Professional Documents
Culture Documents
Producedby:FisheriesandAquaculture
Department
Title:Thedietofprawns...
Moredetails
THA:75:008/80/WP/12
by
MichaelB.New
SeniorFisheriesBiologist(Aquaculture)
ProgrammefortheExpansionofFreshwater
PrawnFarminginThailand
(FAO/UNDP/THA/75/008)
Bangpakong,Chacheongsao
Thailand
1980
HyperlinkstononFAOInternetsitesdonotimplyanyofficialendorsementoforresponsibilityforthe
opinions,ideas,dataorproductspresentedattheselocations,orguaranteethevalidityoftheinformation
provided.ThesolepurposeoflinkstononFAOsitesistoindicatefurtherinformationavailableonrelated
topics.
Thiselectronicdocumenthasbeenscannedusingopticalcharacterrecognition(OCR)software.FAO
declinesallresponsibilityforanydiscrepanciesthatmayexistbetweenthepresentdocumentandits
originalprintedversion.
THEDIETOFPRAWNSa
by
MichaelB.Newb
LECTUREI:WHATISKNOWNABOUTPRAWNNUTRITION?
1.INTRODUCTION
Theknowledgeofshrimpnutritionislessthanthataboutfishnutrition.Thisfirstpaperdealswiththeresult
ofcontrolledexperimentsthesecondisonamorepracticallevelanddealswithrealcommercial,rather
thanresearch,conditions.
Ifirstreviewedthistopicfouryearsago(New,1976).Inthesameyear,Biddlepresentedapaperonthe
sametopic,butrestrictedtothenutritionoffreshwaterprawns,atameetingintheUSA.Hispaperwas
largelybasedonminebutcontainedsomeadditionalpublicationandformedpartofabookonshrimp
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farming(Biddle,1976).Today'slecturehasbeenbasedonthesetworeviews,updatedbyanexamination
oftheliteraturepublishedsince1976.Abibliographyonthistopicisinpreparation(New,1980).
Thenotesthatfollowarearecordofmylecturenotesandthereforeonlysummarisewhatwasactually
said.
Briefcommentsweremadeontheconfusionintheterminology(i.e.,thewordshrimpversusprawn)the
differencesbetweenmarineandfreshwaterprawnsandbetweenpenaeidsandcarideansthemethodof
crustaceangrowth,etc.
Crustaceansabsorbnutrientsdirectlyfromthewateraswellasfromingestedmaterialthusphysiology
andnutritionislikelytobecomplex.Thislecturedealsonlywithnutrition(strictlydietaryexperiments),not
withphysiology.
Mostdietaryworkhasbeencarriedoutbyfeedstuffmanufacturerstheresultsarethereforeproprietary.
Theimportanceofdietinprawnculturemustbeborneinmindfoodwillbethesinglemostexpensive
runningcostinrearingprawnswhenthelabouroffeedingandofproducinglivefeedsistakeninto
account.Feedingprobablyrepresents5060%ofrunningcosts.
aThisworkingpaperisbasedontwolecturesgiventothe2ndInlandAquacultureCoursegivenattheNationalInlandFisheriesInstitute,
Bangkhen,Thailand(JuneAugust1979).
bSeniorFisheriesBiologist(Aquculture,(UNDP/FAO/THA/75/008).
SomeofthestatementswhichImakeaboutprawnnutritionmaybeapparentlyconflictingThisonly
demonstratesthepaucityofknowledgeonthesubjectandthefactthatcrustaceannutritionalresearchis
veryyoung.
2.POSTLARVALNUTRITION
a)PalatabilityandPhysicalDietaryStructure
Bothpalatabilityandstructureofthedietaffectingestion.
Themannerinwhichprawnseatinfluencestheneedfordietarywaterstability.
Wetdietsappeartogivesuperiorgrowthratesandsurvival.
Carideanslackthegastricmillingapparatuspresentintheanteriorchamberoftheproventriculusof
penaeids.Dryfeedsmaycongesttheproventriculusandhamperenzymaticmixingwithfoods.Penaeids
maytoleratedryfoodsbetter.Wetdietshoweverposemanufacturingandstorageproblems.
Theprocessingofdietaryingredientscandamagetheirnutritionalqualitythereissomeevidencethat
compoundingdoestoo.Caremustthereforebetakeniningredientselection.
Manybindersareusedinexperimentalrationsforpostlarvalprawnsthereisnoevidencethatthoseused
areinhibitorysomeareinfactnutritiousaswellasbeingbinders.Someexamplesofbinders:
Agar
Gelatin
Carboxymethylcellulose
Polyvinylalcohol
GuarGum
Sodiumalginate
Manucol(Glycolpolyesteralginate)
Collagen
LStarch
StickyRice
HighGlutenWheat
AmaizoStarch
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Dietswhicharenotwaterstablepolluteenvironmentmore.
Ingestionrateisgovernedbythefillingofthedigestiveglandthisisregulatedbyfoodsize.Thereforethe
animals'millingabilityisimportant.
Moreworkonfoodformandsizeisneededprawnswillacceptmostfoodspresentedthisdoesnotmean
thatthiscriterioncanbeusedtojudgetheefficacyofafeed.
Wetdietsarelesspracticalhowever,waterstabilitymaybelessimportantifthelabourtofeedseveral
timesadayorautomaticfeedersareavailable.
Foranydiet,thefinalarbiterofitscharacteristicswillbeitsefficiencvbythisIdonotmeanfood
conversionrateproteinconversionrate,etc.,Imeanthecostofthefoodneededtoproduceeach
kilogramofmarketableprawns.
Thechemosensorypropertiesofdietsarealsoimportant.Dietsshouldreleaseattractantsubstances.Live
feedsandfreshanimalproductsdothis.Prawnsareattractedeventootherprawnswhicharedying.This
phenomenonisquiteseparatetothetopicofpheromones.
