Reproduction and Development in Mollusca

304 Pages·2018·5.25 MB·English

by  Pandian

Series on
Reproduction and Development in Aquatic
Invertebrates
Volume 2
Reproduction and
Development in Mollusca

Series on
Reproduction and Development in Aquatic
Invertebrates
Volume 2
Reproduction and
Development in Mollusca
T. J. Pandian
Central Marine Fisheries Research Institute
Kochi, Kerala
India
p,
p,
A SCIENCE PUBLISHERS BOOK
A SCIENCE PUBLISHERS BOOK
Cover Acknowledgement
Cover Acknowledgement
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·         Top-right photo of the Galapagos fur seal: Reproduced by kind courtesy of J.J. Alava
·         Top-right photo of the Galapagos fur seal: Reproduced by kind courtesy of J.J. Alava
·         Bottom-left photo of the Juan Fernandez fur seal: Reproduced by kind courtesy of L.P. Osman
·         Bottom-left photo of the Juan Fernandez fur seal: Reproduced by kind courtesy of L.P. Osman
·  Bottom-right photo of the Mediterranean monk seal: Reproduced by kind courtesy of A.A. Karamanlidis/MOm
·  Bottom-right photo of the Mediterranean monk seal: Reproduced by kind courtesy of A.A. Karamanlidis/MOm
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NNaammeess: :A Plaanvda,i aJnu,a Tn .J Jo.seì, editor.
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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 1950s, the multi-
volume series by L. E. Hyman accumulated bits and pieces of information
on reproduction and development of aquatic invertebrates. Through a few
volumes published during 1960s, A. C. Giese and A. S. Pearse provided a
shape to the subject of Aquatic Invertebrate Reproduction. Approached
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 in large numbers are appearing in 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 explanation 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
Molluscs are unique for the presence of a protective external shell, defensive
inking, distribution from the depth of 9,050 m to an altitude of 4,300 m, gamete
diversity, the use of nurse eggs/embryos to accelerate the first few mitotic
divisions in embryos, the occurrence of natural androgenics, gigantism
induced by elevated polyploidy, the complementary role of mitochondrial
genome in sex determination by nuclear genes and the uptake of steroids
from surrounding water. Not surprisingly, many books were authored or
edited on one or other aspect of molluscs. This book represents perhaps the
first attempt to comprehensively project the uniqueness of molluscs, covering
almost all aspects of reproduction and development from aplacophorans to
vampyromorphic cephalopods.
The book is organized in nine chapters. Considering their uniqueness,
three chapters are devoted to shell and reproduction, shell and acidification,
and aestivation. In view of its importance, regeneration is woven with the
claimed asexual reproduction. Molluscan strategy is to carry shell(s) to escape
from predators, limit energy expense on motility, and allocate the thus saved
energy on reproductive output. Being proliferative breeders, molluscs inject
a huge volume of fodder into the trophic dynamics, contribute 22 mt to the
global food basket, colonize thermal vents and cold seeps by successfully
engaging endosymbionts; whereas the adults move at ‘a snail pace’, their
larvae embark on trans-Atlantic dispersal; they also serve as intermediary
hosts to the dreadful disease causing schistosomes. For the first time, the
second chapter attempts to relate shell thickness to reproductive output,
and chirality to mating and insemination between dextrals and sinistrals.
With increasing size/age, the allocation for shell growth is progressively
diminished but with increasing reproductive output. At low temperatures
and higher altitudes, energy cost of calcification of shell increases. The shell
size, shape and thickness are phenotypic traits. Like other animals, the 2%
shell-less molluscs depend on chemical defense; aplysiaids and cephalopods
are known for inking and escape.
From the fairly long third chapter, it has become apparent that, the
presence of shell in molluscs has afforded iteroparity and comparatively
longer life span in prosobranchs and bivalves but its absence semelparity
and short life span in opisthobranchs and cephalopods. Notably, > 25%
viii Reproduction and Development in Mollusca
of molluscs is hermaphordites. In shell-less molluscs, gonochorism has
facilitated faster growth and relatively larger body size in cephalopods but
hermaphroditism small body size in opisthobranchs. Gastropods display
great powers of regeneration inclusive of ‘brain’ and nervous system. The
claimed asexual reproduction in the pteropod Clio pyramidata is more of
regeneration involving multi-potent stem cells. The fifth chapter recognizes
the occurrence of seasonal aestivation in snails and tidal aestivation in
bivalves. For the first time, the effect of varying durations of aestivation on
growth and reproduction in a freshwater snail and supratidal inhabitant
bivalves are comprehensively described. For the first time, the scope for
employing aestivating, schistosome-infected snails as ‘biological weapon’ is
also explored.
The sixth chapter commences with description of spawning induction and
gamete preservation, indicates the presence of heterogamety in less than a
dozen gastropods and bivalves, and reports the rare occurrence of androgenics
in bivalves; the induced triploid and tetraploid bivalves grow many times
faster than their counterpart diploids, produce more and tastier meat with
higher glycogen content and are available in markets during off season. The
unique but perhaps ‘vestigial’ presence of Doubly Uniparental Inheritance
(DUI) in bivalves and its possible causative complementary role of sperm
mitochondrial genes in sex determination by nuclear genes is brought to
light. Irrespective of sexuality in molluscs, sex is irrevocably determined by a
few genes harbored on autosomes. Sex differentiation is solely accomplished
by neurohormones. The genes responsible for biosynthesis of vertebrate-
type steroid hormones are indicated to be missing in molluscan genome.
Surprisingly, molluscs absorb and accumulate steroid hormones from
surrounding waters. The ongoing debate on the role played by the steroid
hormones in regulation of reproductive cycle is discussed. While tributyltin
induces sex reversal from female to male in fishes, it only superimposes
masculine traits on female neogastropods. Trematode parasites exploit many
molluscs as intermediate host and partially or completely sterilize them.
Sex ratio represents the cumulative end product of sex determination and
differentiation processes. In molluscs, sex ratio of 1♀ : 1♂ is maintained at
population level. Among different families of a molluscan species, sex ratio
varies from nearly all females to nearly all males. But the ratio remains
constant between successive broods in a family. Hence, the ratio is more a
family trait than that of species. The primary sex differentiation is determinate
and not amenable to environmental influence. However, the secondary
differentiation from male to female in protandrics is labile, protracted and
amenable to environmental factors like food supply and temperature.
The eighth chapter highlights the effects of acidification on molluscan
shell. Since the advent of industrial era, the oceans have absorbed 127 billion
metric tons of CO, resulting in decreased pH, which renders the acquisition
2
of calcium costlier. Consequently, the molluscan shells suffer from pitting,
dissolution and loosening of the hinge to valve mechanism. However, with
Preface ix
abundant food supply, many molluscs do adapt and not suffer from shell
dissolution.
This book is a comprehensive synthesis of over 907 publications carefully
selected from widely scattered information from 249 journals and 87 other
literature sources. The holistic and incisive analyses have led to harvest
several new findings related to reproduction and development in molluscs
and to project the uniqueness of mollusca. Hopefully, this book serves as
a launching pad to further advance our knowledge on reproduction and
development of molluscs.
September, 2016 T. J. Pandian
Madurai, 625014