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A SCIENCE PUBLISHERS BOOK
Series on
Reproduction and Development in Aquatic
Invertebrates
Volume 1
Reproduction and
Development in Crustacea
Series on
Reproduction and Development in Aquatic
Invertebrates
Volume 1
Reproduction and
Development in Crustacea
T. J. Pandian
INSA Senior Scientist
Central Marine Fisheries Research Institute
Kochi-682018, Kerala
India
p,
A SCIENCE PUBLISHERS BOOK
Cover Illustrations: Peripheral figures show representatives of over 96% crustaceans brooding eggs on one or
the other parts of their body. Brooding eggs is a hallmark strategy of crustaceans. The remaining 4% crusta-
ceans (in the centre of the picture) shed their eggs (for more details see Fig. 2.10).
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Preface to the Series
Invertebrates surpass vertebrates not only in species number but also in
diversity of sexuality, modes of reproduction and development. Yet, we know
much less of them than we know of vertebrates. During the 1950s, the multi-
volume series by L. H. Hyman accumulated bits and pieces of information
on reproduction and development of aquatic invertebrates. Through a couple
of volumes published during 1960s, A. C. Giese and A. S. Pearse provided
a shape to the subject of Aquatic Invertebrate Reproduction. Approaching
from the angle of structure and function in their multi-volume series on
Reproductive Biology of Invertebrates during 1990s, K. G. Adiyodi and R. G.
Adiyodi elevated the subject to a visible and recognizable status.
Reproduction is central to all biological events. The life cycle of most
aquatic invertebrates involves one or more larval stage(s). Hence, an account
on reproduction without considering development shall remain incomplete.
With passage of time, publications are pouring through a large number of
newly established journals on invertebrate reproduction and development.
The time is ripe to update the subject. This treatise series proposes to (i) update
and comprehensively elucidate the subject in the context of cytogenetics and
molecular biology, (ii) view modes of reproduction in relation to Embryonic
Stem Cells (ESCs) and Primordial Germ Cells (PGCs) and (iii) consider cysts
and vectors as biological resources.
Hence, the first chapter on Reproduction and Development of Crustacea
opens with a survey of sexuality and modes of reproduction in aquatic
invertebrates and bridges the gaps between zoological and stem cell
research. With capacity for no or slow motility, the aquatic invertebrates have
opted for hermaphroditism or parthenogenesis/polyembryony. In many
of them, asexual reproduction is interspersed within sexual reproduction.
Acoelomates and eucoelomates have retained ESCs and reproduce
asexually also. However, pseudocoelomates and hemocoelomates seem not
to have retained ESCs and are unable to reproduce asexually. This series
provides possible explanations for the exceptional pseudocoelomates and
hemocoelomates that reproduce asexually. For posterity, this series intends
to bring out six volumes.
August, 2015 T. J. Pandian
Madurai-625 014
Preface
Being arthropods, the crustaceans molt to grow and/or reproduce. Of 52,000
species, > 96% crustaceans brood their eggs on their body. Consequently,
they share the available resources among intensely competing processes
(i) growth including molting, (ii) breeding and (iii) brooding. Hence, they are
unique among aquatic invertebrates and render the study of reproduction and
development a fascinating field of research. Not surprisingly, many books are
written but they are limited to one or another taxon or aspect of reproduction
and development. This book represents the first attempt to comprehensively
elucidate almost all aspects of reproduction and development covering from
anostracan Artemia to xanthid crabs.
Commencing with resource allocation to reproduction, the second and
third chapters display that the investment on generation time as a fraction
of life span averages to 28, 48 and 85% in tropical, temperate and Arctic
crustaceans. Consequently, the tropical cladocerans, for example, produce 10
times more neonates than their temperate counterparts. That their hallmark
strategy of brooding embryos costs considerable time and resource investment
is demonstrated by the fact that the egg shedding calanoids and penaeids
produce 3-20 times more eggs than the brooders. A second consequence of
brooding is the liability of the brooders to reproductive senescence. For the
first time, the book has adduced adequate evidence to establish the fact that
brooding gonochorics, parthenogenics and hermaphrodites of all taxons
across Crustacea do undergo reproductive senescence. With inclusion of
many species characterized by sessile, sedentary or slow motile groups,
gonochorism in crustaceans is limited to 92%; the remaining 8% employ
alternate sexuality of parthenogenic and hermaphoditism. To escape from
inbreeding, parthenogenics sporadically or regularly produce sexual males
and females, or intertwine hybridization and polyploidization. On the other
hand, hermaphrodites employ sequential hermaphroditism or androdioesic
mating system.