Prawnsarestimulatedintofeedingactivitybysubstancessuchasbetaine(trimethylammoniumhydroxide),
morin(afragrantaromaticcompound),eggwhiteproteinsandindividualnonessentialaminoacidssuch
asglutamicacid,glycineandtaurine.OneAmericancompanyhasevenpatented(forwhatthat'sworth)
amethodofinducinghuntingandfeedingreactionsinprawnsbytheuseofamixtureofmonosodium
glutamatewithsodiumorpotassiumaspartate.
b)LipidsandSterols
Lipids
Withcarbohydrates,theyformsourcesofenergyiffeedenergyistoolowanimalswillutiliseother
nutrients,e.g.protein,tosatisfydemandforenergy,whichisverycostlyiffeedenergyistoohigh,reduced
foodintakewillresultandthereforenotenoughproteinwillbeingestedtogivegoodgrowth.
However,optimumcalorieproteinratiosforprawnsarenotyetunderstood(althoughonegroupof
scientistsclaim40%proteinand3.3Kcal/gisagoodcombinationforPenaeusmonodon).
Iamgoingtotalkaboutcarbohydrateslaterletusconsiderlipidsfirst.
Thereseemstobesomeevidencethatprawnscannottoleratehighlevelsofdietaryfat.Ifthisisso,thenit
isinmarkedcontrasttofish.Thefactthatlipaseactivityinprawnsislimitedhasbeendemonstrated
(palmiticacidisincorporatedintotissuethreetimesasrapidlyastripalmitin).
Oneworkershowedthat10%ormoreofabeeftallow,cornoilandmenhadenoilmixture
depressedgrowthandsurvival.Anotherfoundthattoincreasethedietarylevelofcodliveroilover5%
producednobenefitsthesameappliedtocornoil.Confusingly,myownpersonalexperienceisthat
Macrobrachiumcantoleratehighoildiets,atleastwhentheyoriginatefromacceleratedfreezedriedegg.I
usedadietcontaining20%oil.Generally,57%levelsaresuggestedcertainlynotmorethan10%.
Thetypeoflipidisalsoimportant.Tissuelipidsmimicdietarylipidsinquality,althoughw3fattyacidsare
retainedinthetissueandw6fattyacidsaremetabolisedforenergy.Thepredominantfattyacidsinprawns
arepalmiticacidandw3polyunsaturatedfattyacids.Oilswithhighw6saturatedfattyacidsandlowlevels
ofw3fattyacidsseeminhibitory.Fatswhicharesuitableinclude:
Linseedoil
Linolenicacid(1%)
Clamoil
Fishoils
Shrimpheadoil
Fatsthatgivepoorresultsifusedaloneare:
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Soybeanoil
Cornoil
Coconutoil
Shrimpfeddietswithshrimpheadoilalsohavemorecarotenoidpigmentanimportantconsumer
characteristic.Theuseofshrimpmealsasasourceofw3fattyacidsiscautionarybecauseofthehigh
variabilityofthesemeals.
Bothlinoleic(w6)andlinolenic(w3)acidsareessentialfattyacidsforprawnsbutthenutritivevalueof
linolenicishigher.
Sterols
Prawns,likeothercrustaceans,cannotconvert(synthesise)acetateintosterols.
Dietarylevelsof0.5%cholesterolseemadvantageous1.0%levelsarenobetter5%cholesterol
depressesgrowth.
0.5%ofstigmasterol,ergosterolorBsitosterolgivesimilarsurvivalto0.5%cholesterolbutpoorergrowth
rates.Shrimpcanconvertthesetocholesteroland,incontrasttothefat,absorbothersterolswell.
Sterolsareimportantbecausetheyare:
Elementsofcellularstructure
Precursorsofsteroidhormones,brainhormones,ecdysonesandvitaminD.
Shrimpcanabsorbdietaryecdysones.
Itisn'tlikelythattherewillbeadeficiencyofcholesterolinanymixeddiet,especiallyifitcontainsfishmeal
dietaryexcessmusthoweverbewatchedinformulation.
Thereiscommercialinterestinthepossibilityofusingdietarysterols,suchasecdysone,etc.,for
suppressinggonadaldevelopment,reducingagression,synchronisingmolting,increasinggrowthrate,etc.
SubstancessuchasmusselsorclamsarerichinsteroidsIdoubtifanycompoundeddiethasyet
producedsuchgoodresultsasmusselasafoodforprawns.
c)ProteinsandAminoAcids
Proteinisofdominantnutritionalimportanceandalsoisresponsibleforthegreatestcostcomponentinany
diet.Ithasthereforereceivedmoreattentionthananyothernutrienttodate.Inspiteofthis,noclearcut
guidestooptimumproteinlevelsinprawndietshaveyetemerged.
Crustaceanproteasesaremainlytrypticnopepsinlikeenzymesarepresenthowever,afullenzymatic
complementisunnecessaryifpredigestedfoodisavailable.Shrimpdigestdetritusandfaeces,for
example:85%ofthemicroorganismsinshrimpgutproducechitinase.Thesebugsmultiplywithinthegut
alsotoprovideanextrasourceofnutritionmostimportantlytheygivetheabilitytotheprawnstodigest
chitinaprotein/carbohydratecomplex.
Nutritionalexperimentsusingcaseinastheonlysourceofprotein,thoughusedinsomemineraland
vitaminassessments,donotsucceedwell.Dietsbasedonchemicallydefinedingredients,e.g.,containing
mixturesofpureaminoacidsorpeptides,donotproducegoodgrowth.Earlyattemptstousechemically
defineddietsforprawnnutritionalstudiesthereforequicklygavewaytodietaryexperimentsusingcomplex
andvariablefeedstuffingredients.Forthisreason,reportsofoptimumproteinlevelforprawndiets
reportedintheliteraturearedifficult,ifnotimpossible,tointerpretbecauseofthepossibleeffectof
concomitantchangesinotherdietarycomponents.
Thereisthusaverywiderangeofoptimumproteinlevelsclaimedbydifferentworkers.Dependingonthe
ingredientsused,optimumproteinlevelsbetween15%andmorethan60%havebeenrecorded.
Comparisonisalsodifficultbecauseexperimentationhasbeenwithdifferentspecies.
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Fromaneconomicpointofview,perhapsthemostencouragingresultwasthatofoneworker,working
withMacrobrachium,whoreportedthatbestresultswereobtainedwithasoytunashrimpmealdietwith
only15%protein.Thedietprovedbetterwitholderprawns(morethan120daysold)younganimals
appeartorequireproteinlevelsinexcessof35%,usingthisrangeofingredients.
