Invertebrate Hormones: Tissue Hormones

INVERTEBRATE HORMONES: TISSUE HORMONES

COMPARATIVE ENDOCRINOLOGY

U.S. von Euler

Fysiologiska Institutionen, Karolinska Institutet, Stockholm, Sweden

H. Heller

Department of Pharmacology, University of Bristol, Bristol, England

Table of Contents

Cover image

Title page

COMPARATIVE ENDOCRINOLOGY

Copyright

CONTRIBUTORS

PREFACE

CONTENTS OF VOLUME I

Part One: INVERTEBRATE HORMONES

Chapter 14: Hormones Controlling Reproduction and Molting in Invertebrates

Publisher Summary

I INTRODUCTION

II PROTOZOA

III CEPHALOPODA

IV CRUSTACEA

V INSECTS

ADDENDUM

Chapter 15: The Structure of Neurosecretory Systems in Invertebrates

Publisher Summary

I INTRODUCTION

II NEUROSECRETORY SYSTEMS OF THE HEAD REGION

III NEUROSECRETORY SYSTEMS OF THE THORAX AND ABDOMEN

IV CONCLUSIONS

Part Two: TISSUE HORMONES

Chapter 16: Kinins: Bradykinin, Angiotensin, Substance P

Publisher Summary

I DEFINITIONS

II BRADYKININ (PLASMAKININ, KALLIDIN)

III ANGIOTENSIN (HYPERTENSIN, ANGIOTONIN)

IV SUBSTANCE P

Chapter 17: Heparin

Publisher Summary

I INTRODUCTION

II EARLY HISTORY

III METHODS OF PREPARING HEPARIN

IV CHEMISTRY

V ACTION MECHANISM

VI THE TISSUE MAST CELLS

VII ANTICOAGULANT THERAPY

Chapter 18: Physiologically Active Lipid Anions

Publisher Summary

I INTRODUCTION*

II PROSTAGLANDIN

III DARMSTOFF

IV BIOLOGICALLY ACTIVE UNSATURATED FATTY ACIDS WITHOUT ALCOHOLIC HYDROXYL GROUPS

V IRIN

VI LIPID-SOLUBLE ACID FROM NASAL MUCOSA

VII ENDOMETRIAL ACIDS IN MENSTRUAL FLUID

VIII SRS-A

Chapter 19: 5-Hydroxytryptamine

Publisher Summary

I INTRODUCTION

II OCCURRENCE AND DISTRIBUTION

III BIOSYNTHESIS AND FATE

IV TURNOVER RATE

V PHYSIOLOGICAL AND PHARMACOLOGICAL ACTIONS: BIOLOGICAL SIGNIFICANCE

Chapter 20: Cholinergic Neurohormones

Publisher Summary

I INTRODUCTION

II NATURALLY OCCURRING, PHARMACOLOGICALLY ACTIVE CHOLINE ESTERS OTHER THAN ACETYLCHOLINE

III ACETYLCHOLINE AS A TRANSMITTER SUBSTANCE

Chapter 21: Adrenergic Neurohormones

Publisher Summary

I DEFINITION OF ADRENERGIC TRANSMITTER SYSTEMS

II NATURE OF TRANSMITTER SUBSTANCES

III DISTRIBUTION OF ADRENERGIC NEUROHORMONES

IV METABOLISM OF ADRENERGIC NEUROHORMONES

V ACTION OF DRUGS ON RELEASE OF ADRENERGIC TRANSMITTERS

VI PLASMA LEVELS AND EXCRETION IN URINE

VII PHYSIOLOGICAL FUNCTIONS OF ADRENERGIC NEUROHORMONES

ACKNOWLEDGMENT

Chapter 22: Histamine

Publisher Summary

I OCCURRENCE AND DISTRIBUTION

II METABOLISM

III PHARMACOLOGICAL ACTIONS

IV ANTIHISTAMINE DRUGS

V PHYSIOLOGICAL SIGNIFICANCE

VI PATHOLOGICAL CONDITIONS

Author Index

Index of Species

Subject Index

COMPARATIVE ENDOCRINOLOGY

A Treatise in Two Volumes

VOLUME I

GLANDULAR

HORMONES

VOLUME II

Part One

INVERTEBRATE

HORMONES

Part Two

TISSUE

HORMONES

Copyright

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.

