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LectureNotesonCompositeFishCultureanditsExtensioninIndia

Producedby:Fisheriesand
AquacultureDepartment
Title:LecturenotesoncompositefishcultureanditsextensioninIndia...
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GENETICSELECTIONANDCARPHYBRIDIZATION
H.A.Khan
Freshwater
Aquaculture
Researchand
TrainingCentre
(CIFRI),Dhauli
P.O.Kausalyagang
ViaBhubaneswar2
Orissa(India)

1INTRODUCTION
Themainpurposeoffishselectionistoimprovetheexistinganddevelopnewbreedsandhybrids
andthusincreasetheirproductivity.Forimprovingthequalityofcommerciallyimportantfish
species,theuseismadeoftheirvariationinmanymorphological,physiologicalandbiochemical
features.Amarkedproprotionofthisvariationishereditary,thelevelofwhichisveryhighinfish
populationandhelpsinfishselectionwork.
Incomparisontothebreedingofdomesticanimalsfishcultureisayoungscience.Fishculturein
ChinaandIndiahasbeeninvoguesincelong.Butthedomesticationoffishandcreationof
breedsdifferingfromtheirwildparentsinhigherproductivitytraitswasactuallystartedonlyafew
centuriesago.Withthenotableexceptionofgoldfish,ornamentalcarpandperhapsthecommon
carp,fewfishcouldbeconsidereddomesticatedeventhoughsomestrainsoftrout,forexample,
aremuchmoreadaptedtohatcheryconditionsthantheirwildcounterparts.Otherspecieslikethe
Chinesecarps,Indiancarps,Tilapiasp.andChannelcatfisharebecomingdomesticated.
Ofallculturablecarps,onlycommoncarphasbeenbredforsufficientlylongtimeanddistinct
breedsofthisspeciesthroughselectiondevelopedare:InUSSRtheUkrainiancarp,Ropsha
carp,hybridsofthefirstgenerationofthedomesticatedcarpandtheAmurWildcarp,theNivchan
carp,theCentralRussiancarp,theKazakhistancarp,theKasnodarcarp,theByelorussianbreed
andtheParrabreedofcarp(Kirpichnikov,1981)inIsraelDor70carp(Wohlfarth,Lahman&
Hulata,1980)andinHungarytheHungarianstrain(Bakos,1979).

2METHODSOFFISHSELECTION
2.1Massorindividualselection
Thetermmassofindividualselectionisusedtodescribeartificialselectionandusein
subsequentreproductionofindividualshavingthebestphenotypes.Thetraitsusedinsucha
selectiondependonspecificpurposeandmayincludeincreasedweightorbodysize,good
exteriorindices,rateofsexualmaturation,thenecessarypigmentation,thedesignpatternof
scaling,resistancetounfavourableenvironmentalconditionsandtodiseasesandcertain
improvedphysiologicalorbiochemicalcharacteristicseasilymeasurableinfishes.Selectionmay
includeinteriorfeaturessuchasnumberofintramuscularbones,sizeofairbladder,etc.
Theefficiencyofmassselectionisexpressedbysimpleequation(Falconer,1960).
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R = ih2=Sh2(1)
Where
R = hereditaryimprovementofcharacterofonegeneration
S
Selectivedifferential(differencebetweenmeanforselectedindividualsandinitial
=
populationmean)
intensityofselectioni.e.selectivedifferentialexpressedinmeansquare
i =
deviations

h2 = heritabilityofcharacter
= variabilityofinitialpopulationexpressedintermsofstandarddeviation.

Nowletusconsiderthepossibilityofincreasingeachofthetermsintherightpartoftheabove
equation,whileperformingtheselectionofafish.
2.1.1Theintensityofselection
Theintensityofselectionmaybequitesubstantialinthosespecieswhichareveryfertile.Insuch
casewecanchooseasmallpartofagrownpopulationforbreedingandrestcanberejected.The
numberofbestspecimensleftforbreedingiscalledtheselectionseveritycoefficient(V)

WhereN=initialnumberofindividuals
n=numberofselectedindividuals
Selectionseveritycoefficientinveryfertilefishlikecommoncarpcanbeeasilybroughtto1%
(1:100)oreven0.1(1:1000).OccasionallyinselectingRopshacarpinfirstyear,stillmorerigid
normsofrejection,keepingforbreedingpurposeonly0.050.02%ofthepopulation(1:2000to
1:5000)havebeenfollowed(Kirpichnikov,1967).Suchanincreaseintheselectionseverity
requiresaverylargenumberofpopulationstobecultured.Theseverityandintensityofselection
arefunctionallycorrelated(Fig.I).AtlowvaluesofV(0.01orless)anyfurtherdecreaseinthose
characteristicshardlyhasanyeffectonthevalueofselectionintensityanddoesnotcompensate
fortheexpenditurenecessarytogrowalargenumberoffishespriortothetimeofselection.Ifthe
fertilityoffishislow,selectionwithratio1:20(V=5%)wouldbepermissible.Afurtherreduction
inseverityisundesirableasitwouldbefollowedbyasuddendropofthevaluei.Withthe
increasedsizeofinitialpopulationandthereforeenhancedimaybeaccompaniedbyreductionin
heritability(h2).Ithasbeenobservedbymanyinvestigatorsthatinyoungstagesofcommoncarp,
whengrown,underconditionofhighdensity,theweightdistributioncurvebecomesmarkedly
asymetricafewshootfryalsocalledjumpersorchampionsappearwhoserateofgrowth
greatlysurpassesthatoftheircompanionsofthesameagethefoodcompetitionbeingthemean
reasonforthissizedifference(Wohlfarth,1977).Inpresenceoffoodcompetition,anincreasein
severityandintensityofselectionaboveacertainlimitwillhaveabadeffect.
2.1.2Variabilityofinitialpopulation
Thevariabilityofinitialpopulation()mustbesufficientlyhighifVisnearzero,evenwithavery
highintensityofselectionandconsiderableheritability(about0.60.8)theefficiencyofselection
mayturnouttobelow.Inmassselection,onlygeneticvariationisimportantbecauseany
increaseinthevalueofnonheriditaryvariablesduetotheincreaseinenvironmentalvariationis
meaninglessbecauseitwillresultinaproportionaldecreaseinheritabilityandwillnotenhance
theefficiencyoftheselection.
2.1.3Heritability
Heritability,inbroadsense,istherelationbetweenthewholegeneticvarianceandtotalvariance:

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Totalvariance,inturn,expandsintoanumberofcomponents.

