Reproduction and Development in Mollusca PDF
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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 ·C o v e rT popag-Le:e Rfte pphroestoen otfa ttihvee Hexaawmaipialens mfoorn akp lsaecaol:p Rheoprrao, dmuocnedo pblya ckoinpdh ocroau, rptoeslyyp olaf cMop. Shuolrlaiv, ashnell-less opisthobranch, ·p t e r o pToodp,- Lperofts pobhroatno cohfs t,h bei vHalavweas,i isacna pmhoonpko dse aanl:d R ceepprhoadluocpeodd bs.y kind courtesy of M. Sullivan · 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 CRC Press Taylor & Francis Group CRC Press 6000 Broken Sound Parkway NW, Suite 300 Taylor & Francis Group Boca Raton, FL 33487-2742 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2017 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business © 2017 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works No claim to original U.S. Government works Printed on acid-free paper Version Date: 2200117700411179 Printed on acid-free paper Version Date: 20170119 International Standard Book Number-13: 997788--11--14398877--14014359--02 (Hardback) International Standard Book Number-13: 978-1-4987-4139-2 (Hardback) This book contains information obtained from authentic and highly regarded sources. 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Trademark Notice:Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging‑in‑Publication Data Library of Congress Cataloging‑in‑Publication Data NNaammeess: :A Plaanvda,i aJnu,a Tn .J Jo.seì, editor. TTiittllee:: TRreoppriocdalu pctinionni paendds d: ebvioel-oepcmoloengyt ,i nth mreoaltlsu, sacnad / cTo.Jn. sPearnvadtiiaonn, C/ eednittroarl, Names: Alava, Juan Joseì, editor. JMuaanr iJnoes eFì iAshlearviae,s F Raecsueltayr cohf ISncsietintucete, ,I Knsotcithui,t eK feorra ltah,e I nOdcieaa.ns and Title: Tropical pinnipeds : bio-ecology, threats, and conservation / editor, FDisehsecrriiepst,i oTnh:e B Uoncaiv Rerastiotny ,o Ff LB :r iCtiRshC CPorelussm, bTiaay, lVora n&c oFuravnecr,i sC Ganroaudpa., 2017. | Juan Joseì Alava, Faculty of Science, Institute for the Oceans and DSeesrciersip: Stieornie: sB oonca r eRpartoodnu, FctLio : nC aRnCd Pdreevsesl,o 2p0m17e.n |t “ Ain sacqieunactiec pinuvbelirstheberrsa tbeos o;k.” Fisheries, The University of British Columbia, Vancouver, Canada. |v Ionlculmudee 2s b| i“bAl iSocgireanpchei cPaulb rleisfehreerns cbeos oakn.”d index. Description: Boca Raton, FL : CRC Press, 2017. | “A science publishers book.” I|| dI Iennncctluliufdideeersss b: bLibiCblilCoiogNgrr a2ap0ph1hi6icc0aa5ll 4 rr1eef7fe0err|e eInnSccBeeNss a a9nn7dd8 i1in4nd9de8ex7x.4e1s.392 (hardback : alk. paper) | IIIdSdBeenNntt i9iffii7ee8rr1ss:4: L9LC8C7CC4N1N4 2 020801 1(6e70-0b50o47o147k00 :7 |a| I lISkSB. BpNNa p9 9e77r8)811419388774110349520 ( h(haardrdbbacakck : )a |l k . paper) | SIISuSBbBNjNe c9 9t7s78: 81L14C39S18H5724: 01S04e60a38ls7 ( (e(A-eb-nboioomoka k:l )sa)l-k-T. proappeicrs). | Pinnipedia. CSSuluabbsjesjeicfctitscs:a :L tLiCoCnSSH: HL: C:S MCea oQlsll Lu(A7sk3n7si-.mP-R6a elTsp)7r-6o-T d2ru0oc1pt7ii co|s nD. .| D| P MCin o5nl9li9up.se7kd9si--a--.dDce2v3elopment. LCCClla asrsessiciffioicrcadat tiaioovnna:i: lLLaCbClCCe aQQt LhL7t43t0p73.sP :./P6/ l3Tc6c7 n26.0 l2o107c1. g|7 o D|v DD/2DC0 1C56 950495-94-.d177c920-3-dc23 LLCC r reeccoorrdd a avvaaiillaabbllee a att h httttppss::////lclcccnn.l.loocc.g.goovv//22001167005047147007 Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com Visit the Taylor & Francis Web site at ahnttdp t:/h/we CwRwC.t aPyrleosrsa Wndefbra snitcei sa.tcom http://www.crcpress.com and the CRC Press Web site at http://www.crcpress.com 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
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