P1:OTE/PGN P2:OTE/PGN QC:OTE/PGN T1:OTE BLBK242-FM BLBK242-Beaumont January15,2010 16:44 PrinterName:YettoCome BIOTECHNOLOGY AND GENETICS IN FISHERIES AND AQUACULTURE Second Edition Biotechnology and Genetics in Fisheries and Aquaculture, Second Edition A. Beaumont, P. Boudry and K. Hoare © 2010 Andy R. Beaumont, Pierre Boudry and Kathrin Hoare. ISBN: 978-1-405-18857-9 i P1:OTE/PGN P2:OTE/PGN QC:OTE/PGN T1:OTE BLBK242-FM BLBK242-Beaumont January15,2010 16:44 PrinterName:YettoCome BIOTECHNOLOGY AND GENETICS IN FISHERIES AND AQUACULTURE Second Edition Andy Beaumont SchoolofOceanSciences BangorUniversity Wales,UK Pierre Boudry Ifremer UMRM100PhysiologieetEcophysiologiedesMollusquesMarins TechnopoledeBrest-Iroise Plouzane´,France Kathrin Hoare SchoolofOceanSciences BangorUniversity Wales,UK A John Wiley & Sons, Ltd., Publication iii P1:OTE/PGN P2:OTE/PGN QC:OTE/PGN T1:OTE BLBK242-FM BLBK242-Beaumont January15,2010 16:44 PrinterName:YettoCome Thiseditionfirstpublished2010 (cid:2)C 2010byAndyR.Beaumont,PierreBoudryandKathrinHoare Firsteditionpublished2003 BlackwellPublishingwasacquiredbyJohnWiley&SonsinFebruary2007.Blackwell’spublishing programmehasbeenmergedwithWiley’sglobalScientific,Technical,andMedicalbusinesstoform Wiley-Blackwell. 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Designationsusedbycompaniestodistinguishtheirproductsareoftenclaimedastrademarks.Allbrand namesandproductnamesusedinthisbookaretradenames,servicemarks,trademarksorregistered trademarksoftheirrespectiveowners.Thepublisherisnotassociatedwithanyproductorvendormentioned inthisbook.Thispublicationisdesignedtoprovideaccurateandauthoritativeinformationinregardtothe subjectmattercovered.Itissoldontheunderstandingthatthepublisherisnotengagedinrendering professionalservices.Ifprofessionaladviceorotherexpertassistanceisrequired,theservicesofa competentprofessionalshouldbesought. LibraryofCongressCataloging-in-PublicationData Beaumont,A.R.(AndyR.) Biotechnologyandgeneticsinfisheriesandaquaculture/AndyR. Beaumont,PierreBoudry,KathrinHoare.–2nded. p.cm. Includesbibliographicalreferencesandindex. ISBN978-1-4051-8857-9(hardback:alk.paper) 1.Fishes–Moleculargenetics. 2.Fishes–Breeding. I.Boudry,Pierre. II.Hoare,K.. III.Title. SH155.5.B432010 639.3–dc22 2009031829 AcataloguerecordforthisbookisavailablefromtheBritishLibrary. (cid:2) Setin10/12ptTimesbyAptaraR Inc.,NewDelhi,India PrintedinSingapore 1 2010 iv P1:OTE/PGN P2:OTE/PGN QC:OTE/PGN T1:OTE BLBK242-FM BLBK242-Beaumont January15,2010 16:44 PrinterName:YettoCome Contents Prefacetothesecondedition ix Acknowledgements xi Chapter1 Whatisgeneticvariation? 1 Deoxyribosenucleicacid 1 Ribosenucleicacid 5 Whatisthegeneticcode? 7 Proteinstructure 7 Sowhataboutchromosomes? 9 Howdoessexualreproductionproducevariation? 11 MitochondrialandchloroplastDNA 16 Chapter2 Howcangeneticvariationbemeasured? 19 DNAsequencevariation 19 DNAfragmentsizevariation 36 Proteinvariation 44 Phenotypicvariation 47 Chapter3 Geneticstructureinnaturalpopulations 49 Whatisapopulation? 49 Howareallelefrequenciesestimated? 51 Whatistherelationshipbetweenallelesandgenotypes? 51 Howdoallelefrequencieschangeovertime? 53 Howdoespopulationstructurearise? 54 Howaregeneticmarkersusedtostudypopulationstructure? 55 Levelsofgeneticdifferentiationinaquaticorganisms 60 Potentialproblemswithallozymesandcodingmarkers 61 mtDNAvariation 63 Microsatellitevariation 66 Populationstructureintheflatoyster 68 Mixedstockanalysis(MSA) 73 v P1:OTE/PGN P2:OTE/PGN QC:OTE/PGN T1:OTE BLBK242-FM BLBK242-Beaumont January15,2010 16:44 PrinterName:YettoCome vi Contents Chapter4 Geneticsofpopulationsizeinconservation andaquaculture 77 Geneticsofsmallpopulationsizeinthewild 78 Geneticmarkersinconservation 82 Geneticsofsmallpopulationsizeinthehatchery 85 Isthereevidenceoflossofgeneticvariationinthehatchery? 87 Howdoeshatcherypropagationaffectheterozygosity? 89 Geneticmarkersforidentificationofhatcheryproduct 93 Geneticmarkersforpathogenidentification 94 Chapter5 Geneticvariationoftraits 99 Qualitativetraits 99 Quantitativetraits 105 Whatkindsoftraitsareimportant? 106 Variationofaquantitativetrait 106 Howcanweestimatenarrow-senseheritability? 109 Realisedheritability 115 Correlatedtraits 118 Whattypesofartificialselectionsarethere? 118 Settingupabreedingprogramme 122 Inbreeding,cross-breedingandhybridisation 123 Currentstatusofselectivebreedingprogrammesinaquaculture 126 Chapter6 Fromgeneticstogenomics 129 Whatisthegenome? 129 Genomemapping 130 Wholegenomesequencing:the‘bigpicture’ 137 QTLmapping 137 ApplicationofQTLsinaquacultureandfisheriesmanagement 138 Marker-assistedselection(MAS):fromQTLstogenomicselection 139 Transcriptomics 139 Chapter7 Triploidsandbeyond:whymanipulateploidy? 