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2.ESSENTIALNUTRIENTSPROTEINSANDAMINOACIDS
2.1Proteins
Theproteinsareamongthemostimportantconstituentsofalllivingcellsandrepresentthelargestchemicalgroupinthe
animalbody,withtheexceptionofwaterthewholefishcarcasscontainsonaverage75%water,16%protein,6%lipid,
and3%ash.Proteinsareessentialcomponentsofboththecellnucleusandcellprotoplasm,andaccordinglyaccount
forthebulkofthemuscletissues,internalorgans,brain,nervesandskin.

2.1.1Composition
Proteinsareverycomplexorganiccompoundsofhighmolecularweight.Incommonwithcarbohydratesandlipids,they
containcarbon(C),hydrogen(H),andoxygen(O),butinadditionalsocontainabout16%nitrogen(N:range1219%),
andsometimesphosphorus(P)andsulphur(S).

2.1.2Structure
Proteinsdifferfromotherbiologicallyimportantmacromoleculessuchascarbohydratesandlipidsintheirbasic
structure.Forexample,incontrasttothebasicstructureofcarbohydratesandlipids,whichisoftencomposedof
identicalorverysimilarrepeatingunits(ie.theglucoserepeatingunitwithinstarch,glycogenandcellulose),proteins
mayhaveupto100differentbasicunits(aminoacids).Itfollowsthereforethatgreatercompoundvariabilitiesand
rangesarepossible,notonlytocomposition,butalsotoproteinshape.

2.1.3Chemicalproperties
Colloidalinnature,proteinsvaryintheirsolubilityinwater,frominsolublekeratintothehighlysolublealbumins.All
proteinscanbedenaturedbyheat,strongacid,alkali,alcohol,acetone,ureaandbyheavymetalsalts.Whenproteins
aredenaturedtheyloosetheiruniquestructure,andthereforepossessdifferentchemical,physicalandbiological
properties(ie.inactivationofenzymesbyheat).

2.1.4Classification
Proteinsmabeclassifiedintothreemaingroupsaccordingtotheirshape,solubilityandchemicalcomposition:
a.Fibrousproteins:insolubleanimalproteinswhicharegenerallyveryresistanttodigestiveenzymebreakdown.
Fibrousproteinsexistaselongatedfilamentouschains.Examplesoffibrousproteinsincludethecollagens(main
proteinofconnectivetissue),elastin(presentinelastictissuessuchasarteriesandtendons),andkeratin(present
inhair,nails,woolandhoovesofmammals).
b.Globularproteins:includeallenzymes,antigensandhormoneproteins.Globularproteinscanbefurther
subdividedintoalbumins(watersoluble,heatcoagulableproteinswhichoccurineggs,milk,bloodandmany
plants)globulins(insolubleorsparinglysolubleinwater,andpresentineggs,milk,andblood,andserveasthe
mainproteinreserveinplantseeds)andhistones(basicproteinsoflowmolecularweight,watersoluble,occurin
thecellnucleusassociatedwithdeoxyribonucleicacidDNA).
c.Conjugatedproteins:theseareproteinswhichyieldnonproteingroupsaswellasaminoacidsonhydrolysis.
Examplesincludethephosphoproteins(caseinofmilk,phosvitinofeggyolk),glycoproteins(mucoussecretions),
lipoproteins(cellmembranes),chromoproteins(haemoglobin,haemocyanin,cytochrome,flavoproteins),and
nucleoproteins(combinationofproteinswithnucleicacidspresentinthecellnucleus).

2.2Proteinfunction
Thefunctionofproteinsmaybesummarisedasfollows:
Torepairwornorwastedtissue(tissuerepairandmaintenance)andtorebuildnewtissue(asnewproteinand
growth).
Dietaryproteinmaybecatabolizedasasourceofenergy,ormayserveasasubstratefortheformationoftissue
carbohydratesorlipids.
Dietaryproteinisrequiredwithintheanimalbodyfortheformationofhormones,enzymesandawidevarietyof
otherbiologicallyimportantsubstancessuchasantibodiesandhaemoglobin.

2.3Proteinrequirements
Thestudyofdietarynutrientrequirementsinfishesandshrimphasbeenalmostentirelybasedonstudiescomparable
tothoseconductedwithterrestrialfarmanimals.Itfollowsthereforethatalmostalltheavailableinformationonthe
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dietarynutrientrequirementsofaquaculturespeciesisderivedfromlaboratorybasedfeedingtrialstheanimalsbeing
keptinacontrolledenvironmentathighdensityandhavingnoaccesstonaturalfoodorganisms.

2.3.1Optimumdietaryproteinlevel
Basedonfeedingtechniquespioneeredanddevelopedforterrestrialanimalsthedietaryproteinrequirementsoffish
werefirstinvestigatedintheChinnoksalmon(Oncorhynchustshawytscha)byDelong,etal,(1958).Fishwerefeda
balanceddietcontaininggradedlevelsofahighqualityprotein(casein:gelatinmixturesupplementedwithcrystalline
aminoacidstosimulatetheaminoacidprofileofwholehen'seggprotein)overa10weekperiodandtheobserved
proteinlevelgivingoptimumgrowthwastakenastherequirement(Fig.2).Sincetheseearlystudiestheapproachused
byworkerstodayhaschangedverylittleifatall,withthepossibleexceptionoftheusebysomeresearchersof
maximumtissueproteinretentionornitrogenbalanceinpreferencetoweightgainasthecriterionofrequirement
(Ogino,1980).Dietaryproteinrequirementsarenormallyexpressedintermsofafixeddietarypercentageorasaratio
ofproteintodietaryenergy.