Avastrangeofhighanimalproteiningredientshavebeenusedinprawnrationsvirtuallyallthe
conventionalfeedstuffingredientsplussquidmeal,prawnmeal,poultryfaeces(becarefuloftoxicity),
singlecellproteins,etc.Macrobrachium,whichseemsmorevegetableinclinedthanPenaeus,thriveswell
onrationscontainingcoconutmeal,soybeanmealandacaciameal.
Theconsensusofopinionseemstobethatproteinlevelsinthe2735%regionarethebest,but
commercialrationsforshrimpusedintheUSAhave2025%levels.
Ithasnotyetbeenshownexperimentallywhetherlipidsparesdietaryproteinfortissueproductionrather
thanenergyutilisation.Workwithisocaloricdiets(increasinglipidlevelswhilereducingproteintoproduce
dietswiththesamelevelofenergy)hasnotbeendoneyet.Optimumproteincalorieratiosareunknown.
Thequantitativeaminoacidrequirementsofprawnsarecomparativelyunknownyetthereforemuchof
theproteinfedinrationsissuretobeunbalancedinitsaminoacidcomposition.Thismaskstheresultsof
proteinexperimentsandisprobablythecauseofreportsthatextremelyhighlevelsarerequired.
Preliminaryresultsindicatedietaryrequirementsfor:
1.4%lysine
1.3%arginine
3.0%menthionine
0.6%tryptophane
Qualitatively,theaminoacidswhichareessentialinthedietofprawnsseemtobesimilartothoseforother
animalsalthoughtherearesomedisagreementonthis.Apparentnonessentialityofsomeaminoacidsis
suspectedtobecausedbytheabilityofthegutbacteriatosynthesisethem.Thefollowingaminoacids
havebeenshowntobeessentialforPalamonserratus,PenaeussetiferusandMacrobrachiumohione:
Arginine
Histidine
Isoleucine
Leucine
Lysine
Methionine
Phenylalanine
Threonine
Tryptophane
Tyrosine
Valine
Thevalueoffreeaminoacids(i.e.,notboundinprotein)asattractantsinprawndietshasbeenmentioned
before.Freeaminoacidsinthebodyofprawnsplayanessentialroleinosmoregulation.
Prawnmealhas,inmanyrations,beenexperimentallyshowntobeaparticularlyusefulhighprotein
ingredientwhich,uptonow,hasbeenunderutilisedinanimalfeedstuffs.Oneresearchworker,working
withP.indicus,anomniverousspecies,suggestedthata60:40ratiooffishmeal:prawnmealgavethe
bestresultsinhisseriesofexperiments.
d)Carbohydrates
ManycarbohydrasesexistincrustaceaincludingLandBamylase,maltase,sacdharase,chitinaseand
cellulase.
Carbohydrates,withlipids,formasourceofdietaryenergyandarealsoimportantinthestorageofenergy
throughglycogen,inchitinsynthesis,andinsteroidandfattyacidsynthesis.
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Wheatstarch,dextrin,andoysterglycogenareassimilatedbetterthanpotatostarch.Notallspecies
assimilatespecificcarbohydrateswiththesameefficiency.
Celluloseisalsopartlydigestedthusprawnscanutilisemarinealgae.
Crustaceansareabletoutilisecomplexpolysaccharides(starches)muchmoreefficientlythansomesimple
sugarssuchasglucose.Thisisinmarkedcontrasttofish.Itispostulatedthatglucoseisrapidlybut
inefficientlyutilisedwhilepolysaccharidesareabsorbedmoreslowlybutmoreefficiently.Somerecentwork
indicatesagainthatglycogenisagoodsourceofcarbohydrateforprawnssoalsoissucrose,asimple
sugarthisisaconfusingpicturetherefore.
Starchesarealsousedindietsasbinders,ashasbeenmentionedpreviously.
Extracellularchitinasesenablethedigestionofchitinfromdietarysourcesorfromcastexuviae.
Dietarycarbohydratehasasparingeffectonthecarbonchainsfromaminoacids,andthereforeondietary
protein,forchitinsynthesis.Chitinsynthesisisrequiredfortheexoskeletonandfortheperiotrophicfaecal
membraneinpenaeidprawns.
Someworkershaveconcludedthatglucosamineisnonessentialinprawndietsbutothershavefoundthat
replacementofglucosaminewithglucosecheckedgrowthrate.Glucosamineisanintermediarybetween
glucoseandchitin,whichisaproteincarbohydratecomplex.
Littleisknownaboutfibrerequirementsinprawndietsbutoneworkermanagedtorestoredecreasesin
growthratecausedbyexcessivelevelsofproteinbyincreasingdietaryfibre(vegetable)contentinthediet.
However,thegastrointestinaltractinprawnsisshort.Thereforethepassageoffoodisrapidandtimefor
digestionlimited.Thereisthepossibilitythatfibremayincreasemotilityandreducetheefficiencyofthe
diet.
Althoughoptimumproteincalorieratiosarenotyetquantifiedadequately,thereissomeevidencethat,ata
25%proteinlevel,1:1and2:1ratiosofcarbohydrate:fatresultinproteincatabolism,whereas3:1and4:1
ratiosreducethiseffectandseemsufficienttosatisfyenergyrequirements.
e)Minerals
Fewstudieshavereportedonthissubjectbutmanydietshaveusedmineralpremixesasakindof
insurancepolicy.
Resultsofearlyexperimentsinvolveddifferentmineralsupplementlevelsandplacedemphasisonthe
Ca:Pratiosinthepremixwithoutreferencetothecontentofthesetwomineralsinotherdietary
ingredients,especiallyinmaterialslikefishandprawnmeal.
Todate,workersreportthatdietaryCA:Pratiosofabove2.4:1seemtodepressgrowth.However,one
workerwithmarineshrimpclaimsthattheCatheprawnsneedcanbetotallyabsorbedfromthe(sea)
watertheyliveinandthatdietaryCaisnonessential.HealsosaysthatMgandFearenonessentialbut
2%P1%Kand0.2%ofatraceelementpremixproveduseful.ThelevelofPinseawaterislowanditis
thereforeclaimedtobeessentialinthedietbecauseprawnfleshishighinit.
f)Vitamins
Manydietsforprawnshavealsousedvitaminmixesdesignedforotheranimalsasaninsurancepolicybut,
inmanycases,enormouslyhighlevelsofvitamins,particularlyofVitaminD,cholineandVitaminChave
beengivenandthepotentialforacutehypervitaminosisorantagonisticreactionsmusthavebeenhigh.