WhereEisenvironmentalvarianceandjisvarianceofinteractionbetweenenvironmentand
genotype.TheE2andj2shouldbereducedandthiscanbedonebycreatingsimilar
homogenousconditionsforgrowing(tobeginwithparentmaintenanceconditions)andreducing
foodcompetitionwhichfrequentlyenhancestheinteractionalvariance.
Heritabilitycanbeincreasedinmanyways:
i. Bycrossingunrelatedindividuals,includingcrossingbetweenlines,breedingstocks,
varieties,subspeciesandoccasionallyspecies
ii. Bythemethodofunifyingfishgrowingconditionstoselection.
Infishculture,theparatypicvariancecanbegreatlyreducedbyadoptingthefollowingmeasures,
theimportantonesare:
i. settingupofsimilarconditionsforallparentspriortoreproduction,
ii. simultaneousperformanceofallcrossesdesignedtoyieldmaterialforselection,
iii. maximalstanderdisationofenvirenmentalconditionsthroughouttheperiodoffishculture,
iv. stockingoffishessimultaneouslyintopondsatamoderatedensityinordertoavoid
excessivefoodcompetition,
v. measurestopreventmixingofindividualsculturedindifferentponds,
vi. selectiontobeconductedattheagemaximallycorrespondingtothemarketingage.
However,inspiteofobservingallabovementionedconditions,theenvironmentalvariance(E2)in
commoncarpandinseveralotherfishesprovestobeveryconsiderable.Acompleteelimination
ofcausesthatproduceunequalconditionsoflifeforindividualslivingtogetherinpondsis
impossible.
Theinteractionbetweenthegenotypeandageinfishisnotveryimportantbuteventhenitexists.
Theearlyphasesoflifeoffisharegreatlyinfluencedbymaternaleffect.Theenvironmental
componentoftotalvarianceofweightisparticularlyhighatthebeginning,butdecreases
thereafterandheritabilityofweightandsizeincreasesbyafactorof23.Geneticdifferencesare
moredetectableatlaterstagethanduringtheearlyphasesoflife.Atlaterstagefishgrowthis,
however,greatlyaffectedbytherateofgonadalmaturation,sowhenselectionisaimedatweight,
itshouldbedoneatthemiddleage.
Occasionally,heritabilitydecreasesbecauseofheterozygousbalance,aphenomenonpeculiarto
manyspeciesofanimalsandplants.Duetogreatnumberofchromosomesincommoncarp(2n=
104)andmanyotherfoodfishtheprobabilityofheterozygousbalancebeingestablishedina
populationappearstobegreat.Hence,itistheindividualswithmaximumheterozygosisthatwill
bethemostviableandfastgrowinginapopulation.
Selectionundersuchconditionsverysoonbecomesinefficient,sincethegeneticvariabilityproves
tobeinthemainnonadditive.Astovariabilityofanycharacter,itistheproportionofadditive
geneticvariabilityinthetotalvariabilityofthegivencharacter.

where
A2=G2D2E2(6)
Withoutbalancedheterogygosis,thevarianceofdominanceandepistasis(D2and2)aresmall
andinitiallytheadditivegeneticvariancedoesnotdiffermuchfromthetotalvariance(G2).If
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thereissubstantialheterozygousbalanceformanypolygenes,thedifferencebetweenh12and
h22willbeverygreat.Atrueheritabilityinspiteofconsiderablegeneticvariability(G2)willbe
nearzero,sincethevariabilityisnotadditive.
Itisdifficulttostriveagainstanestablishedheterozygousbalanceastheequilibriumofgenotype
provestobeverystable.Theonlywaytocorrect,thesituationistoperformnew,sufficiently
remotecrossingswhichmaydestroythepolygenesystemandtherebycontributetoanincrease
inheritability.Table1showsthephenotypicandgeneticparametersforquantitativecharactersin
commoncarp.
Table1.Phenotypicandgeneticparametersforquantitativecharacterincommoncarp(
=mean,standarddeviation),CV=CoefficientofVariation,S.E.=Standard
error,Heritability(h2)asestimatedfromsire(S),dam(D)andfamily(F)
Componentsorvariation
Traits
Weightoffingerlings
Bodyweight
Drymatter
Fatcontent
Nindrymatter
4monthwt.(g)

Tolerancetohypoxia
Weightgain(g)

CV h2s S.E. h2DS.E Nooffamilies

h2fS.E

0.100.20
0.25
0.150.18
0.140.15
0.150.17
71 17 23

179 50 28
366 81 22 0.47(b)

0.48F
0.12F
0.15F

9F
9F
9F
17off/parents

Authors
Kirpichnikov(1972)
Smisek(1979)
do
do
do
Nagyetal
(1980)
do
Brodyetal.(1981)

Fromtheabovetableitisevidentthatweightheritabilityincommoncarpforjuvenilesisratherlow
andthatforbodyweightofadultsisofmediumsizeandhigherforyounganimals.Selectionfor
weightisalwayspossible,butitdoesnotalwaysgivethedesiredresults.Inthepresenceoffood
competition,moreaggressiveindividuals,whoareabletosnatchfoodfromothersinthepond
(Moav&Wohlfarth,1967)maynotnecessarilybethebestatassimilatingfood.Itisevidentthat
forimprovingtherateofgrowthmassselectionshouldbecombinedwithtestingforrelatives.
Massselectionshouldbereplacedbytestingforrelativesonlyincaseswhensuchcharacteristics
asfatcontent,biochemicalcompositionofmeatanddegreeofboninessaretobeknown,orifthe
principaltaskistoraisethefishproductionofponds.