Fig.2.Typicaldoseresponsecurve
Todateover30fishandshrimpspecieshavebeenexaminedinthismannerandtheresultsshowauniformlyhigh
dietaryproteinrequirementintherange2457%,orequivalentto3070%ofthegrossenergycontentofthedietinthe
formofprotein(Table1).Whilstahighproteinrequirementmighthavebeenexpectedforcarnivorousfishspecies,such
asplaice(Pleuronectesplatessa50%Coweyetal,1972)orsnakehead(Channamicropeltes52%WeeandTacon,
1982),thefactthatarelativelyhighproteinrequirementwasalsoobservedintheherbivorousgrasscarp
(Ctenopharyngodonidella4143%Dabrowski,1977)suggeststhattherequirementmayinpartbeafunctionofthe
methodologyusedforitsdetermination.Theusebydifferentworkersofdifferentdietaryproteinsources,nonprotein
energysubstitutes,feedingregimes,fishageclassesandmethodsforthedeterminationofdietaryenergycontentand
dietaryrequirementleaveslittlecommongroundfordirectcomparisonstobemadewithinorbetweenfishspeceis.For
example,thehighproteinrequirementobservedforgrasscarpfry(4143%Dabrowski,1977)almostcertainlyarose
fromallexperimentalfishbeingfedarestrictedration(fishfedonlytwicedaily,andfixedonthelowestrecordedad
libitumfeedtake)andconsequentlyfishfedthelowerproteindietsnotbeingabletoconsumesufficientfeedtomeet
theirdietaryproteinandenergyrequirements.Acriticalreviewofthemethodsusedfortheestimationofdietaryprotein
andaminoacidrequirementsinfishandcrustaceahasbeenmadebyTaconandCowey(1985)andCoweyandTacon
(1983)respectively.
Thehighdietaryproteinrequirementoffishandshrimpisgenerallyattributedtotheircarnivorous/omnivorousfeeding
habit,andtheirpreferentialuseofproteinovercarbohydratesasadietaryenergysource(Cowey,1975).Incontrastto
terrestrialfarmanimalsfishandshrimpareabletoderivemoremetabolizableenergyfromthecatabolismofproteins
thanfromcarbohydrates.

2.3.3Abioticfactorstemperatureandsalinity
Theinfluenceofwatertemperatureonproteinrequirementandfishgrowthhasbeenthesubjectofnumerous
investigations.TheearlystudyofDeLongandcoworkerswithfingerlingChinooksalmon(O.tshawytscha)wassaidto
showanincreaseinthedietaryproteinrequirementfrom40%to55%withanincreaseinwatertemperaturefrom8.3C
to14.4C(DeLong,etal.,1958).Morerecently,asimilarriseindietaryproteinrequirementwasreportedinfingerling
Stripedbass(Moronesaxatilis)from47%to55%withanincreaseinwatertemperaturefrom20.5Cto24.5C(Millikin,
1983Table1).Incontrast,fingerlingrainbowtrout(Salmogairdneri)showednodifferenceingrowthatdietaryprotein
levelsof35%,40%and45%attemperaturesof9C,12C,15Cand18Cinonestudy(Slingeretal.,1977)orin
anotherstudywithtemperaturesof9C,15Cand18C(ChoandSlinger,1978).Althoughdistincttemperatureeffects
wereobservedintermsofgrowth,thegreaterabsoluteneedforproteinatthehigherwatertemperatureswas
apparentlysatisfiedthroughtheincreasedconsumptionofthelowerproteindiets.Theselatterstudiesareinlinewith
thehypothesisthatanincreaseinwatertemperature(uptoanoptimumlevel)isaccompaniedbyanincreasedfeed
intake(Brett,etal.,1969Choubert,etal.,1982),increasedgrowthrateandmetabolicrate(Jobling,1983)andafaster
gastrointestinaltransittime(Fauconneau,etal.,1983RossandJauncey,1981)underconditionswherefoodsupplyis
notlimiting.Theweightofevidenceisthatincreasedwatertemperaturedoesnotleadtoanincreasedprotein
requirement.Inbothcaseswheresucharequirementwasclaimed,theeffectofwatertemperatureondietaryprotein
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requirementwasinvestigatedbycomparingtheresultsobtainedinsuccessiveexperimentsatdifferentwater
temperatures.Inaddition,thesuboptimalgrowthandincreasedfeedintakeobservedwithfishfedthehigherprotein
dietssuggeststhattheadlibitumfeedingregimeemployedinfactledtoarestrictedfeedintake.

TABLE1.Dietaryproteinrequirementoffishandshrimp(expressedas%ofdrydiet)
Dietaryprotein Sizerange Feeding
requirement 1
regime2

Species
FISH
Oreochromis
mossambicus
Oreochromis
niloticus
O.niloticus
O.niloticus

Culture
system

Reference

40

Fingerling

6%bw/d

Indoor/tank

Jauncey(1982)

35

Fry

15%bw/d

Indoor/tank

Santiagoetal.,(1982)

2830
25

Fry/fing.
Fingerling

6%bw/d
3.5%bw/d

Indoor/tank
Indoor/tank

O.niloticus

35

Fingerling

4%bw/d

O.niloticus

1929

Juvenile

3%bw/d

30

Grower

22.5%bw/d

Outdoor/pond Viola&Zohar(1984)b

30

Fingerling

6%bw/d

Indoor/tank

Toledo,Cisneros&Ortiz(1983)

36
56
34
35

Fingerling
Fry
Fingerling
Fingerling

8.8%bw/d
20%bw/d
10%bw/d
5%bw/d

Indoor/tank
Indoor/tank
Indoor/tank
Indoor/tank

T.zilli

3540

Fingerling

4%bw/d

Indoor/tank

Cyprinuscarpio
C.carpio
C.carpio
Ctenopharyngodon
idella

35
34
38

Grower
Fingerling
Fingerling

5%bw/d
Ad.lib.
Ad.lib.

Indoor/tank
Indoor/tank
Indoor/tank

Davis&Stickney(1978)
Winfree&Stickney(1981)
Winfree&Stickney(1981)
Mazidetal.,(1979)
Teshima,Gonzalez&Kanazawa
(1978)
Jauncey(1981)
Muraietal.,(1985)
Ogino&Saito(1970)

4143

Fry

Fixed(?)

Indoor/tank

Dabrowski(1977)

Mugilcapito

24

Fingerling

Ad.lib.

Indoor/tank

PapaparaskevaPapoutsoglou&
Alexis(1985)

O.niloticus/aureus
hybrids
Oreochromis
aureus
O.aureus
O.aureus
O.aureus
Tilapiazilli

DeSilva&Perera(1985)
Wang,Takeuchi&Watanabe(1985)
Teshima,Kanazawa&Uchiyama
Indoor/tank
(1985)
Wannigama,Weerakoon&
Outdoor/cage
Muthukumarama(1985)a

I.punctatus

2942

I.punctatus

45

I.punctatus
I.punctatus

25
36

I.punctatus

25

I.punctatus
Alosasapidissima
Pangasiussutchi
Chanoschanos
Channa
micropeltes
Fugurubripes
Chrysophrysaurata
Moronesaxatilis

35
42.5
25
40

Fixed(1
4%bw/d)
Fixed(1
Grower
4%bw/d)
Fixed(34
Grower
45kg/ha/d)
Grower
Ad.lib.
Fingerling
3%bw/d
Fixed(3
Juvenile
4%bw/d)
Juvenile/grow. 3%bw/d
Fingerling
Ad.lib.
Fry/fing.
10%bw/d
Fry
10%bw/d

52

Grower

50
38.4
47

Fingerling
10%bw/d
Fingerling/juv. Ad.lib.
Fingerling
Ad.lib.