Studyofthevitaminrequirementsofprawnshas,asyet,receivedlittleattention,asisthecasewith
mineralrequirements.
Qualitatively,mostoftheBgroupofvitaminsseemessentialinthedietofcrustacea,asarevitaminsCand
E.VitaminDmaycomepartlyfromdietarysourcesbutitcanbesynthesisedfromergosterol.Ithasbeen
showntobenonessentialininsectdiets.VitaminKmayactuallybeantagonistictosomespeciesof
crustacea.
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VitaminAisprobablyinessentialbutitsprecursorsmaybeessential.OftentheoriginisBCarotene,which
isconvertedintoastaxanthinaswellandisindirectlyresponsibleforcrustaceanpigmentation.Prawnmeal
canbeusedasasourceofcarotenoidsbutthesecanbedestroyedbyexcessiveheatduringthe
productionofprawnmealandalsobyexposuretolightandatmosphericoxygen.Thequalityofprawn
mealutilisedisthereforeimportant.Spirulina,abluegreenalga,isabetterprecursorofastaxanthinthan
alfalfaorcorngluten.Artemiais,ofcourse,agoodsourceofcanthaxanthin.
VitaminCseemstobeessentialinprawndietsandappearstoaccelerategrowthatlowlevelsbutdepress
growthathighlevels.Stress(throughhighdensitycultureofdiseaseorbadwaterquality)cancause
vitaminCdepletionintissuestissuelevelscanbemaintainedbydietaryinclusionofVitaminC.One
workerhasclaimedthat0.51.0%dietaryvitaminCisthequantitativerequirementbutanother,testinga
rangeoflevelsbetween0%and1%suggestedthat0.3%wasoptimalandthathigherlevelsdepressed
growth.
Theonlyothervitaminwhichseemstohavebeenexaminedinprawndietsisinositolanda0.4%levelhas
beenpostulatedasoptimal.
g)OtherSubstances
Sofar,Ihaveonlyseenonereportofanantibioticusedinprawndiets.Thiswasinanexperiment
designedaspartofadiseaseinvestigationbutthereweresomeinterestingsideeffects.Inashortterm
experiment,usingdietarylevelsof1001000mg/kgofdryfoodofoxytetracycline,feedintakedroppedto
betweenandofthatofthecontrols.Foodconversionefficiencyhoweverincreasedand,whilelarger
shrimp(av0.46g)grewlesswell,smallshrimp(av.0.14g)actuallygrewfasterthanthecontrol.
h)FeedingRateandFrequency
Finally,inthissectiononthenutritionofpostlarvalprawns,afewwordsaboutfeedingmethods.
Dietaryexperimentshavebeencarriedoutmostlybyfeedingtodemandbutfeedingbasedonestimates
ofbiomasswouldbemoreefficient.
Ingestionratesareinverselyrelatedtoanimalsize.Feedingratesof10%ofbiomass(sometimesashigh
as100%)areemployedinthefirsttwopostlarvalmonthsbutthesedeclineto35%.
Littleisknownabouttheeffectsofvaryingenvironmentalconditionsonfoodconsumptionandtheefficiency
offoodutilisation.
Innature,prawnseatfrequentlyandrapidly.Theyalsohaveshortdigestivetractsandlittletimefornutrient
absorption.Feedingthreeorfourtimesperday,ratherthanonce,mayprovebetterandalsosolvesomeof
thedietarystabilityproblems.Decreasedingestionrateshavebeenobservedaftermorethan6hours
exposurebyprawnstothesamefood.Anotherimportantobservationisthat,eveninanapparentlywell
bounddiet,morethan90%ofitsthiaminecontentleachesfromthedietwithin18hours.Thissortofeffect
isprobablyexperiencedbyotherwatersolublenutrients.
3.LARVALNUTRITION
Althoughpenaeidandcarideanlarvaehavedifferentfoodrequirements(thistopicisdealtwithmorefullyin
mysecondtalk),theemphasisinlargescalelarvalproductionhasinbothcasesbeenontheuseof
mixturesoflivefeedswithothernaturalmaterials.Iamnotgoingtosayverymuchaboutthetopicof
experimentalartificialdietsforlarvae,becausenotmuchisknownyet.Iwillbetalkingmoreabout
practicaldietsinmysecondtalk.
Therehasbeensomepreliminaryworkonthedevelopmentofcompoundeddietsforprawnlarvaeanditis
tobeexpectedthatthistypeofresearchwillgrow,withthehighcostoflivefoods(thecostoflivefood
production,aswellasthecostofArtemiacysts,forexample,mustbeborneinmind)actingasan
importantstimulanttosuccess.
Thephysicalproblemsinmakingartificallarvalfoodinclude:
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1.Nutrientsmustnotleachout
2.Thefoodmustnotpollutetherearingwater
3.Thefoodmustbeavailablephysicallyclosetothelarvae
4.Itmustbeofasizewhichlarvaecantacklethezoeastagesofmarineprawnlarvaecanonlyeat
algaeatapproximately310ubutthemysisstagescaneatArtemianauplii,attentimesthesize
MacrobrachiumlarvaeontheotherhandcantackleArtemianaupaliialmostimmediatelyafter
hatching:theycanalsouselargerparticlesoffoodbutiftheparticlesaretoolarge,theweightof
severallarvaedragslarvaeandfoodtothebottomofthetank.
5.Itmustattractthelarvae
6.Itmustbedigestible
IfalarvaldietforMacrobrachiumistoconsistofparticlesofasizerangebetween250uand1500ufor
exampleandifeachparticleistorepresentabalancedmixtureofallthedietaryingredients,thelatterhave
tobegroundextremelyfineindeed.
Neutraldietarybuoyancyinwaterisdifficulttoachievebindingthedietwithoutmakingitinedibleisalso
difficult.
