Comparative Endocrinology

285 Pages·1963·4.805 MB·English

by  U.S. Von Euler (Eds.)

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
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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.