Table Of ContentIntersexuality
in the Animal Kingdom
Edited by R. Reinboth
With 221 Figures
Springer-Verlag
Berlin Heidelberg New York 1975
Professor Dr. Rudolf Reinboth
Institut fUr Zoologie
der Johannes Gutenberg-Universitat
0-65 Mainz, SaarstraBe 21
ISBN-13 978-3-642-66071-9 e-ISBN-13: 978-3-642-66069-6
001: 10.1007/978-3-642-66069-6
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© by Springer-Verlag Berlin' Heidelberg 1975.
Softcover reprint of the hardcover 1s t edition 1975
Library of Congress Cataloging in Publication Data. Symposium on Inter
sexuality in the Animal Kingdom, Akademie der Wissenschaften und der
Literatur, 1974. Intersexuality in the animal kingdom. Bibliography: p.lncludes
index. 1. Hermaphroditism--Congresses. 2. Sex--Cause and determination-
Congresses. I. Reinboth, Rudolf, ed. II. Title. [DNLM: 1. Hermaphroditism-
Congresses. 2. Sex determination--Congresses. 3. Sex reversal, Gonadal-
Congresses. WJ712 S 986i 1974]. OP267.S94. 1974. 591.1'6'67. 75-4585
The use of registered names, trademarks, etc. in this publication does not
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Preface
When Richard Goldschmidt' coined the term "intersexuality" in 1915,
he intended it to apply to normally dioecious species which exhibit
some kind of mixture between male and female characters. However, as
knowledge of the bewildering variability present in the sexual orga
nization of members of the animal kingdom has increased, the original
meaning of the word has changed. Today many authors define inter
sexuality as "the presence of both male and female characteristics,
or of intermediate sexual characteristics, in a single individual".2
This more extensive and widely accepted concept justifies the title
of our book •.
Among all the anatomical and physiological features of living organisms
the reproductive system has a unique importance for the perpetuation
of the species. Conversely, reproductive processes are of little or
no account for the viability of the individual. Therefore, within the
framework of general biology reproduction has all too often been looked
at solely from the point of view of genetics. Lively discussions about
genotypic versus phenotypic sex determination long dominated the sci
entific literature on sexuality in animals; this one-sided emphasis
has tended to obscure many important facets of an organism's ability
to reproduce.
Recent developments in current biological research have brought the
classic problem of sex differentiation into focus again, and the rapid
progress being made in comparative endocrinology has added a new di
mension to the study of reproductive biology.
Whatever is understood by intersexuality, the term always implies
some deviation from the usual gonochoristic pattern and acknowledges
the indisputable fact that sex depends on a strict dualism of "male
ness" and "femaleness", at least at the level of the gametes. The
basically disjunctive character of the two types of sexual potency
means that the occurrence of intersexual phenomena links the two
extremes, so offering an insight into nature's differentiation be
tween "male" and "female" and the processes involved.
Such ideas were the leitmotif for a symposium that brought together
research workers from such varied biological disciplines as endo
crinology, developmental physiology, genetics, cytology, ecology
and evolutionary theory. The papers presented there have been edited
for this book, which gives a fairly detailed picture of the problems
'Biol. Zbl. 35, 565 (1915).
2Atz,J.W.: In: Intersexuality in Vertebrates Including Man (C.N.
Armstrong and A.J. Marshall, eds.). New York: Academic Press, 1964.
VI
and objectives of current research on intersexuality. No claim is
made to present an exhaustive review of current knowledge, but the
reader will find ample information in the literature cited.
At first sight the heterogeneity of the contributions is rather
startling. Some authors reported details of original studies where
as others tried in different ways to provide a synoptic review of
certain topics. Studies on intersexuality are being carried out on
various groups in quite different directions. For example, in teleost
fishes surprising discoveries of a wide array of different types of
natural intersexuality have aroused the curiosity of research workers
in different disciplines, whereas in other classes (e.g. turbellaria,
echinoderms, etc.) the scope for study is more limited because less
is known about the biology of these animals. The reader may look in
vain for information on some taxonomic classes in which intersexual
organization is either the rule (trematodes and cestodes) or fairly
cornmon (nematodes) because of the few recent findings available.
Professor Nigon of the University of Lyon, France, an expert on
nematodes, wrote to me: "Since my own studies on sexuality in nema
todes (of which the investigation of intersexuality represented only
a limited part), this subject does not appear to have been pursued
further anywhere in the world. I am aware of only sporadic observa
tions in this field ... which cannot be compared with the active
experimental studies being carried out on other invertebrates."
Despite such unevenness, there was unanimous agreement among the
contributors that the exchange of ideas about intersexuality was
stimulating and deserved a wider audience. Moreover, the historical
subdivisions of the biological disciplines have not favoured attempts
to examine intersexuality as a highly complex phenomenon, requiring
joint efforts from various angles.
No single theme is likely to emerge as the "essence" of this book,
but one may risk the opinion that the problem of "bisexual potency",
a term introduced by Max Hartmann in 1923,3 is an important reference
point. Answers have yet to be found to the question as to which pro
cesses are involved in inducing (bi-?) potential (germ) cells to stop
mitotic divisions and to enter a development (by meiosis or similar
events) which leads irreversibly to the formation of clearly polarized
male and female gametes.