Somelarvaldietshavehoweverbeenformulated:
1.Microcapsuleswithcrosslinkednylonproteinwalls:thesecanbeproducedinarangeofsizesfrom
5uto250uwiththepropertyofneutralbuoyancy.Macrobrachiumlarvae,Artemiaandcrabzoea
havebeenshowntoingestthem.
2.Flakedandgel(alginate)andfreezedriedfeedshavebeentried.Gelandfreezedrieddietswere
neutrallybuoyant.
Sofar,resultsoflarvalproductionexperimentsusingsuchdietshavenotbeengoodunlessliveArtemia
arealsofedoranextractofArtemiaisincludedinthediet.
LECTUREII:WHATFEEDSAREUSEDINCOMMERCIALPRAWN
PRODUCTION?
1.INTRODUCTION
InmyfirsttalkIspokeaboutresearchworkdesignedtoincreaseourbasicknowledgeofthenutritional
requirementsofprawns.Inthecaseofpostlarvaldiets,suchknowledgeisfundamentallyimportantto
thosewhowishtocultureprawnsinenvironmentallycontrolledsystemswherenaturalfeedsmaybe
unavailable.Inthecaseoflarvae,artificialfeedsarecriticallyneededtoremoveorlessendependenceon
scarceandcostlyresourcesofsuitablelivefeed.InthissecondtalkIamgoingtospeakaboutthosefeeds
whichareusedinhatcheriesandpondswhereprawnsarereared.
Intheprevioustalk,Ispokefirstaboutpostlarvaldietsbecausemoreprogresshasbeenmadewiththem
thanwiththedevelopmentofartificiallarvaldiets.InthistalkIamgoingtoreversetheorder,sinceprimary
attentionhasbeenpaidtohatcherydietsratherthantodietsforuseinpondsforpostlarvalrearing,where
naturalfoodsarealsoavailable.
2.FEEDSFORLARVALPRAWNS
Intheirnaturalenvironmentitislikelythatprawnlarvaeconsumeavastrangeofmaterialswhichare
presentinthewatertablewiththemanythingprobablywhichpresentsitselftothemandisoftheright
particlesize.Thisincludeslivephytoplanktonandzooplanktonofmanytypesandnonliveparticles.
InmyfirsttalkIgeneralisedbetweencarideanandpenaeiddietaryrequirementsasmuchasIcould.Inthis
sectionofthistalkImustseparatethetwogroupssincetheyhaveratherdifferentrequirementsintermsof
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bothfoodandfeedingtechniques.Thereasonforthesedifferencescanbeunderstoodthroughan
examinationthedifferenceinthelifehistoryofPenaeusspp.andMacrobrachiumspp.Inmakingthese
comparisonsImustmakeitclearthattherearedifferencesbetweendifferentspeciesineachgroupaswell
so,whenItalkaboutthelengthofeachlarvalstage,forexample,Iamnottalkingaboutanyparticular
speciesbuttryingtopointoutmajordifferencesbetweenthetwogroupsofprawns.Basicdifferencesare
summarisedinthefollowingtable:
PENAEUSSPP. MACROBRACHIUMSPP.
0.25mm
0.60.7mm
Notcarriedbyfemale
EGG Carriedbyfemaleabout20days
Highfecundity:100,000
Lowerfecundity:10100,000
1,000,000
0.250.5mm
NAUPLIUS Doesnoteatlivesoffyolk None
Periodlasts23days
0.52.2mm
ZOEA Eatsalgae
Periodlasts5days
(Larvae=Zoea)
2.25mm
26mm
EatsArtemiaandother
EatsArtemiaandotherparticulatefoodfromfirst
MYSIS particulatefoods
orsecondday.Algaearenonessential
Periodlasts45days
Totallarvallife1825days
Totallarvallife1113days
METAMORPHOSIS
POSTLARVAE 56.7mm:Pelagic 6mm:Benthic
7.5:Benthic
Hastobeweanedslowlyoff Canbefedbenthicorartificialdietsimmediately
livefeedsontononlivediets aftermetamorphosis.
Thusitisnecessarytoculturealgaeinmarineshrimphatcheriesbutnotinfreshwaterprawnhatcheries
(althoughthiscommentisqualifiedlater).ThelarvallifeofpenaeidsisshorterandArtemiaareonly
requiredforabout5daysMacrobrachiumrequireArtemiaforthewholeofthelarvallifeof18days+.
Conversely,penaeidshavearequirementforlivefoodsintheirearlypostlarvallife,whereas
Macrobrachiumdoesnot.
Thushatcheriesdesignedformarineprawnproductionusequitedifferenttechniquestothosein
freshwaterprawnhatcheriesthisdoesnotonlyapplytofeedingregime.Perhapsthesedifferencesareso
greatastomakehatcheriesattemptingtorearbothtypesofprawninadvisable.Iwillthereforedealwith
thefeedingofmarineandfreshwaterprawnsseparately.Itmustbestressedthateverydifferenthatchery
usesadifferentfeedingregimeandfrequentchangesaremade,basedonlocalexperienceandonfood
availability.IcannothopetocoverallthesevariationsandthereforeIamjusttalkingaboutexamplesfor
illustrativepurposes.
a)PenaeidLarvae
Newlyhatchedmarineprawnlarvaeliveofftheireggyolkandthenaupliidonotfeed.However,chemical
fertilisationoftheirrearingtanksispractisedfromthefirstdaysothatphytoplanktonareavailablefor
consumptionassoonastheanimalsbecomezoea(insomehatcheries,phytopplanktonareculturedin
separatetanks).
After23daysthenaupliibecomezoeaandbegintofeedonphytoplankton.Diatomsarebest.Species
suchasSkeletonema,Thalassiosira,MelosiraandNitzschiahavebeenused,aswellasTetraselmis.
Whengrownwiththelarvae,phytoplanktonconcentrationsof5,00020,000cells/mlareused.23g/m3of
KNO3and0.20.3g/m3ofNaH2PO4willachievethislevel.Ifdiatomsfail,assometimesunpredictably
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happensinphytoplanktonculture,formulafeed,yeastandsoybeancake,crumbledtolessthan100u,has
beensuccessfullysubstituted.