2.2TheSelectionforRelatives
Theselectionforrelativesinvolvestoagreatextenttheselectionforgenotypesthepositive
characteristicsofindividualschosenforsubsequentreproductionareknownfromananalysisof
theircloserelatives.Therearetwoformsofselectionforrelatives:(a)familyselectionand(b)
evaluationofspawnersbyprogenytesting.Asinmassselection,theefficiencyofselectionfor
relativeisexpressedbyasimpleequation.
R=if.fhf2(7)
Inthisequationintensityofselection(if)equalsthedifferencebetweenmeanforselectedfamily
andpopulationmeanexpressedinmeansquaredeviations,ofcharacterisesthevariabilityof
familymeansandh2showsheritabilityofdifferencesbetweenfamilyeans.Infamilyselectionthe
valuefandselectionintensityarereducedthanthesevaluesinmassselection,butthe
heritabilitymeanscanreachveryhighvaluesifthefamilitiesaregrowntogetherundersimilar
conditions.
2.2.1Familyselection
Familyselectionisgenerallyusedwhenhereditaryofselectedcharacteristicislow.Thisrequires
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farmingofseveralfamiliesoroffspringsfromdifferentpairsorsmallpotentialgroupsunder
optimumconditions.Thememberofcrossingsmaybe810orwhenculturefacilitiesexist,itmay
beashighas15or20.
Theselectionintensity(f)dependstoaverygreatextent,onthenumberoffamilies.The
variabilityofthefamilymeans(f)isnotgreatandcanbeincreasedonlyattheexpenditureof
enlargingitsgeneticcomponent.Theheritabilityofmeans(hf2)isincreasedanditapproaches
whenallfamiliesarekeptunderstandardconditions.Themostimportantconditionsforfamily
selectionare:
i. Sufficientgeneticvariabilityoffamilymean
ii. Homogenityofconditionsforgrowingseparatefamiliesorsufficientreplicatesof
experiments.
2.2.2Progenytesting
Thesimplestmethodinvolvescomparisonofoffspringsobtainedfromdifferentpairsornestsof
parents(Fig.2a)inthiscasetheevaluationreferstothecombinationofindividuals.Inthe
simplifieddiallelecrosses(Fig.2b)whicharegenerallypractised,malesorfemalesareseparately
crossedwithoneormoreindividualsoftheoppositesex.Thematuringofeachofthemales
testedwithtwofemalesprovidesasufficientlyreliableevaluationofthebreedingqualitiesofthese
males.Acompletediallelecross(22,33,55,1010,etc.)alsoenablesthefishbreederto
selectthebestindividualsbelongingtoothersex,sincethenumberofoffspringincreasesits
proportiontobesquarenumberofparentsofonesexsubjectedtoexamination(Fig.2c),onehas
tofacetheproblemofgrowinglargenumberofoffspringsunderuniformstandardconditions.The
progenytestingisalsotimeconsumingandrequiresoneortwoyears.
TestingofspawnersofeithersexofcommoncarphasbeendoneforselectioninUSSR
(Kirpichnikov,1966)andinIsrael(Moav&Wohlfarth,1967)thatabout70%ofthedifferences
betweenprogenymeanscanbeattributedtogeneticvariance.Thusprogenytestsprovidean
effectivetoolforidentifyinggeneticallysuperiorprogeniesandparentalgroups.Theconditions
andrequirementsforprogenytestingaresimilartothoseusedinfamilyselection.

2.3CombinedSelection
ComparisionoftheequationR=Sh2andRf=Sf,h2fallowsonetodeterminewhichofthe
selectionmethodsisbetterforfishbreeding,ifSh2>Sf,h2fthenmassselectionistobepreferred
overfamilyselectionorviceversa.Selectionforrelativeswillbemoreeffectivefortraitswithvery
lowheritabilityi.e.lessthan0.5andwhenh2ishigherthan0.5,massselectionismoreefficient
thanthefamilyselection.Whenh2=0.5,familyandindividualselectionareofequalefficiency
(Falconer,1960).Withfish,selectionshouldbebasedonacombinationofmassandfamily
selection.Massselectionisonlyofinterestwhengrowthrateistheonlytraitofeconomic
importanceandishighlyheritable(Gjedrem,1963).Theresponseofcombinedselectionis
theoreticallyequaltothesumtotalresponseofeachoftheselectionmethodsused.
Rs=Rf+Rm+Rpr,whereRf,RmRPrrefertotheeffectivenessoffamilyselection,mass
selectionandprogenyselection.
Thefirststepincombinedselectionconsistsofcrossesbetweenheterogenousunrelatedparents,
suchcrossesareaimedatobtainingasmallnumberofprogenyupto10incommoncarp
breeding.Duringthecultivationofthesefamiliestheirreproductivepropertiesareevaluated.
Thesepropertiesincludeviability,growthrate,thequalityofflesh,etc.sothatbestfamiliescanbe
selected.Thesecondstageincludesmassselectioninseveralofthebestfamilies.Atthethird
stageparentsareexamined,usingprogenytestingParentsofjustonesexwheretheonsetof
maturityoccursearlieraretested(malesincommoncarpbreeding).Thistestingistobe
completedbythetimeofonsetofmaturityofindividualsoftheothersex.