Indoor/tank
Indoor/tank
Indoor/tank

Kanazawaetal,(1980)
Sabaut&Luquet(1973)
Millikin(1983)

M.saxatilis

55

Fingerling

Ad.lib.

Indoor/tank

Anguillajaponica
Micropterus
dolomieui
Micropterus
salmoides
Pleuronectes
platessa
Salvelinusalpinus
Salmogairdneri

44.5

Fingerling

Ad.lib.

Indoor/tank

Millikin(1982)g
Nose&Arai(1973)

45.2

Fry/fing.

Ad.lib.

Indoor/tank

Andersonetal.,(1981)

4041

Fingerling

Ad.lib.

Indoor/tank

Andersonetal.,(1981)

50

Juvenile

Ad.lib.

Indoor/tank

Coweyetal.,(1972)

3643.6
42

Juvenile/grow. Ad.lib.
Grower
Fixed(?)

Indoor/tank
Indoor/tank

Jobling&Wandsvik(1983)
Austreng&Refstie(1979)

S.gairdneri

40

Fingerling/juv. Fixed

Indoor/tank

Satia(1974)h

S.gairdneri

4045

Fingerling/juv. Ad.lib.

Indoor/tank

Ictaluruspunctatus 35

Grower

2%bw/d

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Outdoor/cage Lovell(1972)c

Indoor/tank
Outdoor/tank
Indoor/tank
Indoor/tank

Page&Andrews(1973)
Murai,Fleetwood&Andrews(1979)
Chuapoehuk&Pthisoong(1985)
Limetal.,(1979)

Indoor/tank

Wee&Tacon(1982)

Outdoor/pond Prather&Lovell(1973)d
Outdoor/pond Lovell(1975)e
Indoor/tank
Indoor/tank

Page&Andrews(1973)
Garling&Wilson(1976)

Outdoor/pond Deyoeetal.,(1968)f

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Zeitounetal.,(1973)i
PRAWN
Macrobrachium
rosenbergii

40

PL0.15g

125%bw/d

Indoor/tank

Millikinetal.,(1980)

M.rosenbergii

15

PL0.12g

Fixed

Outdoor/tank Boonyaratpalin&New(1982)j

M.rosenbergii

35

PL0.10g

5%bw/d

Outdoor/tank Balazs&Ross(1976)k

M.rosenbergii

27

PL1.90g

5%bw/d

Outdoor/pond Stanley&Moore(1983)l

Penaeusindicus

3040

PL142day

Fixed

Indoor/tank

P.indicus

43

PL0.41.1g

1015%bw/d

Indoor/tank

Penaeusaztecus

40

PL24135mg 10050%bw/d

Indoor/tank

P.aztecus

4351

PL0.41.3g

Fixed(?)

Indoor/tank

Penaeussetiferus
Penaeus
merguiensis

2832

Juveniles4g

5%bw/d

Indoor/tank

ZeinEldin&Corliss(1976)n
Andrews,Sick&Baptist(1972)

5055

Juv.38g

Fixed(?)

Indoor/tank

AQUACOP(1978)n

P.merguiensis

3442

PL0.3g

Fixed(?)

Indoor/tank

Sedgwick(1979)n

Penaeusmonodon 55

PL2mg

Fixed(?)

P.monodon
P.monodon

34
40

PL5mg
100%bw/d
PL25mg0.7g 10010%bw/d

Outdoor/tank Bages&Sloane(1981)o
Indoor/tank Khannapa(1977)
Indoor/tank Khannapa(1977)

P.monodon

40

Juv.13g

Fixed(?)

Indoor/tank

AQUACOP(1977)n

P.monodon

45.8

PL0.51g

Fixed(?)

Indoor/tank

Lee(1971)n

Penaeusvannamei 36

Juv.420g

Fixed(?)

Indoor/tank

P.vannamei
Penaeusstylirostris
P.stylirostris
Penaeus
californiensis
P.californiensis
Penaeusjaponicus

3035
3035
44

PL32mg0.5g (?)
PL45mg
(?)
PL5mg
(?)

Indoor/tank
Indoor/tank
Indoor/tank

Smithetal.,(1985)n
Colvin&Brand(1977)
Colvin&Brand(1977)
Colvin&Brand(1977)

44

PL5mg

(?)

Indoor/tank

Colvin&Brand(1977)

30
5257

Juv.1g+
PL0.8g

(?)
Ad.lib.

Indoor/tank
Indoor/tank

Colvin&Brand(1977)
Deshimaru&Yone(1978)

P.japonicus

>40

Juv.12g

Fixed(?)

Indoor/tank

P.japonicus

54

PL0.61g

Ad.lib.

Indoor/tank

Balazs,Ross&Brooks(1973)n
Deshimaru&Kuroki(1974)

PL0.10.2g

Fixed(?)

Indoor/tank

Forster&Beard(1973)n

SHRIMP

Palaemonserratus 3040

Bhaskar&Ali(1984)m
Colvin(1976)
Venkataramiah,Lakshmi&Gunter
(1975)

1Fishsizerange:fry00.5g,fingerling0.510g,juvenile1050g,grower50gandabove.
2Feedingregime:%bw/dfixedfeedintakeexpressedasapercentageofbodyweightperday,orAdlibitumfeedingtwotofourtimesdaily.
aNodifferenceinproteinrequirementatthreestockingdensitiesof400,600and800fish/m3,using5m3cages.
b200m2earthenponds,fishdensityof2/m2,pondsalsofertilizedwithpoultrylitteratarateof5kg/pond/week.
cFishstockingdensityof300/m3.
d/eFishstockingdensityof9880/hectare.
fPlasticlinedponds,withfishstockingdensityof30003700/hectare.
gIncreaseddietaryproteinrequirementreportedforfingerlingstripedbassfrom47to55%withanincreaseinwatertemperaturefrom20.5to24.5C.
hFeedintakefixedwithinallgroupstothelowestrecordedAdlibitumfeedintakeobserved.
iProteinrequirementsaidtoincreasefrom40to45%withincreasingsalinity.
jOutdoorconcreteponds,5animals/m2,infrequentwaterexchange,allanimalsfedatsamefixedratebasedonhighestrecordedintake.
koutddorfibreglasstanks,17animals/m2,highwaterexchange.
lAnimalshousedwithinpensinearthenpond,10animals/m2.
mAllanimalsfedatfixedrateof5mgfeed/larvae/day(PL110),15mgfeed/larvae/day(PL1150),and20mgfeed/larvae/day(PL2442).
nAllanimalsfedtoexcessonceortwicedaily.
oDietformulatedto55%crudeprotein,butactuallevelafterdietprocessingwas45%.