Youcanalterthepredominantspeciesofphytoplanktonpresentbutthisisbestachievedifthealgaeare
grownseparatelyfromthelarvae.Thevariablequalityofseawateroftencausesproblemsinphytoplankton
cultureandartificialseawaterissometimesusedinhatcheries.Inmarineshrimplarvaltanks,brownwater
istakenasanindicationthatpredominantlydiatomsarepresent,whichisgoodgreenwaterheraldsa
failure.Theconverseisthecasewithfreshwaterprawnlarvaltanks.
Afteranother5days,thezoeachangeintothemysisstageandareabletotakeArtemia.Theyarenow
aboutthesamesizeasMacrobrachiumlarvaearewhentheyarefirsthatched.Artemiaareusually
suppliedtothemysislarvaeatdensitiesvaryingfrom0.33nauplii/ml.
Afterafurther45daysorso,thelarvaemetamorphoseintopostlarvaebutcontinuetobepresentedwith
Artemiaoverthefirst45days,whiletheyarechangingfromanomnivoroustoamainlycarnivorousdiet.
Atthisstagetheyareconsuming5090Artemianaupliieachperday.Afterthattheyarefedanaturalfood
suchasprocessedmolluscanfleshoraformulafeedaswellasArtemia,whichisdiscontinuedaftertwo
moredays.
Itisimportanttorealise,asisthecasewiththefeedingofMacrobrachiumlarvae,thatitisthedensityof
foodparticlesperunitofwatervolumethatismostimportant,notthenumberofparticlesperlarvapresent
inthetank.Thisiswhy,forefficientutilisationoffoodavailable,optimisationoflarvalrearingdensityis
sought.
Theabovedescribesatypicallarvalfeedingsystemforpenaeidsoneworkerhashoweverclaimed
successfulrearingoftwospeciesofPenaeusfedentirelyonamonocultureofthediatomChaetoceros
gracilisatconcentrationsof30,000100,000cellsperml.
b)Caridean(MacrobrachiumRosenbergii)Larvae
Unlikepenaeids,theyhavenoessentialrequirementforalgae.
TheywilleatlivinganimalfoodslikeArtemia,therotiferBrachionusandpreparedfoodsalmost
immediatelyafterhatching.
BythetimeMacrobrachiumlarvaehavereachedtheninthstagetheyare,likepostlarvalpenaeidprawns,
consumingabout80Artemianaupaliiperday.
Onceagain,thedensityoffoodisimportantand,whereArtemianaupliiarethesolesourceoffood,a
densityofnaupliiofbetween5and10/mlisessential,whateverthedensityoflarvaeinthetank.
LarvaewillgrowandsurvivewellandmetamorphoseonadietofArtemianaupliialone,butthereis
evidencethatthepresenceofotherfoodsafterthefirstfewdaysresultsinbetterproduction.Forthis
reasonandtoeconomiseintheuseofArtemia,otherfoodsareusedtoreplacepartoftheArtemia,such
assievedfishflesh,cookedeggcustard,frozenadultArtemia,etc.Althoughthereisevidencethatthe
nutritionalvalueofArtemianaupliicanbeimprovedbyallowingthemtograzeonphytoplanktonbefore
feeding,adultArtemiaareunsuitableasafoodasmanyevadethelarvae.
Otherlivefoodshavebeenused,suchasBrachionus,fisheggs,Daphnia,oysterlarvae,etc.,butnoneof
themandnoartificialfoodhasyetbeenfoundwhichcansuccessfullytotallyreplaceArtemiaasalarval
foodforprawnlarvae.ThecriticalrequirementoftheseandsomefishlarvaeforArtemiamakesthesupply,
demandandcostofbrineshrimpanoverwhelminglyimportanttopic.SuccessfulinsulationofArtemia,
leadingtotheproductionofThaigrowncystsinsaltponds,hasbeenreported(Tunsutepanich,1979).
Thisworkgiventhepossibilityofcheaper,localsupplyofArtemiawhichis,afterall,anidealprepackaged
larvalfood.
AlthoughMacrobrachiumlarvaedonotrequirephytoplankton(althoughtheywillingestalgae,evidence
showsthattheydonotderiveanynutrientsfromthephytoplanktontested),thereisevidencethatthe
presenceofphytoplanktoninlarvalrearingwater(greenwater),contrarytopenaeidlarvalculture,is
beneficial.Greenwaterisclaimedtoimprovewaterqualityandtoreducediseaserisks,butthereal
reasonsforitssuccessareunknown.Phytoplanktonarecapableofconvertingtheexcretaryproductstoxic
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tolarvae(unionisedNH3),whichareproducedbythelarvaethemselvesandalsobytheArtemianauplii
presentandbythedegradationofotherdecomposingfeeds,tolessharmfulnitrates.Manyhatcheries
however,includingthegovernmentoneclosetoushere,arecurrentlysuccessfullyusingaclearwater
systemwhichdoesnotinvolvegreenwater,usingonlyArtemianaupliiandfishfleshormussel/eggcustard
asfeeds.Oneofthebigproblemswiththeuseofagreenwater,orpolycultureofphytoplanktonspeciesis
thatitsproductionisunpredictable.
Asmentionedabove,theuseofotherfeedsenablesthequantityofArtemianeededtobereduced.For
example,inourhatcheryinEngland,whichbynecessitywasenvironmentallycontrolled,weoriginallyused
thenaupliifrom50gofArtemiacystsperdayper600litretank.Thiswasequivalenttoapproximately8
nauplii/ml/dayandanavailability(notaconsumption)of200nauplii/larvae/day.Byusingsievedmackerel
flesh(afishsimilartoskipjacktunainitscharacteristicsasafood),gradedtolessthan250uandfedata
rateof50gper600litresperday,wewereabletoreducethequantityofArtemiacystsusedto5gpertank
perdaytheequivalentofnearly1naupliipermlperdayortheavailabilityof20naupliiperlarvaeper
day.Ineachcasewewereraisingbatchesof20,00030,000larvaepertank(equivalentto3350larvae
stockedperml)andwewereusingsome5litresofphytoplankton(Chlamydomonas)atanoriginaldensity
of1,500,000cellsperml.Thusthedensityofphytoplanktonintherearingtankswasapproximately12,500
cells/mlonly:thesealgaewereaddedtoimprovethequalityofArtemia,notasafoodforprawnlarvae.