3NEWTRENDSINFISHSELECTION
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3.1Applicationofgeneticmarkerinfishselection.
Theinformationregardingthepatternofinheritanceofmorphological(bothquantitativeand
qualitative),physiologicalandbiochemicaltraits,particularlythoserelatedtoyieldcapacitycanbe
directlyutilizedinplanningselectionwork.Thelaterinformationhelpsindecidingwhethertogo
formassselectionorselectionofrelativesandisusefulindevelopingasystemofcrosses.Agood
exampleofsuchanapplicationoftheinformationcanbeseeninthepurificationofthebrood
stockoftheRopshacommoncarpfromtherecessivegene.
InthestudyonhomoandhetorozygosityinthescaledcarpinthegeneS,(mirrorandscattered
scale),thebackcrossingiscarriedoutthus:
1)

2)

S(S?)
scaled
S(s?)
scaled

nnssnn
scattered
nnSsnn
scaledheterozygous

Intheformercase,thescaledandscatteredoffspringoftheheterozygousparentsareproduced
ina1:1ratio.Inthelattercase,theratiois3:1(threescaledperonescattered).Theentire
offspringofhomozygousscaledparentsinanyofthetwocrossingshaveacompletelyscaled
integument.From1956to1964,469parentsofRopshacarpwereexaminedbythistechnique,
247werehomozygous.Thisworkenabledcompleteeliminationoftheoccurrenceofthescattered
carpasearlyasinthe5thselectedgeneration(Kirpichnikov,1971).
Useofgenesformarkingthebreedingstocksisanotherimportantpromisingtrendandpresently
geneticmarkershavebeensuccessfullyusedforlabellingdifferentstocksofcommoncarpe.g.,
selectionofCentralRussiancarpincludingthebreedingoftwostocksdifferingintheS(patternof
scale)andD(pigmentationofspinesandhead)loci(Kirpichnikov,1981)andtwostrainsof
commoncarpinIsraelmarkedwithcolourrecessivegene(MoavandWohlfarth,1967).Gene
markersarehelpfulinestablishingtheoriginofcertainfishbreeds.Ifthecorrelationbetween
markergenesandselectivetraitissignificant,theselectioninvolvingsuchmarkersmaybe
considerablyaccelerated.

3.2Artificialgynogenesis
Gynogenesisisaspecialtypeofsexualreproductionrequiringinseminationwhennucleusof
sperm,whichhaspenetratedtheovum,undergoesinactivationintheeggplasmanddevelopment
ofembryoiscontrolledexclusivelybymaternalnucleus.Thechromosomesofspermsare
eliminatedsoonafterfertilization.Thetechniqueofartificialgynogenesisisbasedoninactivation
ofspermbyirradiationanddiploidizationofthefemalechromosomesetbyacoldshock.This
methodhasbeenemployedsuccessfullyonCyprinuscarpio(Golovinskaya,1968)andintwo
Indianmajorcarpsviz.,CatlacatlaandLabeorohita(GeorgeJohnetal.,1984).Alsothe
productionofmonosexbroodsforpopulationcontrolhasbeenattemptedwithgrasscarp.
Thegreatadvantageofthistechniqueisthatitconstitutesadefactovegetativereproductionthat
canmaintainandmultiplyasinglesuperiorgenotyperegardlessofitslevelofheterozygosity.
Anotheradvantageisthatgynogeneticallyreproducingpopulationstendtohaveonlythefemale
sex.Thisprovidesmeansofselectingfemaleswithhighergenetictendencyforgynogenesis.

3.3Polyploidy
Infishesnaturalgynogenesisisintimatelyassociatedwiththephenomenonofhybridization,
apomxisandpolyploidy.Triploidamieoticparthenogenesisandrogenesismayoccurasaresultof
backcrossesofparthenogeneticdiploidfemaleswithmalesofrelatedbisexualspecies.Incaseof
thecrossbetweentriploidgynogeneticfemaleswithdiploidmales,tetraploidgynogenetic
individualsmaydevelop.Triploidshavebeenproducedbycoldshocktreatmentofthefertilized
eggsofcommoncarp(Garvaietal.,1980).Thesefishesareexpectedtobesterile.The
integgenerichybridsbetweengrasscarpandeithercommoncarporbigheadcarpappeartobe
largelytriploid(AllenandStanley,1981).ThepopulationofCarassiusauratusglibelioinJapan
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hasbeenreportedtocontaintetraploids.

4CROSSINGASAMETHODOFINCREASINGTHEHETEROGENITY
OFTHESELECTIONMATERIAL
Theselectionresponsedoes,toalargeextent,dependonthelevelofheterogenityofthe
selectedgroups.Crossesbetweenunrelatedindividualsenrichthestrainincreasingthegenetic
componentoftheirveriationandtherebyfacilitatingselection.Anotherresultisthedisappearance
ofanyharmfuleffectofinbreeding.Crossingalsoensuresthepreservationandperfectionofthe
reproductivequalitiesofthebreedandallowsheterosistobeutilizedtoitsutmostinevery
generation.Crossinghelpsinimprovingthebreedingqualityofthelocalbreedbymakinguseof
thefewvaluabletraitsofanotherbeed(improver)andinincreasingtheviabilityofthebreedby
introducinggenesresponsiblefortheresistancetoenvironmentalfactorsanddiseases.In
accordancewiththeaboveobjectives,followingtheoriginalcrossing,thereproduction,ofthe
hybrid,populationiscarriedoutbymeansofthereproductive,introductory,absorptiveoralternate
crossing.