Veryfewstudieshavebeenundertakenconcerningtheeffectofsalinityonproteinrequirement.Experimentsconducted
withfingerlingrainbowtrout(aeuryhalinefish)arereportedtoshowanincreaseintheabsolutedietaryrequirementfor
proteinfrom40%to45%withasalinityincreasefrom10to20partsperthousand(Zeitoun,etal.,1973Table1).
However,noincreaseindietaryproteinrequirementwasobservedinasimilarexperimentconductedwithCohosalmon
fingerlings(O.kisutch)Zeitounetal.,1974).Inviewofthespeculativemethodforarrivingatdietaryrequirementfrom
thedoseresponsecurve(Zeitounetal.,1973),andthelackofinformationontheproteinrequirementsofthesefish
speciesinfullstrengthseawater(35partsperthousand),therearenofirmdatademonstratingthattheprotein
requirementsoffishareelevatedwithincreasedsalinity.Thereisnoinformationontheeffectofsalinityontheprotein
requirementofshrimp.

2.4Aminoacids
Althoughover100differentaminoacidshavebeenisolatedfrombiologicalmaterials,only25ofthesearecommonly
foundinproteins.Individualaminoacidsarecharacterisedbyhavinganacidiccarboxygroup(COOH)andabasic
nitrogenousgroup(generallyanaminogroup:NH2).Inviewofthepresenceofbothacidicandbasicgroups,amino
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acidsareamphoteric(ie.havebothacidicandbasicproperties)andconsequentlyactasbuffersbyresistingchangesin
pH.Thechemicalstructureofthemorecommonlyoccurringaminoacidsisshownbelow:

2.5Aminoacidfunction
Aminoacidsoccupyacentralpositionincellularmetabolismsincealmostallbiochemicalreactionsarecatalysedby
enzymescomposedofaminoacidresidues.Aminoacidsareessentialforcarbohydrateandlipidmetabolism,forthe
synthesisoftissueproteinsandmanyimportantcompounds(ie.adrenalin,thyroxine,melanin,histamine,porphyrins
haemoglobin,pyrimidinesandpurinesnucleicacids,choline,folicacidandnicotinicacidvitamins,taurinebilesalts
etc),andasametabolicsourceofenergyorfuel.

2.6Aminoacidrequirements
Fornutritionalpurposes,aminoacidsmaybedividedintotwogroupstheessentialaminoacids(EAA),andthenon
essentialaminoacids(NEAA).EAAarethoseaminoacidsthatcannotbesynthesizedwithintheanimalbodyorata
ratesufficienttomeetthephysiologicalneedsofthegrowinganimal,andmustthereforebesuppliedinareadymade
forminthediet.NEAAarethoseaminoacidsthatcanbesynthesizedinthebodyfromasuitablecarbonsourceand
aminogroupsfromotheraminoacidsorsimplecompoundssuchasdiammoniumcitrate,andconsequentlydonothave
tobesuppliedinareadymadeforminthediet.
ThedietaryEAAforfishandshrimpareasfollows:
Threonine
Leucine
Methionine
Lysine
Arginine

Valine
Isoleucine
Tryptophan
Histidine
Phenylalanine

AlthoughtheNEAAarenotdietaryessentialnutrients,theyperformmanyessentialfunctionsatthecellularormetabolic
level.Theyaretermeddietarynonessentialnutrientsonlybecausethebodytissuescansynthesizethemondemand.
InfactitisoftenquotedthattheNEAAarephysiologicallysoessentialthatthebodyensuresanadequatesupplyby
synthesis.Fromafeedformulationviewpoint,itisimportanttoknowthattheNEAA'scystineandtyrosinecanbe
synthesizedwithinthebodyfromtheEAA'smethionineandphenylalaninerespectively,andconsequentlythedietary
requirementfortheseEAAisdependentontheconcentrationofthecorrespondingNEAAwithinthediet.
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2.6.1Optimumdietaryessentialaminoacidlevels
(a)Doseresponseandcarcassdepositionmethod:ThequantitativeEAArequirementsoffishhavetraditionallybeen
determinedbyfeedinggradedlevelsofeachaminoacidwithinanaminoacidtestdietsoastoelicitadoseresponse
curve(forreviewseeKetola,1982CoweyandLuquet,1983Wilson,1985).Dietaryrequirementisthenusuallytaken
atbreakpointonthebasisoftheobservedgrowthresponse.Inadditiontogrowth,severalworkershavealsoused
freeaminoacidlevelswithinspecifictissuepools(wholeblood,bloodplasmaormuscleKaushik,1979),orthe
oxidationofradioactivelylabelledaminoacids(administeredorallyorbyinjectionWalton,CoweyandAdron,1982)as
thecriterionforestimatingdietaryrequirement.Withintheaminoacidtestdietsusedtheproteincomponentissupplied
almostentirelyintheformofcrystallineaminoacidsorincombinationwithselectedwholeproteinsources(commonly
eithercasein,gelatin,zein,glutenorfishmeal)theaminoacidprofileofthetotalproteincomponentofthedietbeing
controlledsoastosimulatetheaminoacidprofileofaspecificreferenceprotein.
Incontrasttotheabovestandardmethodwherefisharefedgradedlevelsofcrystallineaminoacids,Ogino(1980a)
determinedthequantitativeEAArequirementoffishsimultaneouslyonthebasisofthedailydepositionofindividual
aminoacidswithinthefishcarcass.IntheOginomethodfisharefedadietcontainingawholeproteinsourceofhigh
biologicalvalue,andthedietaryEAArequirementcomputedonthebasisoftheobserveddailyEAAtissuedeposition
value.
Table2summarisestheknownquantitativeEAArequirementsoffishstudiedtodateusingtheabovementioned
techniques.Quantitativedietaryrequirementsforall10EAAshavebeenestablishedforonlyfivefishspecies(common
carpC.carpio,rainbowtroutS.gairdneri,channelcatfishI.puntatus,JapaneseeelA.anguilla,andtheChinook
salmonO.tshawytscha).AtpresentthereisnoquantitativeinformationonthedietaryEAArequirementsofshrimpin
themainthishasbeenduetothepoorgrowthobservedwithshrimpfedsyntheticaminoacidtestdietsandtheinherent
problemsofnutrientleachingduetotheextendedfeedinghabitsofshrimp.
Althoughnumerousindependentstudieshaverecentlybeenperformedontheaminoacidrequirementsofrainbow
trout,significantdifferencesinrequirement(gaminoacid/100gprotein)existwithinandbetweenindividualfishspecies
(Table2).Forexample,differencesoftheorderof65%,72%and114%wereobservedbetweenindependent
laboratoriesforthelysine,arginineandmethioninerequirementoffingerling/juvenilerainbowtrout.Similarly,inter
speciesvariationsrangedfrom22%forvalinetoashighas122%fortryptophan.Whilstonewouldhaveexpectedthe
quantitativeEAArequirementsoffishtodecreasewithageanddecreasingproteinsynthesis(growth),onemaywell
questionwhetherornottheobservedvariationsinrequirementarerealormerelyanartifactofthemethodemployed.In
contrasttothevariationsinrequirementobservedforthesamefishspeciesfedconventionalaminoacidtestdiets,there
wasnosignificantdifferenceintheEAArequirementofcarpandtroutonthebasisofthecarcassdepositionmethodof
Ogino(1980a).However,thedietaryrequirementsobservedarewithintherangereportedforfishfedaminoacidtest
diets(Table2).
Comparedwiththeconventionalmethodoffeedinggradedlevelsofindividualaminoacids,thecarcassdeposition
methodofOgino(1980a)offersnumerousadvantages:
Fisharefedrationsinwhichtheproteincomponentissuppliedintheformofawholeproteinofhighbiological
value.Aminoacidrequirementscanthereforebeascertainedinfishdisplayingoptimalgrowth.
ThedietaryrequirementforalltenEAAscanbedeterminedsimultaneouslyinonesingleexperiment.Using
conventionalaminoacidtestdietsupto10separateexperimentshavetobeperformed,eachexperiment
involvingtheuseofuptosixdietaryregimesemployingvaryingdietaryconcentrationsofthesingleEAAunder
test.
QuantitativeEAArequirementscanequallybeestablishedforfirstfeedingfryandbroodstockfishwithnolossof
precision.