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United Kingdom Edition published by

ACADEMIC PRESS INC. (LONDON) LTD.

Berkeley Square House, London W.1

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 DUNÉR,     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, Stockholm, 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 E. SILVA,     Department of Pharmacology, Faculty of Medicine, University of São Paulo, Ribeirão Preto, Sãn Paulo, Brazil (64)

U.S. VON EULER,     Fysiologiska Institutionen, Karolinska Institutet, Stockholm, Sweden (209)

V.P. WHITTAKER,     Department of Biochemistry, Agricultural Research Council Institute of Animal Physiology, Babraham, Cambridge, England (182)

PREFACE

The aim of this book is to give readers with some basic knowledge of animal morphology, physiology, and chemistry, a systematic and comprehensive 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 originating 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 communication—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 hormone-producing system, consisting of secretory neurons (otherwise undistinguishable 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 chemical 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 messengers 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 identification 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 resemblances 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 publishers for permission to use illustrations and tables which have appeared in previous publications.

U.S. VON EULER and H. HELLER

April 1963

CONTENTS OF VOLUME I

Hypothalamic Control of Anterior Pituitary

Claude Fortier

Neurohypophyseal Hormones

H. Heller

Pituitary Hormones Affecting the Chromatophores

j . N. Karkun and F. W. Landgrebe

Female Gonadal Hormones

Weiert Velie

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 O. Greep

Pancreatic Hormones: Insulin

F. G. Young

Pancreatic Hormones: Glucagon

J. Berthet

Comparative Biochemistry of Adenohypophyseal Hormones

Choh Hao Li

The Physiology of the Adenohypophyseal Hormones

E. Knobil and R. Sandier

AUTHOR INDEX • INDEX OF SPECIES • SUBJECT INDEX

Part One

INVERTEBRATE HORMONES

∼14∼

Hormones Controlling Reproduction and Molting in Invertebrates¹

LAWRENCE I. GILBERT,     Department of Biological Sciences, Northwestern University, Evanston, Illinois

Publisher Summary

This chapter describes hormones that are responsible for controlling reproduction and molting in invertebrates. The chapter also describes that ecdysone, the molting hormone of insects, triggers gametogenesis in certain symbiotic flagellates 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. It has been found in an extended series of investigations on seven species of roaches that virgin female adults produce volatile sex attractants that act via the male’s antennal chemoreceptors. In addition, production of this pheromone is under the control of the corpus allatum. Allatectomy results in a failure of pheromone production that can be corrected by implantation of active glands. It appears that the active principle regulating pheromone production may be identical to that controlling egg maturation as pheromone release ceases at those stages in the reproductive cycle when the gonadotropic hormone is not produced.

I. INTRODUCTION

II. PROTOZOA

III. CEPHALOPODA

IV. CRUSTACEA

A. Introduction

B. X-Organ Sinus Gland

C. Y-Organ

D. The Ovary

E. Androgenic Gland

F. Sex Reversal

G. Summary

V. INSECTS

A. Introduction

B. Hormonal Control of Metamorphosis.

C. Salient Problems

ADDENDUM

REFERENCES

I INTRODUCTION

It is more than 100 years since Berthold (1849) established the endocrine function of the mammalian testis. The endocrine control of sexual development 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 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 decisive experiments have been conducted. The information has been most exactly reviewed by Scharrer (Scharrer, 1953, 1955a; Scharrer and Scharrer, 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.

TABLE I

HORMONAL CONTROL OF REPRODUCTIVE PROCESSES IN SOME INVERTEBRATE GROUPS

II PROTOZOA

Recently Cleveland and his associates (Cleveland, 1959; Cleveland and Burke, 1960; Cleveland et al., 1960) have shown that ecdysone, the molting hormone of insects, triggers gametogenesis in certain symbiotic flagellates 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 culture, 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 hormone 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 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 environment (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 developed enlarged gonads. Wells and Wells (1959) in an elegant set of experiments 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 indicates 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 control 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.