4.1Thereproductivecross(Fig.3a)
Thisisusedwhenmanyusefultraitsaretobecombinedfrombothcrossbreedsorspecies.This
iseasilyachievedwhenhybridsarecompletelyfertilebutrequirethoroughselectioninallhybrid
generations.Examplesofsuchcrossesare:TheUkrainianandRopshacommoncarpbreeds,the
Hungariancarp,etc.

4.2Theintroductorycross(Fig.3b):
Thisisusedwhenitisrequiredtointroduceoneorseveralvaluabletraitsofanotherstrainor
speciesintothelocalhighlyproductivebreed.F1hybridsoftwoformsarethenbackcrossed
manytimeswithindividualsofthelocalbreed,whoseimprovementsintended.Inthisprocess,the
backcrosshybridspossessingthedesiredtraitsofdonorstrainareusedforsubsequent
reproductiontoimprovethebreed.Ifthesetraitsaredeterminedbydominantclearlymanifested
genes,theproblemofconservationofrequiredcharacterscanbesolvedrelativelyeasily,
otherwiseincaseofrecessivegenesorwheninheritanceispolygenic,theriskoflossof
charactersisveryhigh.

4.3Theabsorbingcross(Fig.3c)
Similartointroductorycross.Aseriesofbackcrossesiscompletedaftertheinitialcrossoftwo
strainsbuthybridsarerepeatedlycrossedwiththeindividualsofstrainsusedforimprovement
andnotoflocalstrain.Strictmeasuresshouldbeadoptedaimingatconservingthemostuseful
traitsofabsorbedstrains,inthiscasethelocalone.

4.4Thealternativecrosses(Fig.3d)
Thisrequiresintermittentcrossingofhybridswiththeindividualsbelongingtothetwoinitialbreeds
asfollowedbyselectionofthenecessarycombinationoftraits.After3or4generations,the
alternatecrossisreplacedbyreproducingone,otherwise,itisdifficulttostabilizethetraitsofthe
newhybridbreed.

4.5Hybridization
F1hybridsiftheypossessheterosiscanbeculturedcommercially(Moav&Wohlfarth,1967).
Heterosisappearstodependontwomaincompensatorymechanisms.Thecombinationofuseful
dominantgenesaccumulatedbybothcrossedformsinhybrids(hypothesisordominance)andthe
increaseinhybridsofthetotallevelofheterozygosity(thehypothesisofoverdominance).An
increaseinthebiochemicalversalityinhybridsoccursinbothcases.Heterosisinnatural
populationisgenerallymanifestedasariseinthefitnessofhybridsintheelevationoftheadaptive
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value.Thisistypicalofmanyintraspecificandcertaininterspecificfishcrosses.Maximalcareis
requiredincarryingoutcommercialhybridization,becauseifF1hybridsareleftinthewater
bodies,therearechancesofcontaminationofthebroodstockoftheinitialforms.Thisleadstothe
deteriorationoftheeconomicallyimportantstrains.Anexampleofsuchhybridizationcanbecited
incaseofdomesticcommoncarp,withitslivingancestor,thewildcarp.Bettercontrolof
commercialhybridizationcanbeconsideredbytheuseofgeneticmarkers,bothparentsand
hybridsmaydifferintheallelesofthegenesresponsibleforthecolourpattern,scalepatternand
certainbiochemicalloci.Somepromisingimportanthybridcombinationsare:(1)Interbreed
hybridsofcommoncarpcrossesbetweenRopshaand'Ukrainiancarps,ofthescaledand
formed,Ukrainiancarps,threestocksoftheKrasnodarcarp.Heterosismanifestedinbetter
survivalandproductivityhasbeenobservedinthecrossesofHungarianandPolishcarps,and
crossesoftheJapaneseYamatocarpwithEuropeanmirrorcarp.(2)Intraspecifichybridsofsilver
carpofAmurandChineseorigin(3)HybridsofdomesticcarpandAmurwildcarp(Cyprinus
carpiohaemalopterus)(4)Intergenerichybridsofcommoncarpandcruciancarp(5)Intergeneric
hybridsofsilvercarpandbighead(6)IntergenerichybridsofCatlacatlaandLabeorohita
(Chaudhuri,1971)andmanyothersspeciesasshowninTable2

