Comparative Endocrinology PDF
Preview Comparative Endocrinology
COMPARATIVE ENDOCRINOLOGY EDITED BY U. S. von Euler Fysiologiska Institutionen, Karolinska Institutet, Stockholm, Sweden H. Heller Department of Pharmacology, University of Bristol, Bristol, England VOLUME II Part One INVERTEBRATE HORMONES Part Two TISSUE HORMONES 1963 ACADEMIC PRESS NEW YORK AND LONDON COPYRIGHT © 1963, BY ACADEMIC PRESS INC. ALL RIGHTS RESERVED. NO PART OF THIS BOOK MAY BE REPRODUCED IN ANY FORM, BY PHOTOSTAT, MICROFILM, OR ANY OTHER MEANS, WITHOUT WRITTEN PERMISSION FROM THE PUBLISHERS. ACADEMIC PRESS INC. Ill Fifth Avenue, New York 3, New York United Kingdom Edition published by ACADEMIC PRESS INC. (LONDON) LTD. Berkeley Square House, London W.l LIBRARY OF CONGRESS CATALOG CARD NUMBER: 63-16982 PRINTED IN THE UNITED STATES OF AMERICA CONTRIBUTORS Numbers in parentheses indicate pages on which the authors' contributions begin. N. AMBACHE, Medical Research Council, Department of Physiology, Royal College of Surgeons, London, England (128) HANS DUNER, Sabbatsbergs sjukhus, Stockholm, Sweden (239) V. ERSPAMER, Institute of Pharmacology, University of Parma, Parma, Italy (159) LAWRENCE I. GILBERT, Department of Biological Sciences, Northwestern University, Evanston, Illinois (1) J. ERIK JORPES, Chemistry Department II, Karolinska Institutet, Stock holm, Sweden (112) FRANCIS G. W. KNOWLES, Department of Anatomy, The Medical School, University of Birmingham, England (47) BENGT PERNOW, Serafimerlasarettet, Stockholm, Sweden (239) M. ROCHA Ε SILVA, Department of Pharmacology, Faculty of Medicine, University of Sao Paulo, Ribeiräo Preto, San Paulo, Brazil (64) U. S. VON EULER, Fysiologiska Institutionen, Karolinska Institutet, Stock holm, Sweden (209) V. P. WHITTAKER, Department of Biochemistry, Agricultural Research Council Institute of Animal Physiology, Babraham, Cambridge, England (182) ν PREFACE The aim of this book is to give readers with some basic knowledge of animal morphology, physiology, and chemistry, a systematic and com prehensive account of endocrine principles from the comparative point of view. It has been written by men who are actively engaged in research in the field which their contribution covers. It can therefore be hoped to present a critical and up-to-date picture of the comparative aspects of endocrinology to the medical scientist and zoologist generally and to furnish an adequately documented background to the research worker who is beginning to take an interest in one of the many endocrine systems described. The subject matter has been divided into three sections. The largest— which forms the contents of the first volume—deals with hormones orig inating in well-defined glandular organs and tissues and also reviews the relationships between the central nervous system and these endocrine complexes. The second section (Volume II, Part 1) discusses hormonal systems of invertebrates, and the third (Volume II, Part 2) contains a description of neurohormones and tissue hormones. This arrangement is based on the following considerations. As originally conceived, an endocrine organ was a discrete anatomical entity which elaborates and stores active principles that are then discharged into the blood to act as chemical messengers or hormones. This definition has the advantage of dividing the two main systems of intercellular communica tion—the endocrine apparatus and the nervous system—into two neat categories. However, it is doubtful whether this definition can now be maintained in its original simplicity. One of the "glands" involved, the neurohypophysis, has turned out to be only one part of a complex hor mone-producing system, consisting of secretory neurons (otherwise undis- tinguishable from neurons in the central nervous system) which in higher vertebrates deliver their products either to the neural lobe—where they are stored and then released to act as long-range hormone in line with the classical concept—or to the median eminence from which they enter the hypophyseal portal circulation to act as releasers of "tropic" hormones of the adenohypophysis. Thus this compound organization functions not only as a bridge between the two systems of communication but bears also a suggestive resemblance to processes in peripheral nerves. The chemi- vii viii PREFACE cal transmitters of peripheral nerve impulses, whether acetylcholine or noradrenaline, appear also to be formed in the cell bodies of the relevant neurons, to be transported along the corresponding axons, and to be concentrated and stored at the endings of these axons, either at their synaptic endings in ganglia or their neuroeffector junctions. Since evidence is accumulating that similar mechanisms of chemical transmission also take place at most of the synaptic junctions of the central nervous systems, a division into "true hormones" and "neurohormones" seems somewhat arbitrary. There exists a further—and rather varied—group of chemical mes sengers whose "systematic" position is not quite clearly defined. The substances in this group, which may be said to contain such biologically active principles as 5-hydroxytryptamine, heparin, and angiotensin, are apparently in some instances also produced by or stored in specific cells but their "physiological range" may or may not be restricted to the organ or tissue in which they originate. The distinction between these tissue factors and the more conventional hormones is again somewhat tenuous, and it was therefore felt that a discussion of their occurrence and properties should not be excluded from the present survey. Initially, and due no doubt to the endeavor to link this new branch of physiology to human endocrine disease, hormone research was almost exclusively concerned with mammals. During recent years, however, the comparative aspect has come very much to the fore. The morphology of endocrine organs throughout the vertebrate phylum has been intensively studied and a good beginning has been made with the chemical identifi cation of hormones of lower vertebrates. The results to date suggest an astonishing constancy in the chemistry of the endocrine principles: the adrenal hormones, both "medullary" and "cortical," for example, do not seem to vary from fish to man, though subtle differences in the composition of protein or peptide hormones have recently come to light. It has also been shown that endocrine mechanisms in certain groups of invertebrates are as important as in vertebrates. Moreover, there are distinct resem blances in organization, as manifested for example by neurohormonal interaction. But the chemistry of invertebrate hormones is very much in its infancy. We wish to thank the contributors and the publishers for their patient collaboration. Our thanks are also due to the authors, societies, and pub lishers for permission to use illustrations and tables which have appeared in previous publications. U.S. VON EULER April 1968 H. HELLER CONTENTS OF VOLUME I Hypothalamic Control of Anterior Pituitary Claude Fortier Neurohypophyseal Hormones H. Heller Pituitary Hormones Affecting the Chromatophores N. Karkun and F. W. Landgrebe Female Gonadal Hormones Weiert Velle Male Gonadal Hormones G. J. van Oordt Adrenocortical Hormones J. G. Phillips and D. Bellamy Chromaffin Cell Hormones U. S. von Euler Thyroid Hormones A. Gorbman Parathyroid Glands Roy 0. Greep Pancreatic Hormones: Insulin F. (?. Young Pancreatic Hormones: Glucagon Berthet Comparative Biochemistry of Adenohypophyseal Hormones Choh Hao Li The Physiology of the Adenohypophyseal Hormones E. Knobil and R. Sandler AUTHOR INDEX · INDEX OF SPECIES · SUBJECT INDEX xiii ~ 14 ~ Hormones Controlling Reproduction and Molting in Invertebrates LAWRENCE I. GILBERT Department of Biological Sciences, Northwestern University, Evanston, Illinois I. INTRODUCTION 1 II. PROTOZOA 2 III. CEPHALOPODA ---- 4 IV. CRUSTACEA 5 A. Introduction 5 B. X-Organ Sinus Gland 5 C. Y-Organ 6 D. The Ovary 7 E. Androgenic Gland 8 F. Sex Reversal 9 G. Summary 9 V. INSECTS — 0 1 A. Introduction 10 B. Hormonal Control of Metamorphosis. 10 C. Salient Problems 4 1 ADDENDUM 3 3 REFERENCES 7 3 I. INTRODUCTION It is more than 100 years since Berthold (1849) established the endocrine function of the mammalian testis. The endocrine control of sexual develop ment and gestation is now well established for mammals and much is known about hormonal mechanisms in other vertebrates. Unfortunately our knowledge of invertebrate hormones is far less complete and most of our information concerns the arthropods. For example, it has been shown 1 Original work from the author's laboratory was supported by grant A-2818 from the National Institute of Arthritis and Metabolic Diseases of the National Institutes of Health. Review of the literature was concluded April, 1961. 1 2 LAWRENCE I. GILBERT that endocrine mechanisms control molting and metamorphosis in insects (Bodenstein, 1954, 1957; Butenandt, 1959; Campbell, 1959; Gilbert and Schneiderman, 1961a; Karlson, 1956; Novak, 1959; Pflugfelder, 1958; Schneiderman and Gilbert, 1959; Williams, 1952; Wigglesworth, 1954, 1957, 1959) and molting in crustaceans (Knowles and Carlisle, 1956; Carlisle and Knowles, 1959; Passano, 1960). The most recent reviews regarding control of reproduction in these two large groups of animals are those of Engelmann (1960a) and Wigglesworth (1960a) for the insects, and Charniaux-Cotton, (1960a) and Carlisle and Knowles (1959) for the crustaceans. As far as other invertebrates are concerned, only a few de cisive experiments have been conducted. The information has been most exactly reviewed by Scharrer (Scharrer, 1953, 1955a; Scharrer and Schar- rer, 1954). In a limited review of this type it would be impossible to discuss in detail the mass of data which indirectly indicates hormonal control of reproduction in many of the animals studied. Some of this information is listed in Table I, but for the most part this review will consider the control of molting and reproduction in arthropods, and particularly the insects. This is as much due to the lack of information regarding other groups as to the author's own interests. IL PROTOZOA Recently Cleveland and his associates (Cleveland, 1959; Cleveland and Burke, 1960; Cleveland et al., 1960) have shown that ecdysone, the molt ing hormone of insects, triggers gametogenesis in certain symbiotic flagel lates that inhabit the gut of the woodroach, Cryptocercus punctulatus. Normally gametogenesis occurs in these protozoa when the host roach molts. Any