IV CRUSTACEA

A Introduction

Both Zeleny (1905) and Megusar (1912) noted that precocious growth resulted when both eyestalks were removed from certain decapod crustaceans. Since that time a large body of information has been gleaned regarding molting in crustaceans although the exact number of hormones involved and the decision as to whether the information collected on brachyurans is applicable to other groups, are still open questions.

The earlier observations on the induction of molting after eyestalk removal were independently rediscovered by Brown and Cunningham (1939) and Abramowitz and Abramowitz (1939, 1940). Brown and Cunningham (1939) conducted the critical experiment indicating the endocrine control of molting when they implanted sinus glands into eyestalkless animals, a maneuver that proved to suppress the effect of eyestalk ablation. The technique of sinus gland removal was a major surgical achievement in that future experiments along these lines could be conducted without blinding the animals and destroying parts of the nervous system (see also Brown et al., 1939; Brown, 1942; Panouse, 1946; Kleinholz, 1947).

B X-Organ Sinus Gland

1 ROLE IN MOLTING

In the brachyurans a group of neurosecretory cells with converging axons that end in the sinus gland (a blood sinus bounded by a connective tissue sheath) comprises the X-organ. Since the axons arise from the medulla terminalis ganglion, the organ has been named the medulla terminalis x-organ (MTGX). In other Crustacea there may be a sensory papilla x-organ (SPX) since the axons emerge from the sensory papilla (Carlisle and Knowles, 1959). The sinus gland presumably stores neurosecretory material from the X-organ and releases it into the blood at the appropriate time. It is a matter of conjecture as to whether the cells of the sinus gland further contribute to the stored material (Passano, 1953; Miyawaki, 1956).

The surgical removal of the X-organ was a most difficult feat, but has been accomplished by several workers (Passano, 1953; Pasteur, 1958). Overwhelming evidence has led to the hypothesis that a molt-inhibiting hormone is secreted by the neurosecretory cells of the X-organ and is stored in the sinus gland (Bliss, 1951, 1953; Bliss et al., 1954; Carlisle, 1953b; Enami, 1951; Passano, 1953, 1954). This material when liberated into the general circulation inhibits molting, and accounts for the earlier results that indicated precocious molting after eyestalk ablation or X-organ extirpation.

Another humoral factor termed the molt-accelerating hormone has been postulated (Carlisle and Dohrn, 1952, 1953). Carlisle (1960) suggests that this material is both secreted by and stored in the x-organ and functions to coordinate the manifold processes constituting the premolt stage. Whether this substance is common to the Crustacea or exists at all is questionable (cf. Passano, 1960).

2 ROLE IN REPRODUCTION

It is now well established that eyestalk ablation in young females or nonbreeding mature females leads to rapid enlargement of the ovary (Panouse, 1943, 1944, 1946, 1947; Brown and Jones, 1947, 1949; Stephens, 1952; Carlisle, 1954) and to oviposition (Brown and Jones, 1949; Stephens, 1952; Démeusy and Veillet, 1952).

Various surgical maneuvers and injection of eyestalk extracts showed that part of the eyestalk secreted a humoral substance that prevented ovarian maturation, the so-called ovarian-inhibiting hormone. Ablation of the eyestalk removes the ovary from inhibition and permits it to mature. The x-organ sinus gland complex located in the eyestalks is the source of this hormone. The neurosecretory complex comprising the X-organ secretes the ovarian-inhibiting hormone which is stored in the sinus gland and finally released to the general circulation. When the titer is sufficiently high there is no ovarian development. A similar mechanism may control testis development in Carcinus (Démeusy, 1953). This hormone may be similar in structure to the queen-substance of bees (Carlisle and Butler, 1958; see Section V,C,2).

C Y-Organ

1 ROLE IN MOLTING

Another endocrine organ described in more than fifty species of malacostracans is the Y-organ (Gabe, 1953a, 1956). This organ is located in the antennary segment of animals possessing a maxillary excretory organ and in the second maxillary segment in forms having an antennary excretory organ. When the Y-organs are bilaterally removed the molting cycle is blocked (Echalier, 1954, 1955, 1959; see also Carlisle, 1957b; Jyssum and Passano, 1957), and it has been generally accepted that the Y-organ is the molting gland of many crustaceans. Eyestalk removal does not result in molting in animals deprived of the Y-organ but implantation of the Y-organ leads to the premolt stage (see Drach, 1936, 1939; Carlisle and Dohrn, 1953 for staging of the molting cycle), and all available evidence indicates that the Y-organ is needed for the initiation of premolt. The morphological and physiological similarity between the Y-organ and the prothoracic glands of insects has been pointed out by Gabe (1953b). Both are nonneurosecretory and release a hormone that initiates the molting process.