5BREEDINGSYSTEMSADOPTEDFORFISHSELECTIONINSOME
COUNTRIES
Inordertoutilizecompletelytheadvantagesassociatedwithheterogenouscrossings,fish
breedingshouldbecarriedoutaccordingtoadefiniteplan,dependingontheknowledgeof
genetics,inbreddepressionandheterosisinfishcrossing.Inmakingcrosses,inducedbreedingof
thespeciesfollowedbyartificialfecundationisemployed.Alsoforcarryingoutgenetic
experimentationonlargescale,anadequatemethodofmarkingfishforindividualorgroup
observationsistobedeveloped.Thebestresultshavebeenobtainedinmarkingexperimental
fishbyfinclippingorbrandmarking(Moavetal.,1960)orsubcutaneousinjectionsoforganic
dyes(dichlorotriazineandothercompounds)(ZonovaandKirpichnikov,1971)orbyinjectingblue,
redoryellowflurescentgranulesintothespine(Smithermanetal.,1983).UnderIndian
conditions,commoncarpwhoninjectedwithProcianMbluestainbytheauthorhasretained
marksforlasttenmonthsandstillthemarksarequitedistinct.
Thevariousfishbreedingsystemsfollowedforthegeneticimprovementofcommoncarpin
differentcountriesaredescribedasfollows:
USSR:Themethodofparallelbreedingoftwoormoregroupsisusedfortheselectionof
Ropshacarp(Kirpichnikov,19711971).Inthissystemtwoorthreegroupsare
concurrentlychosenwithinabreed,withoutintermingling,allowinginsideeacha
moderateinbreedingandcarryingoutselectionineachgeneration.Forcommercial
purposesfishfromdifferentgroupsarecrossedtoavoidcloseinbreeding(Fig.4).
Israel:
(a)Breedingingroupswithfamilyselection:
Eachyearabout20pairsofcommoncarparespawnedandprogenytested.Thetestedpairs
mayincludethesethatexcelledinpreviousyears.Theymayalsoincludeagroupoffullsibsofa
superiorprogenyofthepreviousyearmassspawnedwithanunrelatedmale.Thefastergrowing
individualsofthebestprogeniesareselectedtoserveasparentsofcrossbreedfry.The
remainingprogeniesarecullod.Thesameprocessisrepeatedwithnewpairseachyear(Moav&
Wohlfarth,1967).
Asthenumberoftestedcombinationincreases,thepossibilityoffindinganew,bestcombination
becomessmallandfurtherimprovementrequiresselectionforcombiningabilitywithinthebest
parentalline.Thisrequirementismetintherecurrentselection(RS)andreciprocalrecurrent
selection(RRS).Thebasicfeatureoftheaboveselectionproceduresthatdifferentiatesthem
fromotherselectionprocedureisthatthepurebredparentsareselectedonthebasisofthe
performanceoftheircrossbredhalfsibsratherthanonthebasisoftheirownperformance(Fig.
6).
(b)Opengenepoolsystem:
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RRSmethodoranyotherbasedonfamilyselectionorprogenytestingisthatitincreases,
inbreedingintheparentlines.Toavoidthis,theuseofopengenepoolsystemmaybemade.This
systeminvolvesthemaintenanceofgeneticallymarkedreservegenepoolsforeachoneofthe
twoparentallines.Eachcycleofprogenytestingmayincludeasmallproportionoftheindividuals
fromthereservegenepools.
Iftheprogenytestingresultsshowthatoneormoreofthegenepoolimmigrantsproduced
superiorcrossbredprogeny,purebredprogenymaybeincorporatedintothepurebredparental
line.Selectionshouldaimatanincreaseincombiningcapacity(Moav&Wohlfarth,1967).This
methodhasalsobeenfollowedandextensivelyusedbySovietfishbreeders(Golovinskaya,
1971).
Hungary:
TheHungarianlandracesofcommoncarpwereusedforgeneticcrossing.Ineachcrossonly
onemaleandonefemalewereused.Thecarpspawnerswereidentifiedbyburnsorbrands,
boaringlineandindividualmarks.Theprogenypopulations,weredifferentiatedbycollective
marks,eachgroupreceivingabrandedstrip2cmlongondifferentpartsoftheirbody.Inplanning
crossingcombinationthefollowingaspectsweretakenintoconsideration(Fig.5).
i. Matingoutstandingfemalesandmalespossessingthesamevalueofstabilizedandfurther
improvementofadvantageouscharacters.
ii. Matingofanoutstandingmaleandfemalehavingtwodifferentcharactersforthe
combinationofadvantageousfeatures.
iii. Matingofonefemalewithoutstandingcharactersandmoremalesforthecontrolof
combiningabilityandfortheselectionofmostsuitablelineofmales.
iv. Matingofrelatedindividualswithinpurelinesfortheproductionofinbredlines.
Theproductivecapacityofcarphybridswasestablishedbytheevaluationofeggfertility,
percentageofsurvivalinthefirstandsecondyearsoflife,increaseinweightduringthesecond
year,foodconversionability,percentageofconsumablefleshandfatcontentofflesh(Bakos,
1979).

6CONCLUSIONS
i. Theefficiencyofmassselectioncanbeenhancedbyincreasingtothepossibleextent,the
valueofthreefactors:intensityofselection,variabilityandheritabilityofthetraittobe
selected.Ofthese,increaseinheritabilityisthemostimportant,whichcanbedoneby
meansofcrossing,creationofuniformconditionsforfishrearingandcarryingoutselection
attheagetobeimproved.
ii. Familyselectionandprogenytestingincarpbreedingprovetobemoreefficientthanmass
selectiononlywhentheheritabilityofthecharacter(h20.10.15).Themaindifficultyin
performingselectionforrelativesistherequirementoflargenumberofpondsforgrowinga
sufficientlylargenumberoffamiliesunderidenticalconditions.
iii. Muchbenefitmaybeexpectedfromcombinedselectioninthecourseofwhich,mass
selection,progenytestingoffemales,andfamilyselectionareperformedsuccessivelyin
onegeneration.
iv. Newtrendsinfishselectionsuchasuseofgeneticmarkers,artificialgynogenesisand
polyploidyhavebeendescribed.InIndianmajorcarps,viz.CatlacatlaandLabeorohita,
thegynogenesishasbeensuccessfullycarriedoutforthefirsttime.
v. Methodsofincreasingtheheterogenityoftheselectionmaterialandbreedingsystems,as
practisedinUSSR,IsraelandHungary,havebeendescribed.

7REFERENCES
Alikunhi,K.H.&H.Chaudhuri,(1959)Preliminaryobservationsonhybridizationofcommoncarp
(Cyprinuscarpio)withIndiancarps.Proc.IndianSci.Congr.,(3):46.
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LectureNotesonCompositeFishCultureanditsExtensioninIndia

