (159), Department of Radiation Biology and Bio physics and Department of Biochemistry, University of Roches ter, School of Medicine and Dentistry, Rochester, New York M. D. Hollenberg (41), Department of Pharmacology and Experimental Therapeutics and Department of Medicine, The Johns Hopkins University School of Medicine, Baltimore, Maryland Benita S. Katzenellenbogen (187), Department of Physiology and Bio physics, University of Illinois, and School of Basic Medical Sci ences, University of Illinois College of Medicine, Urbana, Illinois XI xii List of Contributors Gerald Litwack (23, 317), Fels Research Institute and Department of Biochemistry, Temple University School of Medicine, Phila delphia, Pennsylvania G. Stanley McKnight* (245), Department of Biological Sciences, Stan ford University, Stanford, California Leon L. Miller (159), Department of Radiation Biology and Biophysics and Department of Biochemistry, University of Rochester, School of Medicine and Dentistry, Rochester, New York Bert W. O'Malley (271), Department of Cell Biology, Baylor College of Medicine, Houston, Texas Jack H. Oppenheimer (119), Division of Endocrinology, Montefiore Hospital and Medical Center, and Department of Medicine, Albert Einstein College of Medicine, Bronx, New York Jeffrey M. Rosen (271), Department of Cell Biology, Baylor College of Medicine, Houston, Texas Gordon H. Sato (391), Department of Biology, University of Cali fornia at San Diego, La Jolla, California Robert T. Schimke (245), Department of Biological Sciences, Stan ford University, Stanford, California David J. Shapiro (245), Department of Biological Sciences, Stanford University, Stanford, California Martin I. Surks (119), Endocrine Research Laboratory, Montefiore Hospital and Medical Center, and Department of Medicine, Al bert Einstein College of Medicine, Bronx, New York Wylie Vale (87), The Salk Institute, San Diego, California * Present address: Faculte de Medecine, Institut de Chimie Biologique, Strasbourg, France. Preface Research directed toward an understanding of the mechanisms of hormone action has been concentrated at the biochemical and molec ular levels using individual hormones in mature or developing cells. Volumes I and II of "Biochemical Actions of Hormones" summarized and evaluated the results of this research, covering virtually every area of the field in which significant advances were made through 1971. Volume III evolved from the realization of further advances in established areas and new approaches not previously developed. A wealth of new information on steroid hormone action is summa rized in four chapters. Although a certain amount of overlapping in formation on the mode of action of estrogens could not be avoided, it comes from some of the foremost laboratories involved with this problem, and, if anything, will make the description of current work on this hormone all the more complete. The recent and previously unreviewed areas of action of hypothalamic-regulating hormones and plasma membrane receptors are presented. These new areas promise to have wide repercussions on endocrine research. Three contribu tions deal with the approaches using genetics and cell culture to ad vance our knowledge of hormone action and of systems by which previously undescribed hormones are being discovered. Two other chapters summarize recent advances in the mode of action of thyroid hormone and of hormones acting on the synthesis of proteins in liver perfusion systems. Originally, only a two-volume treatise had been planned. How ever, as mentioned above, new advances signaled the need for a third volume. Additional volumes will be published if new knowledge warrants them. I wish to commend the staff of Academic Press for their excellent cooperation in the publication of these volumes. GERALD LITWACK xiii Contents of Previous Volumes Volume I Hormonal Responses in Amphibian Metamorphosis Earl Frieden and John J. Just The Developmental Formation of Enzymes in Rat Liver Olga Greengard Regulation of Protein Synthesis by Growth and Developmental Hormones /. R. Tata The Regulation of Some Biochemical Circadian Rhythms Ira B. Black and Julius Axelrod Hormones and Transport across Cell Membranes Thomas R. Riggs Binding of Hormones to Serum Proteins Ulrich Westphal Insulin and Protein Synthesis K. L. Manchester Mineral ocorticoids Isidore S. Edelman and Darrell D. Fanestil Parathyroid Hormone and Calcitonin Howard Rasmussen and Alan Tenenhouse Mechanism of Action of Thyrotropin E. Schell-Frederick andj. E. Dumont The Thymus as an Endocrine Gland: Hormones and Their Actions Allan L. Goldstein and Abraham White xv χνι Contents of Previous Volumes Plant Hormones Carlos O. Miller AUTHOR INDEX—SUBJECT INDEX Volume II The Present Status of Genetic Regulation by Hormones Gordon M. Tomkins and Thomas D. Gelehrter Cyclic AMP and Hormone Action R. W. Butcher, G. A. Robinson, and E. W. Sutherland Multiple Hormonal Interactions. The Mammary Gland Roger W. Turkington The Catecholamines G. A. Robison, R. W. Butcher, and E. W. Sutherland Subcellular Actions of Glucocorticoids Gerald Litwack and Sanford Singer Insulin Actions on Carbohydrate and Lipid Metabolism Irving B. Fritz Estrogens and Progestins Elwood V. Jensen and Eugene R. DeSombre Androgenic Regulation of Tissue Growth and Function H. G. Williams-Ashman and A. H. Reddi Mechanism of Action of Gonadotropins and Prolactin Ralph I. Dorfman The Mechanism of Action of Adrenocorticotropic Hormone James J. Ferguson, Jr. 25-Hydroxycholecalciferol: A Hormonal Form of Vitamin D Hector F. DeLuca and Mark J. Melancon, Jr. Insect Hormones G. R. Wyatt AUTHOR INDEX—SUBJECT INDEX CHAPTER 1 Hormones and Regulation of Cell Division: Mammalian Cell Cultures as an Experimental Approach Hugo A. Armelin I. Introduction 1 II. Establishment of "Growth Responsive" Cell Lines: Its Empiri cism and Pitfalls 2 III. Control of Cell Proliferation and "Cell Cycle": Prospective Models 6 IV. Growth Factors in Mammalian Cell Cultures: Experimental Facts and Tentative Concepts 12 V. Fibroblast Cell Lines as Experimental Prototype Models: Present Status 16 References 20 I. INTRODUCTION Studies by classical endocrinologists developed the concept that tropic hormones (proteins, such as the gonadotropins, or steroids, such as estrogens) are endocrine regulators involved in the control of mammalian cell proliferation in vivo. The experimental conclu sions of these studies were derived from observations of the effects of endocrine organ ablation (hypophysectomy, ovariectomy, adrena- lectomy, etc.) and hormone replacement therapy (administration of 1