Table2.Quantitativeessentialaminoacid(EAA)requirementsofselectedfishspecies.Valuesare
expressedinorderasapercentageofthedietaryproteinandasapercentageofthedrydiet(the
denominatorbeingthepercentageproteininthediet)
Species
Cyprinuscarpio
Ictaluruspunctatus
Oncorhynchustshawytscha
Oncorhynchusketa
O.keta
Oncorhynchuskisutch
Anguillajaponica
Salmogairdneri
S.gairdneri
S.gairdneri
S.gairdneri
S.gairdneri
S.gairdneri
Dicentrachuslabrax

Simulatedaminoacid(AA)
profileofproteinsource
Casein:gelatin(38:12)
Wholehen'segg
Wholehen'segg
Fishbodyprotein
Fishbodyprotein
Wholehen'segg
?
Wholehen'segg
Wholehen'segg
Fishmeal
Zein:fishmeal(1:1)
Casein:gelatin(3:2)
Whitecodmuscle
Fishmealcomposite

Feedingregime1
Ad.lib.4f/d
3%bw/d,3f/d
Ad.lib.3f/d
Ad.lib.2f/d
Ad.lib.2f/d
Ad.lib.3f/d
?
?
Fixed(?)
4.5%bw/d,3f/d
Ad.lib.4f/d
2%bw/d,3f/d
25%bw/d,4f/d
1.5%bw/d,2f/d

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Initialbodyweight
(g)
0.54.0
210
24
1.1
1.1
24
?
1214
12
1.59
2030
27
514
35

Arginine
3.3(1.3/38.5)
4.3(1.03/24)
6.0(2.4/40)

6.0(2.4/40)
3.9(1.7/42)
>4.0(1.4/35)
5.45.9(2.52.8/47)

3.43(1.2/35)

3.54.0(1.61.8/45)

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Oreochromismossambicus
C.carpio

Fishmealcomposite
4%bw/d,3f/d
1.7
CalculatedonthebasisoftissuedepositionofEAA,withfishfeda
wholeproteinsourceofhighbiologicalvaluehavingaprotein
digestibilityof80%,andafeedingrateof3%bw/dforbothspecies
(carp6274g,2025Ctrout68127g,1518C)

S.gairdneri

<4.0(1.59/40)
3.8(1.52/40)
3.5(1.4/40)

Species

Histidine

Isoleucine

Leucine

Lysine

Methionine2

C.carpio

2.1(0.8/38.5)

2.5(0.9/38.5)

3.3(1.3/38.5)

5.7(2.2/38.5)

2.1(0.8/38.5)a

3.1(1.2/38.5)

2.34(0.56/24)

I.punctatus

1.54(0.37/24)

2.58(0.62/24)

3.5(0.84/24)

5.1(1.5/30)

O.tshawytscha

1.8(0.7/40)

2.2(0.9/41)

3.9(1.6/41)

5.0(2.0/40)

O.keta
O.keta
O.kisutch

1.6(0.7/40)

1.7(0.7/40)

4.8(1.9/40)

A.japonica

1.9(0.8/42)

3.6(1.5/42)

4.8(2.0/42)

4.8(2.0/42)

S.gairdneri
S.gairdneri

3.7(1.3/35)
6.1(2.9/47)

S.gairdneri

S.gairdneri

S.gairdneri

S.gairdneri

4.3(1.95/45)

D.labrax

O.mossambicus

4.1(1.62/40)

C.carpio
S.gairdneri

1.4(0.56/40)
1.6(0.64/40)

2.3(0.92/40)
2.4(0.96/40)

4.1(1.64/40)
4.4(1.76/40)

5.3(2.12/40)
5.3(2.12/40)

Species

Phenylalanine4

C.carpio
I.punctatus
O.tshawytscha
O.keta
O.keta
O.kisutch
A.japonica
S.gairdneri
S.gairdneri
S.gairdneri
S.gairdneri
S.gairdneri
S.gairdneri
D.labrax
O.mossambicus
C.carpio
S.gairdneri

1.34(0.32/24)
1.5(0.6/40)

Methionine3

2.1(0.9/42)d

2.9(1.2/42)