Although the Y-organ degenerates in some crabs after the last molt (Carlisle, 1957b) and thus shows further similarity to the prothoracic glands, it continues to exist and function in many species of Crustacea leading to an adult life of persistent molts and increasing size. In other Crustacea in which the Y-organ remains intact in nonmolting adults, the X-organ presumably maintains a high circulating titer of molt-inhibiting hormone. It is believed that cessation of secretion of the molt-inhibiting hormone removes the reins of inhibition from the Y-organ, resulting in secretion of the molting hormone which then sets in motion all those processes leading to the premolt, and finally to ecdysis in an almost auto-catalytic manner.

2 ROLE IN REPRODUCTION

Echalier (1954) and Arvy et al. (1954) showed that Y-organ extirpation led to a rapid decline in the rate of gametogenesis and to gonad atrophy. This only occurred in animals operated on before maturity, and Y-organ removal in mature crustaceans had no effect. It is not yet known whether ablation of the Y-organ leads to a general metabolic disturbance ultimately causing a decline in the rate of gametogenesis or whether the Y-organ secretes a substance specific for gonad maturation.

D The Ovary

Most information about the endocrine role of the ovary has come from experiments involving ovary removal. This has been accomplished by irradiation (Callan, 1941; Haemmerli-Boveri, 1926; Knowles and Callan, 1940), parasitic castration (cf. Reinhard, 1956), and surgical extirpation (Takewaki and Nakamura, 1944; Charniaux, 1952; 1953, Charniaux-Cotton, 1953; cf. Charniaux-Cotton, 1960a). The data reveal that ovariectomy affects behavior, and in particular suppresses secondary brooding characteristics. None of the morphological secondary sexual characteristics are affected. The mature ovary presumably secretes a hormone that releases secondary brooding behavior.

E Androgenic Gland

The androgenic gland described first by Charniaux-Cotton (1954) in the amphipod Orchestia gammarellus is the most impressive sex gland in the invertebrates. It was later described in other species of Crustacea (Balesdent-Marquet, 1958; Carlisle, 1959a,b,c; Charniaux-Cotton, 1956a, 1958a,b; Duveau, 1957; see also Katakura, 1959; Knowles and Carlisle, 1956) and has also been called the vas deferens gland by Carlisle because of its proximity to that anatomical structure. The most work regarding structure and function has been performed by Charniaux-Cotton (1954, 1955, 1956a,b, 1957a,b,c, 1958a,b, 1960b, cf. review, 1960a). The androgenic gland is a solid strand of cells near the vas deferens. The physiological significance of the androgenic gland was elucidated by means of some remarkable extirpation and implantation experiments and by histological study.

Implantation of the androgenic gland into an immature or developing female caused masculinization: ovaries rapidly transformed into testes and there was a progressive acquisition of male secondary sex characters. These included transformation of external characters such as the slender female claw to the stout male claw and the development of male sex ducts in the recipient female. The behavior of these masculinized females was also dramatically transformed as they attempted to copulate with normal females and actually mated with them.

Extirpation of the androgenic gland from the male resulted in the cessation of differentiation of secondary sexual characters and spermatogenesis, finally causing degeneration of the testis. An ovary implanted into a male after removal of the androgenic gland survived while the same ovary implanted into a normal male was readily transformed into a testis.

As summarized by Charniaux-Cotton (1960a, p. 430), The androgenic glands are the exclusive source of a hormone responsible for the development of all the characters of the male sex; spermatogenesis, differentiation of the sperm-duct primordia, growth of the secondary sex characters, and male sexual behavior. The secretion of the androgenic hormone can take place without any nervous coordination; in fact, the androgenic gland carries out its normal activity regardless of the region into which it is implanted.