The critical reader is also asked to keep in mind that one of the
unavoidable obstacles facing the editor of such a symposium is that
English is not the mother tongue of all contributors. Both publisher
and editor made great efforts to achieve an evenness in grammar and
style, by revising and sometimes even retranslating. Nevertheless,
rapid publication called for an acceptable compromise that did not
allow for "smoothing the rough edges".
Please note that the index is merely meant to supplement the key words
of the article titles. Terms basic to the general contents (e.g. gonado
genesis, germ cells, gametogenesis, ovary, testis, gonoducts, sex dif
ferentiation, etc.) have deliberately been omitted because they are
dealt with in almost every contribution.
3Studia Mendeliana Brlinn, 203 (1923).
~I
The Symposium on Intersexuality in the Animal Kingdom was held in the
Akademie der Wissenschaften und Literatur at Mainz on 8-12 July 1974.
The participants are much obliged to the hosts, who provided a most
pleasant environment for our discussions. We acknowledge the generous
financial help granted by the Deutsche Forschungsgemeinschaft, Bonn,
the Akademische Auslandsamt der Universit~t Mainz, the government of
Rheinland-Pfalz, the Vereinigung der Freunde der Universit~t Mainz,
the Humboldt-Stiftung, Bonn, Farbwerke Hoechst AG, and C.H. Boehringer
Sohn, Ingelheim. Members of the staff of the Akademie der Wissenschaf
ten and the Zoological Institute of the University of Mainz gave us
broad support. Dr. J.A.P. Mehl as copyeditor has worked intensively
and constructively on most of the manuscripts. Without the personal
engagement and cooperation of my secretary, Mrs. A. Reichow, the
whole project would never have materialized.
The publisher met all our interests in a very efficient and always
friendly and sympathetic way.
To all persons and institutions who lent us their support I am deeply
bound in gratitude.
March 1975 R. Reinboth
Contents
INVERTEBRATES
Sex and Sex Determination in Coelenterates •••.•..•.••••.••....••
P. Tardent
Sexual Differentiation in the Fissiparous Strain of Dugesia
gonoaephaZa • • . • • • • • • . • • • • • • • • • • • • • • • • • • • • . • • • • • • • • • • • • • • • • • • • • • 14
Th. Lender and C. Brian90n
Sexuality and Neurosecretion in Freshwater Planarians •.•••.•.••• 20
M. Grasso
Sex Differentiation in Regeneratingd' I ~ Nemertine Chimeras ••.••• 30
J. Bierne
Sex Reversal in the SyUinae (Polychaeta: Annelida) 41
M. Durchon
Mutual Influence on the Sexual Differentiation in the Protandric
Polychaete Ophryotroaha pueriUs •• • • • • • • • • . • • • • • • • • • • • • • • • • • • • • • • 48
H.-D. Pfannenstiel
Neurosecretory Phenomena during Reproduction in Oligochaeta .••.. 57
H. Herlant-Meewis
Study of Sex Control of Gametogenesis by Organ Culture in the
Oligochaete Annelid Eisenia foetida f. typiaa Sav. .•••..••••..... 64
M. Lattaud
Factors of Primary Sexual Differentiation in the Simultaneous
Hermaphrodite Eisenia foetida (Oligochaeta: Lurnbricidae) •...•.• 72
J.C. Relexans
Sex-Determination in BoneUia ••••••••••••••••••.••.••••••••••.••• 84
R. Leutert
Hermaphroditism and Gynandromorphism in Malacostracan Crustacea. 91
H. Charniaux-Cotton
Temperature-Sensitive Intersexuality and Its Determinism in
Orahestia gammareZZa Pallas .•.•.••..•.•••.•....••.••••..••..••.. 106
T. Ginsburger-Vogel
Parasite-Induced Castration and Intersexuality in Insects ••.••.. 121
w.
Wiilker
Hermaphroditism in Insects. Studies on Iaerya purahasi •••••••••.••• 135
M. Royer
x
Sex Specific Cell Differentiation in Different Types of
Intersexes of Lymantria dispar L. .•............................ 146
G.C. Mosbacher
Structural and Endocrinological Aspects of Hermaphroditism in
Pulmonate Snails, with Particular Reference to Lymnaea
stagnaZis (L.) •...••..••.•..........•..•••....•.......•.•.•... 158
J. Joosse
Protandric Hermaphroditism in Prosobranch Gastropods ........... 170
S. Le Gall and W. Streiff
The Sexuality of Pelecypod Molluscs ............................. 179
P. Herlin-Houtteville and P.E. Lubet
Hermaphroditism in Echinoderms. Studies on Asteroids ............ 188
R. Delavault
VERTEBRATES
The Gonadal and Adenohypophysial Functions on Natural Sex
Reversal ................................•.................... 201
S.T.H. Chan, Wai-sum 0, and S.W.B. Hui
Ovarian and Testicular Intersexuality in Two Protogynous
Mediterranean Groupers, EpinepheZus aeneus and EpinepheZus guaza ... 222
J. Brusl~ and S. Brusl~
Hermaphroditism in Cichlid Fishes ............................... 228
H.M. Peters
An Unusual Approach to Experimental Sex Inversion in the Teleost
Fish, Betta and Maeropodus .....••......•.•••.......•...•....... 236
P. Becker, H. Roland, and R. Reinboth
The Possible Significance of Sex-Chromatin for the Determination
of Genetic Sex in Ambisexual Teleost Fishes .................. 243
J.A.P. Mehl and R. Reinboth
Sex Determination and Differentiation among Uniparental
Homozygotes of the Hermaphroditic Fish RivuZus marmoratus
(Cyprinodontidae: Atheriniformes) ...........•................ 249
R.W. Harrington, Jr.