1.572.14(0.550.75/35)e

1.0(0.5/50)

2.0(1.0/50)

12(0.51/50)

<1.33(0.53/40)
1.6(0.64/40)
1.8(0.72/40)

Phenylalanine5

Threonine

Tryptophan

Valine

Reference

6.5(2.5/38.5)

3.9(1.5/38.5)

0.8(0.3/38.5)

3.6(1.4/38.5)

Nose(1979)

2.0(0.5/24)

5.0(1.2/24)

2.2(0.53/24)

0.5(0.12/24)

2.96(0.71/24)

NRC(1983)

4.1(1.7/41)

2.2(0.9/40)

0.5(0.2/40)

3.2(1.3/40)

NRC(1983)

3.0(1.2/40)

0.73(0.29/40)
0.5(0.2/40)

Akiyamaetal,(1985)
Akiyamaetal,(1985a)
Klein&Halver(1970)

5.2(2.2/42)

3.6(1.5/42)

1.0(0.4/42)

3.6(1.5/42)

Nose(1979)

3.3(1.32/40)
3.4(1.36/40)

0.45(0.25/55)

0.6(0.24/40)
0.5(0.2/40)

2.9(1.16/40)
3.1(1.24/40)

Kimetal.,(1983)
Ketola(1983)
Rumseyet.al.(1983)
Kaushik(1979)
Waltonetal,(1982)
Waltonetal,(1984)
Thebaultetal.,(1985)
Jackson&Capper(1982)
Ogino(1980a)
Ogino(1980a)

3.4(1.3/38.5)

2.9(1.2/42)k

2.9(1.16/40)
3.1(1.24/40)

1Feedingregime:indicatesfeedinglevelandnumberoffeedingsperday.
2Inthepresenceofdietarycystine
a2%
b0.24%
c1%
d1%
e0.3%
f2%
g0.74%
h1%
3Intheabsenceofdietarycystine.
4Inthepresenceofdietarytyrosine
h1%
i1%
j0.4%
k2%
5Intheabsenceofdietarytyrosine.

(b)Carcassanalysismethod:Interestingly,recalculationofthedataobtainedbyOgino(1980a)showsthatthereisno
differencebetweentherelativeproportionsofindividualEAAsrequiredinthedietandtherelativeproportionsofthe
same10EAAspresentwithinthefishcarcass(TaconandCowey,1985).Asimilarrelationshiphasalsobeenseenin
thegrowingpigandchick(Boorman,1980),andtoalesserextentwithinthefourfishspeciesforwhichEAA
requirementshavebeendeterminedusingaminoacidtestdiets(Fig.3).Similarly,WilsonandPoe(1985)obtaineda
regressioncoefficientof0.96whentheEAArequirementpatternforthechannelcatfishwasregressedagainstthe
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wholebodyEAApatternfoundina30gchannelcatfish.Sincetheaminoacidcompositionoffishbodytissuedoesnot
differgreatly(ifatall)betweenindividualfishspecies(NjaaandUtne,1982WilsonandCowey,1985),itfollows,
therefore,thatthepatternofrequirementfordifferentspecieswillalsobesimilar.Althoughnotproven,itisnot
unreasonabletosupposethatasimilarrelationshipalsoexistsforshrimpsandfreshwaterprawns.Forcomparative
purposesTable3presentsthedietaryEAArequirementpatternforfish,asdeterminedbyOgino(1980a),togetherwith
thecarcassEAApatternofwholefishbodytissue,Penaeusjaponicus

Figure3RelationshipbetweenpatternofEAArequirementsfoundbyfeedingexperimentsusingaminoacidtestdiets
withcarp(),Japaneseeel(),channelcatfish()andchinooksalmon(o)andthepatternofthesameaminoacidsin
fishcarcass.Thelevelofeachaminoacidisrepresentedasapercentageofthesumofall10EAA'sineachpattern.
Thelinerepresentscoincidenceofrequirementandtissuepatterns.larvaeandjuveniles,Penaeuspaulensisjuveniles,
shortneckedclamtissue(Venerupisphilippinarumregardedasanexcellentandidealnaturalfoodformarineshrimp),
andthetailmuscleofMacrobrachiumrosenbergii.Onthebasisoftheaminoacidprofilespresenteditwouldappearthat
shrimphaveahigherdietaryrequirementforarginine,tryptophanandtyrosine,andalowerdietaryrequirementfor
valine,threonineandlysinethanfish.

Table3.MeanfishdietaryEAArequirementpattern(%)andEAApatterninbodytissueofwholefish,short
neckedclam,marineshrimp,andthefreshwaterprawn.

EAA
Threonine
Valine
Methionine
Isoleucine
Leucine
Phenylalanine
Lysine
Histidine
Arginine

Fish
requirement
(Ogino,
1980a)
10.6
9.5
5.4
7.5
13.5
9.5
16.8
4.8
11.6

Wholebody
fishtissue
(Wilson&
Cowey,1985)
9.2
9.5
5.5
8.0
14.6
8.3
16.9
5.2
12.3

Shortnecked
clamtissue
(Deshimaruet
al,1985)
9.6
8.5
5.4
6.8
14.0
7.7
14.7
4.4
15.5

P.japonicus
larvae(Teshima,
Kanazawa&
Yamashita,1986)
5.9
8.8
5.7
9.1
12.1
8.6
13.1
4.5
14.1

P.japonicus
juveniles
(Deshimaru&
Shigeno,1972)
8.2
8.3
5.4
8.6
15.0
9.0
15.8
4.5
15.2

P.
Paulensis
M.rosenbergiitail
juveniles
muscle
(unpublished (Farmanfarmian&
data)
Lauterio,1980)
6.7
7.5
13.6
7.3
7.0
6.5
6.9
7.4
12.6
14.8
9.2
7.3
15.4
17.1
4.4
4.5
14.3
20.6

Tryptophan

1.7

1.7

2.7

6.3

NA

NA

Cystine*

2.7

2.0

2.7

2.4

2.1

3.0

NA
NA

Tyrosine*

6.5

6.6

7.8

9.2

7.8

6.7

6.6

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NAdatanotavailable(notanalysed).
*Nonessentialaminoacids.AllvaluesareexpressedasapercentageoftotalEAApluscystineandtyrosine.