145 Howisitdone? 146 Productionofgynogensandandrogens 148 Identificationofploidychange 149 ValueofTriploids 151 Tetraploids 155 Gynogensandandrogens 157 P1:OTE/PGN P2:OTE/PGN QC:OTE/PGN T1:OTE BLBK242-FM BLBK242-Beaumont January15,2010 16:44 PrinterName:YettoCome Contents vii Chapter8 Geneticengineeringinaquaculture 161 TheDNAconstruct 162 Transgenedelivery 164 Transgeneintegration 168 Detectingintegrationandexpressionofthetransgene 169 Resultsoftransgenesiseffortsinfish 171 Somuchfortransgenics–whataboutcloning? 176 Genethics 176 Glossary 181 Index 193 Colourplatesappearbetweenpages52and53 P1:OTE/PGN P2:OTE/PGN QC:OTE/PGN T1:OTE BLBK242-FM BLBK242-Beaumont January15,2010 16:44 PrinterName:YettoCome Preface to the second edition Therateofchangeinthefieldofmoleculargeneticshasbeenextremelyfastsincethe development of DNA sequencing methodologies and the polymerase chain reaction. The new millennium has seen increasing miniaturisation and sophistication of these techniques,andthishasledtosecond-generationsequencingmethodsthatcanproduce hundreds of thousands of DNA sequences from an organism in a very short space of time.Becauseofthepaceofthesetechnologicalchanges,somepartsofthefirstedition ofthisbookquicklybecameoutdated. This second edition benefits from the addition of a new author (Dr Pierre Boudry) and in updating the book to take account of new technologies, we have taken the opportunitytocorrecterrors,improveandreorganisethematerialfromthefirstedition. Several chapters retain the same headings but with new material added to bring them rightuptodate.Usingmaterialfromthefirsteditionandaddingfreshtextandfigures, we have introduced a new chapter based around the genetics of population size that links directly to problems in aquaculture and in conservation. In addition, we have providedanewchaptercalled‘Geneticstogenomics’thatcoversrecentdevelopments in genetic mapping, the search for quantitative trait loci and introduces the subject of transcriptomics–thetranscriptionofDNAsequencetoRNA. We have retained the introductory-level nature of the book but have increased the numberofreferencesattheendofeachchapterformoreadvancedreaders. ix P1:OTE/PGN P2:OTE/PGN QC:OTE/PGN T1:OTE BLBK242-FM BLBK242-Beaumont January15,2010 16:44 PrinterName:YettoCome Acknowledgements The following have kindly given us permission to use some of their material in the illustrations for the book: Dr Ann Wood, Dr Karen Abey, Dr Shelagh Malham, Dr CraigWilding,ChrisBeveridge,AlixTaylorandNatalieHorton.Wethankthescientific publishingcompaniesthathaveallowedustousefiguresfrompublishedsourcesandall suchcompaniesareacknowledgedinfigurelegends. WethankDavidRobertsfromtheSchoolofOceanSciences,BangorUniversity,for photographyandassistingwithpreparationoffigures. Dr Pierre Boudry thanks the members of the Working Group on the Application of GeneticsinFisheriesandMariculture(InternationalCouncilfortheExplorationofthe Sea)forsharingideasandknowledge. This second edition of the book would not have appeared without the support and encouragementofNigelBalmforthandthekeeneditorialassistanceofKateNuttalland CatrionaDixonofWiley-Blackwell. xi P1:SFK/UKS P2:SFK BLBK242-COLOUR-PLATE BLBK242-Beaumont December31,2009 12:20 PrinterName:YettoCome PlateB2.1 Athermalcyclerinwhichpolymerasechainreactioniscarriedout.During polymerasechainreactiontheheatedlidisclosedoverthemicro-titreplatewhichispositionedin theheatingblock. PlateB2.2 Agarosemini-gelshowingpipettingofDNAsampleintoseparatewellspositioned acrossthegel.Thegelispreparedbypouringitintoamouldandallowingittoset.Thewellsare castintothegelduringthisprocess.Samplesarecolouredtomonitortheprogressof electrophoresis. Biotechnology and Genetics in Fisheries and Aquaculture, Second Edition A. Beaumont, P. Boudry and K. Hoare © 2010 Andy R. Beaumont, Pierre Boudry and Kathrin Hoare. ISBN: 978-1-405-18857-9 1 P1:SFK/UKS P2:SFK BLBK242-COLOUR-PLATE BLBK242-Beaumont December31,2009 12:20 PrinterName:YettoCome PlateB2.3 SlabgelDNAsequencershowingtheupperandlowerbuffertrays.Theverythingel issandwichedbetweentwoglassplatesconnectedtotheupperandlowerbuffertrays.Inset showsdetailofthesharks-toothcombrestingonthetopofthegel.Upto98samplescanbe appliedindividuallyintoeachofthesmallchambersbetweentheteeth. PlateB2.4 ReadoutfromacapillarygelDNAsequencerusingtheSangermethod.Each colouredpeakrepresentsasinglebase(blue,cytosine;red,thymine;black,guanine;green, adenine)andthesequenceofbasesisgivenabovethegraphic. 2
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