Ingeneral,inmixedfoodregimes,otherfoodsarepresentedduringthedaytime,whenutilisationcanbe
judgedbyeye,atseveraldifferentfeedingtimes.TheArtemianaupliiportionofthedietisnormally
presentedinthelateeveningasthelastfeedasameansofensuringthatfoodsuppliesdonotrunout
duringthenight.Itisessentialthatlarvaeareabletofeedcontinuously.
3.DIETSFORPOSTLARVAE
Undernaturalconditions,postlarvalprawnsconsumeaverywiderangeoffoods.
Penaeidsareconsideredtobeeitheromnivorousscavengersordetritusfeeders.Thenaturaldietofsuch
speciesasP.mondon,P.indicusandP.merguiensisconsistsofmolluscs,smallcrustaceans,polychaetes,
algaeanddetriuts.P.indicuscaneatlargercrutaceans.Allcanbecannibalistic.Metapenaeusspp.seem
tobemorevegetariananddetritusinclined.Thus,polycultureofpenaeids,ratherthanmonoculture,may
beamoreefficientwaytoutiliseapond.
Macrobrachiumpostlarvaeseemtofeedmostactivelyatnightandareomnivorous,eatingaquaticinsects,
fish,molluscs,othercrustaceans,algae,leavesandstemsofaquaticplants,plantseeds,detritus,etc.
ThereisabeliefthatMacrobrachiumspp.aremorevegetarianintheirdietthanpenaeids,buttheywillnot
thriveonadietofalgaealone.
Bothmarineandfreshwaterprawnshaveaccesstochitinfrominsects,crustaceaandbacteria.Thus
prawnmealmaynotbesoessentialindietsdesignedtobefedinpondsasitseemstobeincaseswhere
thecompoundeddietisthesolefeed.Obviously,inthepondcultureofprawns,dietcanbeveryvariedas
thereisagreatdealofnaturalfoodavailable.Thecompositionofsupplementarydietsmaynothavetobe
ascarefullyconsideredasmightatfirstbeexpected.However,inhighlyintensivesystems,wherenatural
foodwillbeinsufficient,thecompositionofsupplementaryfeedsmaybecomecritical.
Iamfirstgoingtodealbrieflywiththestimulationofnaturalpondfoodbeforegoingontotalkabout
supplementaryfeeds.
Inthecaseofmarineprawnculture,naturalfoodproductionisnormallyencouragedbyfertilisation.
Generallyfertilisationand/orfeedingisnecessaryifproductionlevelsaretoberaisedabove400
500kg/ha/yr.
Thepracticeoffertilisationbringswithitsomeproblemssuchasdensephytoplanktonbloomswhichmay
cause(inadditiontogrossturbidity)supersaturationofDO2bydayandDO2depletionatnight.The
subjectofpondfertilisationisawholeseparatetopiconitsownandIsuspectthatyouwillbereceiving
otherlecturesaboutthisinrelationtofishculturegenerally.
Whentalkingaboutfertilisationregimes,therecanbenotruegeneralisationsincethesystemoffertilisation
suitedbesttoeachindividualponddependsonthetypeoffertiliserlocallyavailable,onwaterquality,on
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soilconditions,onthefeedingregimetobeemployed,etc.Thefollowingaregivenasexamplesofsome
fertilisationprogrammeswhichhavebeenrecommendedformarineshrimpculture:
1.Sundrythepondsapply170kg/haofcottonseedmealaddafurther56kgtothewatereveryweek.
2.Fornurseryponds(thefirsttwomonths),encouragethegrowthoflablab,whichisabasisofblue
greenalgaeinwhichdiatomsandotherplantsandanimalsgrow,byapplying350kg/haofchicken
manuretothedriedpondalternatively,50100kg/haof18:46:0(N:P:K)or100150kg/haof16:20:0
(N:P:K)canbeusedtheseamountsarethenaddedtothewateragaineverytwoweeks.
3.Inproductionponds,toencouragethegrowthoflumut,basicallyafilamentousgreenalgae,addan
initial180200kg/haof16:20:0(N:P:K)fertilisertothewater,followedby90100kg/haofthesame
fertilisereveryotherweek.
4.Forphytoplanktongrowth,diatomsbeingpreferableforpenaeids,useof20:1or30:1ratiosofN:P
aresuggested.Ratiosof1:1arebestforflagellateproduction.
Note:Ammoniafertilisersaremostlyabsorbedbythesoil.Theyarethereforebetterforbluegreenalgae
production.Nitratefertiliserstendtoremaindissolvedinthewaterandarethereforebetterfor
phytoplanktonproduction.
Theamountoffertilisertouseforphytoplanktonproductionisbestassessedonatrialanderror
system.Tostartwith,1ppmofNand0.1ppmofPissuggestedfurtherapplicationsshouldbebased
onresults.
PondsforcultivationofMacrobrachiumareoftennotintentionallyfertilisedbutsomefarmersdosomainly
toencourageaphytoplanktonbloomtodiscouragefilamentousalgaeandprovidecover.Waterdepthand
flushingratesarealsomanipulatedtoencouragephytoplanktongrowthratherthanfilamentousalgaean
interestingcontrasttotheuseoflumutinmarineshrimpcultureharvestingtechniquesalso,ofcourse
differ.
NowwecometothepointwhereIshouldsaysomethingaboutsupplementalfoodforpostlarvalprawns
andIcouldeasilyendmylecturesimplybysayingyounameit,theyfeedit.Probablymuchofthe
supplementalfoodaddedactsasafertiliserandincreasesthebiologicalproductivityofthepondasa
wholeratherthanactingasatrueprawnfeed.Certainlyitseemsthat,particularlyinMacrobrachium
culture,muchofthepelletedfoodaddediseatenbysmallfish,whichthemselvesformasourceoffoodfor
theprawns.Largerfish,ofcourse,notonlybecomecompetitorsfortotalpondnutrientsbutmaybe
predatory.