IntheabsenceoffirmquantitativeinformationonthedietaryEAArequirementsofshrimpandthemajorityoffarmed
fishspecies,dietaryrequirementcaninitiallybecomputedonthebasisofthecarcassEAApatternpresentwithin35%
oftheknowndietaryproteinrequirementsofthesaidspeciesonageneralbasisEAAs(includingtheNEAAscystine
andtyrosine)constituteabout35%ofthetotaldietaryproteinrequiredbyfish(Table2).Thusifashrimporfishisknown
tohaveadietaryproteinrequirementof45%,thendietaryEAArequirementwouldbecomputedonacarcassEAA
patternof35%ofthedietaryproteinlevel.Forexample,ifthecarcassEAApatternforlysineis16.9%ofthetotalEAA
pluscystineandtyrosinepresent,thendietaryrequirementlevelforlysinewouldbe
thedrydiet(ie.45%proteinfishration).

or2.66%of

AsaguidelineTable4presentsthecalculateddietaryEAArequirementsoffishandshrimpatvaryingdietaryprotein
levelsbasedonthemeancarcassEAApatternofwholefishtissueandshortneckedclamtissuerespectively(short
neckedclamtissueisusedhereintheabsenceofameancarcassEAApatternforshrimp).

Table4.CalculateddietaryEAArequirementsoffishandshrimpatvaryingdietaryproteinlevels(values
areexpressedasapercentofthedrydiet)

EAA
FISH

25

30

Dietaryproteinlevel(%)
35
40
45
50

55

Arginine
Histidine
Isoleucine
Leucine
Lysine
Methionine

1.07
0.45
0.70
1.28
1.49
0.48

1.29
0.55
0.84
1.53
1.77
0.58

1.51
0.64
0.98
1.79
2.07
0.67

1.72
0.73
1.12
2.04
2.37
0.77

1.94
0.82
1.26
2.30
2.66
0.87

2.15
0.91
1.40
2.55
2.96
0.96

2.37
1.00
1.54
2.81
3.25
1.06

12.3
5.2
8.0
14.6
16.9
5.5

Cystine*

0.17

0.21

0.24

0.28

0.31

0.35

0.38

2.0

Phenylalanine

0.73

0.87

1.02

1.16

1.31

1.45

1.60

8.3

Tyrosine*

0.58

0.69

0.81

0.92

1.04

1.15

1.27

6.6

Threonine
Tryptophan
Valine
SHRIMP

0.80
0.15
0.83

0.97
0.18
1.00

1.13
0.21
1.16

1.29
0.24
1.33

1.45
0.27
1.50

1.61
0.30
1.66

1.77
0.33
1.83

9.2
1.7
9.5

Arginine
Histidine
Isoleucine
Leucine
Lysine
Methionine

1.36
0.38
0.59
1.22
1.29
0.47

1.63
0.46
0.71
1.47
1.54
0.57

1.90
0.54
0.83
1.71
1.80
0.66

2.17
0.62
0.95
1.96
2.06
0.76

2.44
0.69
1.07
2.20
2.31
0.85

2.71
0.77
1.19
2.45
2.57
0.95

2.98
0.85
1.31
2.69
2.83
1.04

15.5
4.4
6.8
14.0
14.7
5.4

Cystine*

0.24

0.28

0.33

0.38

0.42

0.47

0.52

2.7

Phenylalanine

0.67

0.81

0.94

1.08

1.21

1.35

1.48

7.7

Tyrosine*

0.68

0.82

0.96

1.09

1.23

1.37

1.50

7.8

Threonine
Tryptophan
Valine

0.84
0.24
0.74

1.01
0.28
0.89

1.18
0.33
1.04

1.34
0.38
1.19

1.51
0.42
1.34

1.68
0.47
1.49

1.85
0.52
1.64

9.6
2.7
8.5

CarcassEAA
pattern(%)

1CarcassEAApatternofwholefishtissue(Wilson&Cowey,1985)
2CarcassEAApatternofshortneckedclam(Deshimaruetal.,1985)
*Nonessentialaminoacids

2.6.2Utilizationoffreeaminoacids
Fishorjuvenileshrimpfedrationsinwhichasignificantproportionofthedietaryproteinissuppliedintheformoffree
orcrystallineaminoacidsgenerallydisplaysuboptimalgrowthandfeedconversionefficiencycomparedwithanimals
fedproteinboundaminoacidsorwholeproteins(Wilsonetal.,1978Robinsonetal.,1981Yamadaetal.,1981
Waltonetal.,1982Deshimaru,1981Deshimaru&Kuroki,1974a,1975).
Ingeneral,dietaryfreeaminoacidsaremorerapidlyassimilatedinfishthanproteinboundaminoacids.Experiments
withrainbowtrout(Yamadaetal.,1981),commoncarp(Plakasetal.,1980)andtilapia(OreochromisniloticusYamada
etal.,1982)fedfreeaminoacidtestdietsshowedthatpeakplasmaaminoacidconcentrationsoccurredsooner(12
24h,24h,2h,respectively)thanwithanequivalentcaseinbaseddiet(2436h,4h,4h,respectively).Furthermore,in
carp.individualfreeaminoacidsappeartobeabsorbedatvaryingratesfromthegastrointestinaltract,and
consequentlypeakplasmaconcentrationsofindividualaminoacidsdonotoccursimultaneously(Plakasetal.,1980).In
juvenileshrimpthesituationappearstobethereverse.Forexample,Deshimaru(1981)showedthattheassimilation
rateofdietaryfreearginineintomuscleproteinbyPenaeusjaponicusjuvenileswasextremelylow(assimilationless
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than0.6%)comparedtothatofproteinboundarginine(assimilationabove90%).However,althoughDeshimaru(1981)
reportednobeneficialeffectongrowthoffreeaminoacidsupplementeddietswithP.japonicusjuveniles,recentstudies
havedemonstratedthatthelarvaeofthesamespeciesiscapableofutilizingaminoacidsupplementeddietsforgrowth
(Teshima,Kanazawa&Yamashita,1986).
Foroptimalproteinsynthesistooccur,itisessentialthatallaminoacids(whethertheybederivedfromwholeproteinsor
aminoacidsupplements)arepresentedsimultaneouslytothetissue.Ifsuchanequilibriumisnotachieved,thenamino
acidcatabolism(breakdown)ensueswithconsequentlossofgrowthandandfeedefficiency.Forthosewarmwaterfish
specieswhichdisplayarapiduptakeandassimilationoffreeaminoacids,itisthereforeessentialthateither(1)the
releaseorabsorptionoffreeaminoacidsfromthedietisreducedsoastominimisethevariationsinabsorptionrate
observedbetweenfreeandproteinboundaminoacids(achievedbycoatingindividualaminoacidswithcasein,zeinor
nylonproteinmembranesMuraietal.,1982Teshima,Kanazawa&Yamashita,1986)or(2)thatthefrequencyoffeed
presentationisincreasedfromtwoorthreefeedsperdaytoupto18feedsperdaysoastominimisethevariations
observedinplasmaaminoacidconcentration(Yamada,Tanaka&Katayama,1981).