Protogynous Hermaphroditism in Fishes of the Family Scaridae .... 263
J.H. Choat and D.R. Robertson
Ecology and Physiology of Sex Reversal in Anthias squamipinnis
(Peters), (Teleostei: Anthiidae) .....•...............•....... 284
L. Fishelson
The Evolution of Hermaphroditism in Fishes ...................... 295
C.L. Smith
Unisexual Female Offsprings in the Salamander, PZeurodeZes waUUi
Michah ...........•.....................................•..... 311
A. Collenot
XI
Autoradiographic Study on the Mechanisms of Testosterone-
Induced Sex-Reversal in Rana Tadpoles ........................ 318
E. Vannini, A. Stagni, and F. Zaccanti
Temperature and Sex Differentiation in Embryos of Two Chelonians,
Emys orbicula:r>is L. and Testudo graeca L ........................ 332
C. Pieau
Evolution of Parthenogenetic Species of Reptiles ................ 340
Ch.J. Cole
Intersexuality in Birds. Study of the Effects of Hybridization
and Post-Embryonic Ovariectomy ............................... 356
L. Gomot
Some Effects of Embryonic Gonad and Non-Gonadal Grafts on the
Development of Primary Sexual Characteristics in the Chick ... 375
J. Thiebold
Intersexuality of the Genital System and "Free-Martinism" in
Birds ..................... 382
c •••••••••••••••••••••••••••••••••••
H. Lutz and Y. Lutz-Ostertag
The Prenatal Development of Bovine Freemartins .................. 392
A. Jost, J.P. Perchellet, J. Prepin, and B. Vigier
The Role of Androgens in Sexual Differentiation of Mammals ...... 407
F. Neumann, W. Elger, H. Steinbeck, and K.-J. Graf
Recent Studies on the Intersexual Programming of the Genetic Rat
Male Pseudohermaphrodite ..................................... 422
A.S. Goldman
Chromosomes and Sex Differentiation ............................. 438
U. Mittwoch
Subject Index ................................................... 447
List of Contributors
BECKER, Peter, Dr., c/o Celamerck GmbH & Co. KG, 6507 Ingelheim (Rhein),
W. Germany
BIERNE, J., Prof., Faculte des Sciences de l'Universite de Reims,
Laboratoire de Biologie generale, 51062 Reims Cedex, France
BRIANCON, C., Dr., Faculte des Sciences, Laboratoire de Biologie
animale, 91405 Orsay, France
BRUSL~, Jacques, Prof., Centre Universitaire de Perpignan, Faculte des
Sciences,. Laboratoire de Biologie marine, 66000 Perpignan, France
BRUSL~, Solange, Maitre-Assistant, Centre Universitaire de Perpignan,
Faculte des SCiences, Laboratoire de Biologie marine,
66000 Perpignan, France
CHAN, S.T.H., Dr., Department of Zoology, University of Hong Kong,
Hong Kong
CHARNIAUX-COTTON, Helene, Prof., Universite Paris VI, Laboratoire
Sexualite et Reproduction des Invertebres, 75230 Paris Cedex OS,
France
CHOAT, J.H., Dr., University of Auckland, Department of Zoology,
Auckland, New Zealand
COLE, Charles J., Dr., American Museum of Natural History, Department
of Herpetology, New York, NY 10024, USA
COLLENOT, Alain, Prof., Universite Paris VI, 75230 Paris Cedex OS,
France
DELAVAULT, R., Prof., Universite d'Orleans, Laboratoire de Biologie
cellulaire et animale, 45045 Orleans Cedex, France
DURCHON, Maurice, Prof., Universite des Sciences et Techniques de
Lille, Biologie Animale, 59650 Villeneuve-d'Ascq, France
ELGER, Walter, Dr., c/o Schering AG, Dept. Endokrinpharmakologie,
1000 Berlin 65, W. Germany
FISHELSON, Lev, Prof. Dr., Tel-Aviv University, Department of Zoology,
Tel-Aviv, Israel
GINS BURGER-VOGEL , T., Maitre-Assistant, Laboratoire de Genetique
evolutive du CNRS, 91190 Gif sur Yvette, France
GOLDMAN, Allen, Associate Professor, M.D., The Children's Hospital of
Philadelphia, Division of Experimental Pathology, Philadelphia,
PA 19104, USA