Supplementalfoodsfallmainlyintothreecategories:
1.Rawmaterialsandwasteproducts
2.Compoundfeedstuffsdesignedforotheranimalsbutusedforprawns
3.Compoundedprawnfeeds
Iproposeonlytogivesomeexamplesofeach,withoutmakingjudgementsaboutwhichisbest.This
judgementmustdependentirelyontheinterrelatingfactorsoffoodavailability,foodcostandproductions
ratesofmarketableprawnsachievedineachspecificlocationgeneralisationisthereforeunwise.
a)Examplesofrawmaterialsandwasteproducts:
Ricebran
Brokenrice
Trashfish
Fishoffal
Deadpoultry
Slaughterhousewaste(poultry,beef,hog,etc.)
Fishprocessingwaste
Prawnprocessingwastes
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Snails
Mussels
Clam
Squid
Byproductsofvegetableoilmanufacture
Alltypesofplantwaste
b)Examplesofcompoundeddietsforotheranimals:
Chickenfeed(1826%protein)
Hogfeed(probably1420%protein)
c)Examplesofcompoundedprawndiets:
Afewdietsareonthemarketbuttheirformulaearekeptsecret.Althoughmajoringredientsmaybelisted,
thequantitiesofeacharenotgiven.RalstonPurinasell20%,25%and,Ibelieve,30%dietsforshrimp
(marine),whichhavealsobeenusedforfreshwaterprawns.
IbelievesomeproprietaryprawncompoundsareonsaleinJapan,thePhilippinesandTaiwan.Iam
hopingthat,inourdiscussiontimeinafewminutes,youwillbeabletotellmewhatfoodsareusedin
prawnproductioninyourowncountries.
ThefollowingisalistoftheingredientsofatypicalformulaforthePurina25%marineshrimpration.In
examiningitonemustremembernotonlythatthequantitiesarenotstatedbutthatitisnormalpolicyfor
feedstuffmanufacturerstoaltertheingredientcompositionofanyoftheircompounds,withoutnotice,
accordingtocurrentavailabilityandingredientcost.
Thelistisasfollows:
Fishmeal Proximateanalysis
Soybeanmeal
Groundwheat
Brewersdriedyeast H2 11%
Driedwhey
SoybeanOil Oil 12.5%
Decalciumphospate Protein 25.5%
Iodisedsalt
Fibre 4.1%
VitaminASupplement
Dactivatedanimalsterol Ash 6.7%
N.F.E. 40.2%
Menadionedimethylpyrimidinol
100.0%
bisulphite
VitaminEsupplement
VitaminB12supplement
VitaminC
Biotin
CholineChloride
Folicacid
PyrodoxineHCL Comments
Thiamin
Niacin Oilover10%
Contains
Calciumpantothenate
VitA
Riboflavinsupplement ContainsVitK
Copperoxide
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Manganousoxide
IronOxide Noprawnmeal
Zincoxide
Containssoybean
Calciumcarbonate
oil
Cobaltcarbonate
ThefollowingmixtureisusedbysomeHawaiianfreshwaterprawnfarmers:
90%BroilerStarterPellets
10%Prawnmeal
ThefollowingisashrimprationusedforpondcultureexperimentsforPenaeusmonodoninTahiti:
Shrimpmeal 8%
Bloodmeal 11%
Meatmeal 21.5%
Wheatgluten 10%
Rice 6%
Groundnutoilcake 17%
Solublefishproteinconcentrate(80%protein) 6%
Codliveroil 4%
Mineralmix 3%
Vitaminmix 5%
Methionine 0.5%
Thoseofyouwhoaregoodatmentalarithmeticandarestillawakeatthislatestageintheafternoonwill
noticethatthislistofingredientsonlyaddsupto92%.Anotherdietquotedonthesamepageasthisone
wasrecordedaddedupto101%abeautifulexampleoftheproblemsinreviewingandtryingtointerpret
theliteratureonprawnnutrition.Icouldgivemanyothers.
Icouldprovideotherformulaeforprawndietstherearemanyintheliteraturebutmosthavebeenusedfor
tankorlaboratoryexperimentsratherthanforcommercialprawnproduction.Inmostprawnfarmsfeeding
isanart,notascience,andthosematerialswhichareclosesttohandandcheapestareutilised.Whilethis
mayresultinprofitnow,intensiveculturewillrequireamuchfullerunderstandingofthedietary
requirementsofprawnsinpondculturethanwehaveavailablenow.
Itisn'treallypossibletosaymuchaboutfeedingratesforsupplementaryprawnrationsbecausethese
alwaysseemtobegovernedbyeyeballjudgementsonhowmuchfoodisuneatenfromyesterdays
feeding?.Thistechniqueislikelytoleadtoalternateperiodsofnearstarvationandsatiation,thelatter
combinedwithdubiouswaterquality.Tablesoffeedingrates,basedonbiomassweightsandprawnsizes,
suchasareusedinfishfeeding,arerequiredforprawns.Oncecompoundedprawndietsbecomereadily
andcommerciallyavailableitistobeexpectedthattheirmanufacturerswillresearchthissubject.One
feedingrateinpondswhichwasrecordedintheliteraturewasforMacrobrachiumfarminginHawaii,
whereaproductionof3000kg/hawasachievedthroughregularcullharvestingtechniques.Here
supplementalfeedwaspresentedatratesbetween28and45kg/ha/day.Onemoreexampleofthe
problemsofstudyingthistopic:thewriterofthereportgivingthisinformationwentontosaythatgallon
ofchow/acre/dayisagoodrateforanewlystockedpond.Suchastatementisalmostincomprehensible!
Howmuchdoesagallonofchowweigh?
4.REFERENCES
Biddle,G.N.,1976.Thenutritionoffreshwaterprawns.In:J.A.HansonandH.L.Goodwin(Editors),
shrimpandPrawnFarmingintheWesternHemisphere,Dowden,HutchinsonandRoss,Inc.,
Pennsylvania,USA.,pp.272291
New,M.B.,1976.Areviewofdietarystudieswithandprawns.Aquaculture,9:101144.
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New,M.B.,1980.AbibliographyofshrimpandprawnnutritionAquaculture,(inpress).
Tursutapanich,A.,1979.CystproductionofArtemiasalinainsaltpondsinThailand.FAOWorkingPaper
THA:75:008/WP/79/9.
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