Allen,S.K.andJ.G.Stanley,(1981)Polyploidyandgynogenesisinthecultureoffishand
shellfish.Int.Counc.Explot.Sea.F.,28:18pp.
Bakos,J.(1979)CrossbreedingHungarianracesofcommoncarptodevelopmoreproductive
hybrids.InAdvancesinAquaculture(T.V.R.Pillay&W.A.Dilleds.)FishingNewsBooks,
England:633635.
Barrackpore,(1980)(1979)CentralInlandFisheriesResearchInstitute,Rep.Cent.Inl.Fish.Res.
Inst.,Barrackporefortheyear1980and1979.
Brody,T.G.W.,Wohlfarth,G.HulataandR.Moav.(1981)Applicationofelectrophoreticgenetic
markerstofishbreeding.IV.Assessmentofbreedingvalueoffullsibfamilies.
Aquaculture,24:175186.
Bhowmick,R.M.,R.K.Jana,S.D.Gupta,andG.V.KowtalandM.Rout.(1981)Studiesonsome
aspectsofbiologyandmorphometryoftheintergenerichybrid,Catlacatla(Hamilton)x
Labeorohita(Hamilton)producedbyhypophysation.Aquaculture,23:367371.
Chaudhuri,H.(1959)ExperimentsonhybridizationofIndiancarpsProc.IndianSci.,Congr.46(4)
:2021.
Chaudhuri,H.(1971)FishhybridizationinAsiawithspecialreferencetoIndia.Rep.FAO/UNDP
(TA)(2926):5159.
Falconer,T.D.S.(1960)Introductiontoquantitativegenetics.Oliver&Boyd.Edinburgh:365pp.
Garvai,J.,S.Poters,A.Nagy,L.HorvathandV.Csanyi(1980)Inducedtriploidyincarp,
CyprinuscarpioD.J.Fish.Bio.,17:667671.
GeorgeJohn,P.V.G.K.ReddyandS.D.Gupta.(1984)ArtificialgynogenesisintheIndianmajor
carps,Labeorohita(Ham.)andCatlacatla(Ham.)Aquaculture.42:161168.
Gjedrem,T.(1983)Geneticvariationinquantitativetraitsandselectivebreedinginfishand
shellfish.Aquaculture,33:5172.
Golovinskaya,K.A.(1968)Geneticsandselectionoffishandartificialgynogenesisofthecarp
(Cyprinuscarpio).FAOFishRep.,(44)Vol.4:251252.
Golovinskaya.(1971)Breedinginfishculture.Rep.FAO/UNDP(TA)(2926):292300.
Ibrahim,K.H.,G.V.KowtalandS.D.Gupta.(1980)Embryonicandlarvaldevelopmentamong
Catlacatla(Ham.)xHypephthalmichthysmolitrix(Val.)Hybrid.J.Inland.Fish.Soc.India,
12(2):6973.
Kirpichnikov,V.S.Methodsofprogenytestingspawnersin(1966)carpfishfarms.Izv.GOS,
NauchnoIssled,Inst.Ozern.Rechn.Rybn.Khoz.,61.TranslatedfromtheRussianby
IsraelprogrammeofScientifictranslations,Jerusalem,IPSTCat.No.(5737):3655
(1970).
Kirpichnikov.(1967)Efficiencyofmassselectionandselectionforrelativesinfishculture.FAO
FishRep.,(44)Vol.4:179194.
Kirpichnikov.(1971)Methodsoffishselection.Aimsofselectionandmethodsofartificial
selection.Rep.FAO/UNDP(TA)(2926):217227.
Kirpichnikov.(1971)Methodsoffishselection2.Crossing,modernmethodsofselection,selection
techniques.Rep.FAO/UNDP(TA)(2926):217227.
Kirpichnikov.(1972)Theoryoffishselection.In:B.1Cherfas(Ed),Geneticselectionand
hybridizationoffish.AcademyofSciencesoftheUSSR.IsraelProgrammeforScientific
Translations,Jerusalem:269pp.
Kirpichnikov.(1981)Geneticalbasesoffishselection.Springer,VerlagBorlin:1140.
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Moav,R.&G.W.Wohlfarth.(1967)Geneticimprovementofyieldincarp.FAOFishRep.,(44).
Vol.4:227.
Moav,R.,G.W.WohlfarthandM.Lahman.(1960)Geneticimprovementofcarp2.Markingfish
bybranding.Bamidgeh,12(2):4953.
Nagy,A.V.Csanyi,J.BakosandL.Hervath.(1980)Developmentofashorttermlaboratory
systemfortheelevationofcarpgrowthinponds.Bamidgeh,32:615.
NaseemHamsa,K.(1971)PreliminaryaccountofanintergenericfertilehybridbetweenCirrhina
mrigalaandLabeorohita.Cent.Inst.Fish.Educ.Rec.Club.Scuvenir,1015.
Roddy,K.P.&T.J.Verghese.(1980)StudiesonthecomparativegrowthrateoftwoIndianmajor
carphybrids,Catlarohuandrohucatla.J.InlandFish.Soc.India12(2),1824.
Singh,S.B.&P.C.Chakraborti.(1970)Observationsonthehybridbetweensilvercarpandgrass
carpSABRAONewsletter2(2):103106.
Smisek,J.(1979)Considerationofbodyconfirmation,heritabilityandbiochemicalcharactersin
geneticstudiesofcarpinCzechoslovakia,Bull.VURHVednany,15(2):36Ref.Anim.
Breed.Abstr.48(5):302.
Smitherman,R.O.,R.A.DunhamandD.Tave(1983).Reviewofcatfishbreedingresearch1969
1981atAuburnUniversity,Aquaculture,33:197206.
Wohlfarth,G.W.Shootcarp.Bamidgeh,29(2):3556.(1977)
Wohlfarth,G.W.,M.LehmanandG.Hulata.(1980)ThestoryofDor70,aselectedstrainofthe
Israelicommoncarp,Bamidgeh,32(1):35.
Zonova,A.S.andV.S.Kirpichnikov.(1971)TheselectionofRopshacarp.Rep.FAO/UNDP(TA)
(2926):233247.
Table2:ShowingimportanthybridsofIndianmajorcarpsandChinesecarpsproducedin
India
Femaleparent
species

Maleparentspecies

1 Interspecifichybrids

Hybrid

Importantcharacteristics

Authority

Labeo
calbasu

Labeorohita

Calbasurohu Attainedfullmaturityin2
yearsF2generation
produced.Varying
charactersintermediate
betweencalbasuandrohu.