procedure that interferes with molting in the roach interferes with gametogenesis in the protozoa. Since the adult insect never molts, the protozoa in adult roaches never undergo gametogenesis. However, injection of ecdysone into the adult roach causes the onset of sex in these flagellates even at concentrations too low to cause molting in the roach. One may ask whether ecdysone acts directly on the protozoa or indirectly through metabolic changes in the host prior to the molt. No experiments have tested the effect of crystalline ecdysone on these protozoa in cul ture, but Cleveland et al. (1960) state that "the fact that some genera of the flagellates react in a remarkably short time and undergo gametogenesis within three hours may best be explained by a direct action of the hor mone on the protozoa. The fact that ecdysone induces gametogenesis in the flagellates of an adult host which, so far as one can see, makes no 14. REPRODUCTION AND MOLTING IN INVERTEBRATES 3 TE GROUPS Reference Cleveland et al., 1960 (see text) Bobin and Durchon, 1952, 1953; De-fretin, 1952; Durchon, 1948, 1949, 1951, 1952, 1953, 1956a, b; Durchon and Frezal, 1955; Gabe, 1954-Hauenschild, 1956; Herlant-Meewis, 1956a, b; Hubl 1953; Michon, 1953; Scharrer, 1941 Kenk 1941 (see also Vandel 1920, 1921) Veillet, 1941 Gabe, 1953a, 1954; Herlant-Meewis, 1959; Laviolette, 1956; Lubet, 1956, 1957 Wells and Wells, 1959 (see text) Carlisle, 1950,1951 (see also Butcher, 1930; Hogg, 1937) A TABLE I HORMONAL CONTROL OF REPRODUCTIVE PROCESSES IN SOME INVERTEBR Endocrine relationship Sexuality induced by insect molting hormone Definite relationship established between activity of neurosecretory cells in the cerebral ganglion and gonad development. Evidence indicates that the humoral substance may inhibit gonad maturation in polychaetes and oli-gochaetes and may also be involved in the maintenance of the clitellum and the process of egg laying Gonads may secrete a hormone necessary for development of the copulatory organs Transformation of larva to sexually mature adult worm may be under endocrine control Adult gastropod gonad liberates some chemical mediator that conditions the accessory glands of the genital tract. As in the annelids, there appears to be a definite relationship between neurosecretory activity and gonad development in gastropods and lammelibranchs, as well as evidence for humoral control of gamete release Optic glands of cephalopods control gonad maturation Neurohumoral mechanism postulated for gamete release in ascidians m hes a Phylu Protozoa Annelida Platyhelmint Phoronidea Mollusca Protochordat 4 LAWRENCE I. GILBERT attempt whatever at molting also suggests the possibility of direct action." From an evolutionary viewpoint, these protozoa appear to have utilized a particular chemical agent in their environment to trigger sexual changes just as many higher animals have utilized physical agents in their envi ronment (e.g., day length). Whether ecdysone actually participates in the same biochemical processes in insects and protozoa is unknown. III. CEPHALOPODA In the study of behavior and learning in cephalopods (Boycott and Young, 1955; Young, 1958; Wells and Wells, 1956, 1957, 1958) the effect of brain lesions on tactile responses was carefully noted. Boycott and Young (1956) observed that after optic tract section, many animals devel oped enlarged gonads. Wells and Wells (1959) in an elegant set of experi ments showed that sexual maturation in Octopus was under hormonal control (see also Wells, 1960). Lesions in a particular part of the brain mass caused a hundredfold increase in the size of the ovary and a 50% increase in the size of the testes. These lesions were always correlated with hypertrophy of the optic glands, two small bodies lying on the optic stalks. This work indi cates that gonad maturation in Octopus is controlled by a hormone (s) released from the optic glands which in turn are regulated by inhibitory nerves. Cutting these inhibitory nerves always results in hypertrophy of the optic glands followed by gonad enlargement. Blinding of Octopus, optic lobe removal, severing the optic nerve, or optic tract section also causes precocious gonad maturation, presumably by affecting certain brain centers which in turn release the optic glands from their rigid con trol by the inhibitory nerves. This result led Wells and Wells to suggest that gonad maturation in Octopus may depend on photoperiod. The optic glands have been found to secrete before the ovary is at a stage competent to respond. However, secretion of the optic glands in these immature animals is thought to be responsible for maturation of the oviducts, ovisac, and oviducal glands (Wells, 1960). Optic glands occur in almost all cephalopods studied and may control gonad maturation in all these forms. Further experiments revealed that it was the highest center of the nervous system, the brain, that ultimately controls gonad maturation. It is believed that this may be characteristic of animals that depend on learning to influence their behavior, and that in the evolution of these organisms special mechanisms were developed to insure a delay in sexual maturity until the brain was fully developed.
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