The fourth chapter explains that the same tissue types are involved in
regeneration of the lost part(s) of chelar appendages and asexual reproduction
of the externae in the exceptional hemocoelomate of the parasitic colonial
rhizocephalan females. They are limited to a few tissues derived from
ectoderm and mesoderm alone. Hence, the so called asexual reproduction
in these rhizocephalans simulates more of regeneration than of agametic
cloning of a whole animal.
viii Preface
The fifth chapter is concerned with cyst and their equivalents like the
ephihippia and diapausing embryos/larvae. In Artemia cyst, protein synthesis
and degradation are ceased, trehalose is used as metabolite, metabolism is
depressed down to 500,000 times and the cyst can survive anoxia for four
years. A number of environmental factors are known to trigger cyst hatching.
However, even the most potent cue does not initiate hatching in all cysts in
an egg bank. Hatching in batches is a crustacean trait. In brooded embryos,
hatching is initiated by the mother with the release of a chemical substance.
However, almost nothing is known on the mechanism, which controls cyst
hatching in different batches.
Interestingly, collections of widely scattered information reveal that
(i) the sex in crustaceans is genetically determined at fertilization and (ii)
the presence of both male and female heterogametic sex determination
mechanisms. The ostracods have explored employing numerically more X
(X-X ) chromosomes to determine the female sex and anostracans used B
1 11
chromosome to determine the male sex.
Chapter seven provides an account of sex differentiation by endocrines
and its disruption by endocrine disruptors, parasites and food. Unlike
vertebrates, endocrine and spermatogenic functions are clearly separated
into distinct androgenic glands and testes, respectively. In crustaceans, sex
differentiation process is irrevocably completed on or before metamorphosis
or birth. Juvenile hormone or its analogs induce(s) parthenogenic male
production in cladocerans. But the ability of these chemically induced males
to fertilize eggs remains doubtful. Limitation of food induces production
of males in cladocerans but females in copepods. Specific alga induces
direct development of females in protrandric hippolytids. For the first time,
cymathoid parasites are recognized and designated as protrandrics and the
other bopyrids as protogynics. Briefly, parasitism in bopyrids has retained
sexual plasticity until their cryptoniscus larvae settle on the definitive hosts
and also eliminated the reproductive senescence.
This book is a comprehensive synthesis of over 972 publications carefully
selected from widely scattered information from 249 journals and 101
other literature sources. The holistic approach and incisive analysis have
led to harvest several new findings and ideas related to reproduction and
development in crustaceans. Hopefully, this book serves as a launching pad
to further advance our knowledge on reproduction and development of
crustaceans.
August, 2015 T. J. Pandian
Madurai-625 014
Acknowledgements
It is with great pleasure, that I wish to thank Drs. N. Munuswamy and
E. Vivekanandan for critically reading parts of manuscript of this book and
offering valuable suggestion. I must thank profusely my student Prof. R.
Gadagkar for persuading me to author this book series and for offering a
token support by the Indian National Science Academy, New Delhi. Besides
my earlier review, my students Drs. C. Balasundaram, S. Katre and S. Kumari
(USA), whose publications from my laboratory, have helped me to author
this book. The Central Marine Fisheries Research Institute, (CMFRI) Kochi
provides the best library and excellent service to the visitors. I wish to place
on record my sincere thanks to Dr. A. Gopalakrishnan and his library staff.
Dr. A. Gopalakrishnan, Director, CMFRI has thoughtfully chosen to engage
and support me to accomplish this book series, for which, I remain thankful.
I also thank Drs. T. Balasubramanian R. Jeyabaskaran, P. Murugasen
and Rajakumar for regularly helping me with publications. Thanks are
due to Drs. J. A. Baeza, (USA) G. Kumarasen, S. M. Naqvi, K. K. Vijayan,
S. S. S. Sarma (Mexico) and S. C. Weeks (USA) for readily providing me
the requested publications. The manuscript of this book was prepared by
Ms C. K. Chitra Prabha, M.Sc., M.Phil. and I wish to record my grateful
appreciation for her competence, patience and excellent work.
To reproduce figures and tables from published domain I gratefully
appreciate the permissions issued by Central Marine Fisheries Research
Institute, CRC Press (Ophelia and others), International Ecology Institute
(Marine Ecology Progress Series, Diseases of Marine Organisms), Oxford and
IBH Publishers (Reproductive Biology of Invertebrates), University of New
England and CSIRO Publishers (Crustacean Parasites: Marine Parasitology).
For permission to reproduce their figures and tables, I remain indebted to
Drs. K. Altaff, C. Balasundaram, CMFRI, Z. M. Gliwics, M. Ikhwanuddin,
N. Munuswamy, K. K. C. Nair, N. Rabet, S. B. Sainath, Scripps Institute
of Oceanography, K. Tande, G. Vogt and S. C. Weeks. For advancing our
knowledge in this area by their rich contributions, I thank my fellow
scientists, whose publications are cited in this book.
August, 2015 T. J. Pandian
Madurai-625 014