L.rohita

L.calbasu

rohucalbasu Inbothhybridsover94%
Do
fertilisationwasobtained,
theirgrowthratewasfar
superiortotheparentLabeo
calbasu.

2 Intergenerichybrids
Cirrhinus
mrigala

L.rohita

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Chaudhuri
(1959&
1971)

mrigalrohu

Relativelysmallhead,
Naseam
deeperbodyslendercaudal Hamza
peduncle.D2/1314P18 (1971)
V9A2/6C19,L.l.4647
Lengthofhead5.21of
caudal4.66heightofbody
4.57intotallength,
pharyngealteeth542.Full
maturityin2years.
Intermediatebody
characters.
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Labeorohita C.mrigala

rohumrigal

Intermediatebody
charactersascomparedto
thoseofparents.Attains
maturityin2years.

Chaudhuri
(1959&
1971)

Labeorohita C.mrigala

rohucatla

Bodycharacteristics
intermediatetoparent
species.Colourationlike
catla,smallhead,mouth
terminal,finraysresembling
thoseofrohu.Broadbodyof
catla,smallheadofrohu
andmoreflesh(54%)than
eitheroftheparents.viz.,
rohu(48%)&catla(45%).
Fastergrowththanrohu.
Fullmaturityattainedin3
years.Fecunditylessthan
bothparentalspecies.F2
generationproduced.
Slightlybettergrowthrate
thanthehybridcatlaxrohu.

Bhowmick
etal(1981)

Greaterbodygirthandfaster
growththanrohu,smaller
headthancatlaandmore
fleshthanboththeparents.
Primarilyplanktonfeederbut
acceptartificialfeed.
Growthslightlyfasterthan
rohubutslowerthancatla.
F1hybridsmaturedin2
years.F2generation
produced.Reddyand
Venghese(1980)reported
growthofhybridmuch
slowerthancatlaand
slightlylowerthanthatof
rohu.

Chaudhuri
(1971)

Chaudhuri
(1971)

Catlacatla

Labeorohita

Catlarohu

Reddyand
Verhese
(1980).

Reddy&
Verghese
(1980)

Catlacatla

L.calbasu

Catla
calbasu

Fastergrowthratethan
calbasu,smallerheadthan
catlaandbodygirthmore
thancalbasu.

L.calbasu

C.catla

Calbasu
catla

Fastergrowththancalbasu.

do

C.catla

L.fimbriatus

Catla
fimbriatus

Fastergrowththan
fimbriatusHeadsmaller
thancatlaandbodygirth
nearertocatla.

do

C.idella

Hypophthalmichthys grasscarp
molitrix
silvercarp

Resembledgrasscarp,
Barrackpore
irregularlargesizedscales (1980)
atsomeplacesalongthe
laterallineinsomecases
activelarvicidaland
weedicidaltendenciesinfry
stage.Malesmaturedin2
yearsandfemalesin3
years.

H.molitrix

C.idella

Roundbodyofgrasscarp
withsmallerscales.Mouth
withlowerlipprotrudingas
insilvercarp.Gillrakers
numerousbutnotfusedas
insilvercarp.Anaverage

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grasscarp
silvercarp

Singh&
Chakraborti
(1970)

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sizeof366mm/475ginone
year.Didnotmature.Non
acceptanceofweeds.
Labeorohita Cyprinuscarpio

rohu
common
carp

Elongateddorsalfinlike
Alikunhiand
parentspecies,butother
Chaudhuri
characteristicsintermediate (1959)
betweenparentspecies,
Sterile.Survivedforseveral
months.

C.mrigala

C.carpio

mrigal
common
carp

Survivalforseveralmonths

Catlacatla

H.molitrix

catlasilver
carp

Bodygirthcomparableto
Ibrahimetal
catla,sizeofheadand
(1980)
scalesnearertosilvercarp
andcolourationslategrey
ondorsalsideandsilveryon
toabdomen

H.molitrix

Catlacatla

silvercarp
catla

Asinglespecimen,survived Barrackpore
for21monthsandattained (1979)
497mm/1.45kg.Showed
roughnessofpectoralfin.

Cyprinus
carnio

L.rohita

common
carprohu

Bodycharacters
Khanetal
intermediatebetweenboth (1984)
parentspecies.Bodyprofile unpublished.
nearertorohu.

Chaudhuri
(1971)

Fig.1RelationbetweenselectionseverityandSelectiveintensity(,1967)NSizeofpopulation
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LectureNotesonCompositeFishCultureanditsExtensioninIndia

requiredforselectinganindividualVselectionseverity(%)iselectionintensity

Fig2Progenytestinginfishbreeding

Fig3TypesofCrossings

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Fig.4.SchemeofcreatingbreedingstockofRapshahybridcarp(kirpichnikov,1967)CC
culturedmirrorcarp,AWCWildCarp.HN+HKHybridsofNovogarad+KurskstrainsHB,
HC,HJhybridsofbackcross,CrissCross(withlinear)andintermediatestrains.Double
framsoutlinesheterosisCombinations?denotesuncontrolledcrossingduringthewar
194144.

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LectureNotesonCompositeFishCultureanditsExtensioninIndia

Fig5.SchemeofproductionofCommoncarphybridsshowingtheprocessofbreeding(Bakos
1979)

Fig.6.Schematicpresentationofacombinationofselection(offemalesoftheparentlineA)plus
moreselectionwithinthepureline(I1,andI2representtwotestedimmigrantsoftheopen
genepooloftheparentlineA)(Moav&Wholfarth,1967)

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