The chemical nature of this androgenic hormone is not known although very preliminary experiments indicate that the active material is most likely not identical with mammalian androgenic substances (Jost and Charniaux-Cotton, 1960).

F Sex Reversal

Certain Natantia are protandric hermaphrodites and normally males undergo sex reversal and become reproducing females. While the animal is in the male phase of its life, it contains developing ovaries, and conversely, the testes may be retained after sex reversal. Thus it is the development and nature of the secondary sexual characteristics that determine whether the animal functions as a male or female.

At a critical molt, the male loses its copulatory appendages and secretes a new exoskeleton that covers the male genital pore. Under favorable conditions the oviduct opening may appear and the animal becomes a physiologically functioning female. This most amazing sequence of events is best expressed by the comment of Carlisle (1960, pp. 13–14), "Thus in Lysmata setica data, in which the one-stage sex reversal is common, I have observed an animal copulate as a male, molt, and within an hour of the first copulation, copulate again as a female."

Experiments by Carlisle (1953a) and Aoto and Noshida (1956) indicate that this sex-reversal molt may be under hormonal control. Eyestalk removal in Lysmata and Pandalus increases the number of males reversing sex at the next molt while eyestalk injections appear to cause the reverse. What glandular structure in the eyestalk or what type of substance is responsible is as yet unknown, although Carlisle (1957a) suggests that increasing and decreasing titers of the ovarian inhibiting hormone may be responsible for sex reversal.

G Summary

Like endocrinologists in other fields, the crustacean endocrinologist has strived to investigate the mode of action of hormones at the cellular and molecular levels. Attempts have been made to correlate biochemical changes at the premolt stage with the action of the crustacean molting hormone. Changes in oxygen consumption, phases of carbohydrate, lipid, and protein metabolism, exoskeleton formation, water uptake, etc., have been followed during the molting cycle (cf. Scheer, 1957; Passano, 1960), but there is as yet no direct evidence that any enzymatically mediated changes are direct results of the molting hormone.

Although the knowledge of hormonal control of reproduction including gonad maturation and the development of secondary sexual characters in the Crustacea is meager, the progress of late has been surprisingly rapid considering that it was only 15 years ago that Brown (1948, pp. 163–164) in his excellent review article stated, many observations support strongly that hormones are operative in the development and maintenance of male and female sexual characteristics. Crucial experiments have not yet been performed to enable us to conclude definitely that specific sex hormones are actually operating, and, if so, in what tissues they arise.

We now know that sex hormones (e.g., hormone from the androgenic gland) exist and several of the endocrine factors controlling egg development, behavior, etc., have been established. Considering the number of orders not yet investigated, crustacean endocrinology promises to be a fertile field for students of growth.

V INSECTS

A Introduction

Insect metamorphosis is the transformation of an immature larval individual into a reproducing adult of very different form, structure, and habit of life, and is under endocrine control. Notwithstanding the tremendous differences between caterpillar and moth, the cells of each appear to contain the same genetic information. Larval life appears to give expression to one battery of genes, adult life to another. We are a long way from understanding the chemical events that ensue between the genes on the one hand and the growth, development, and metamorphosis of the organism as a whole on the other. In many cases we have only begun to comprehend how organ systems themselves change morphologically during metamorphosis (Wigglesworth, 1954, 1957, 1959; Finlayson, 1956) let alone how cells and cell organelles transform during development. Although many gaps exist in our knowledge, it is possible to outline a reasonable picture of several aspects of insect metamorphosis.

B Hormonal Control of Metamorphosis

As in most higher organisms, the postembryonic growth of insects appears to be under hormonal control, at least during part of the life history. Although there are differences among insect groups, a basic pattern of hormonal control emerges. This pattern is seen to good advantage in the giant Saturniid silkmoths of which Hyalophora cecropia, Samia cynthia, and Antheraea polyphemus are well-known examples. The life cycle of these insects is typical of the Holometabola and consists of four larval molts, a larval-pupal molt; and a pupal-adult molt (fig. 1). The larval-pupal-adult transformations are what we term metamorphosis. In these silkmoths metamorphosis is usually interrupted soon after pupation