2.6.3Aminoacidcompositionandproteinquality
Onthebasisoftheabovediscussionsitisevidentthattheproteinqualityofafeedingredientisdependentuponthe
aminoacidcompositionoftheproteinandthebiologicalavailabilityoftheaminoacidspresent.Ingeneral,thecloserthe
EAApatternoftheproteinapproximatestothedietaryEAArequirementofthespecies,thehigheritsnutritionalvalue
andutilization.Forexample,Table5presentsthechemicalscoreorpotentialproteinvalueofsomecommonlyused
feedproteins.Chemicalscoresof100indicatethatthelevelofaparticularEAAwithinthefeedproteinisidenticaltothe
dietaryEAArequirementlevelforfish(whenexpressedasapercentageofthetotalEAAspluscystineandtyrosine)as
determinedbyOgino(1980a).ThechemicalscoreoftheproteinistakentobethepercentageoftheEAAingreatest
deficitrelativetothedietaryrequirementpattern.Thismethodofassessingproteinqualityisbasedontheconceptthat
thenutritivevalueofaproteindependsprimarilyontheamountoftheEAAingreatestdeficitinthatprotein,compared
toareferenceprotein(inthiscasethereferenceproteinisthedietaryEAArequirementsoffishasdeterminedby
Ogino.1980a).ItcanbeseenfromTable5thatcomparedtofishmealorfishmuscle,whichhasawellbalancedEAA
profileandhighchemicalscore(c.80),themajorityofproteinsourcespresentedhaveaminoacidimbalanceswhich
renderthemunsuitableasasolesourceofdietaryproteinforfishwithincompletedietsintendedforintensivefarming
systems.TheaimoffeedformulationistomixproteinsofvariousqualitiestoobtainthedesiredEAApatternofthefish
orshrimpspeciesinquestion(completedietfeeding).
However,theaboverelationshipbetweenproteinqualityandEAApatternwillonlyholdtrueiftheindividualaminoacids
areequallybiologicallyavailabletotheanimal.Forexample,undercertainconditionssomeoftheaminoacidsmaybe
unavailablebecausetheproteinsinthedietareincompletelydigested.Thus,forcarnivorousfishandshrimpspecies,
thecellulosecellwallwithinplantproteinsourcesmayrendertheproteinspresentwithinthecellinaccesibletothe
digestiveenzymes.Inothercases,digestionmaybehinderedbythepresenceofenzymeinhibitorswithinthefood
proteintrypsininhibitorwithinrawsoybeans.Althoughitispossibletoinactivatetheseinhibitorsbymoderateheat
processing,underconditionsofexcessiveheattreatmentproteinsbecomemoreresistanttodigestionduetopeptide
bondformationoccurringbetweenthesidechainsoflysineanddicarboxylicacid.Thefreeepsilonaminogroupsof
lysineareparticularlysusceptibletoheatdamage,formingadditioncompoundswithnonproteincompounds(ie.
reducingsugarssuchasglucose)presentinthefoodstuff(Cockerell,Francis&Halliday,1972).Thisreactionisknown
astheMaillardreaction,andrendersthelysinebiologicallyunavailable.Substancesotherthanreducingsugarswhich
areknowntoreactwiththefreeepsilonaminogroupoflysineincludegossypolphenolbasedcompoundpresentin
cottonseedmeal.Anestimateofthebiologicalavailabilityofaminoacidswithinfeedproteins,andhenceanindicatorof
proteinquality,canbemadebychemicallymeasuringthefreeoravailablelysinecontentofthefeedprotein(Cowey,
1979).

Table5.Chemicalscoreandlimitingessentialaminoacidsofsomecommonlyusedfeedproteins1
1stlimiting
aminoacid

Feedstuffs

Source2

Thr

Chickpea

64* 89

86

72

100 166 129 Met

Mungbean

59* 110 54* 48* 127 121 124 94

79

114 123 123 Cys

Cowpea

65*

103 61* 59* 116 116 116

100 75

127 134 129 Cys

Yellowlupin

66* 81

Limabean

84

110 57* 74

Broadbean

77

103 30* 41* 115 118 98

118 77

98

Haricotbean

80

103 43* 67* 120 121 118

83

127 104 129 Met

Safflower

68* 125 63* 141

Crambe

98

121 67* 218

117 104 83

86

66* 104 111

Palmkernel

62*

113 94

133

95

89

72

78

41*

Cottonseed

65* 102 52* 118

92

94

122 89

Sunflower

65* 124 83

137

115 104 109 91

42* 119 159 165 Lys

Linseed

71

156

111

90

43* 100 174 182 Lys

Sesame

58* 98

91

105 86

Coconut

65* 114 61* 96

115 112 95

Groundnut

55* 99

117 100 107 117 53* 100 196 141 Met

Val Met Cys Ils

63* 104

20* 126

122 93

109 148

39* 133

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Leu Phe Tyr

119 110 113

117 125 85

94

Lys His

64* 117 192 135 Met

135 118 125 106 72

111

99

Arg Trp

92

112 98

106 Met

160 118

101 100 43* 121 181 118

105 92

98

Met
Lys

200 Lys

225 311

Lys

52* 117 205 141 Met/Lys

114 33* 114 211 153 Lys

92

37* 81

217 123 Lys

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Rapeseed

93

118 83

77

74

131 112 159 Cys

Soybean

74

Potatoproteinconcentrate

89

101 46* 130

128 115 105 97

76

106 123 176 Met

125 63* 96

128 120 112

149 74

73

73

118

Leafproteinconcentrate

84

127 57* 56* 112 120 122 129 71

90

96

141 Cys

Spirulinamaxima

87

136 52* 30* 159 118 105 123 55* 75

111

165 Cys

Saccharomycescerevisiae

93

116 63* 85

139 112 91

Torulopsisutilis

94

118 54* 81

144 98

M.methylotrophus

97

134 89

59* 115 107 115

Wholehen'segg
Fishmuscle
Fishmeal(herring)
Fishmeal(white)

8
9
4
4

77
83
76
81

125
98
127
106

130
85
78
93

Fishproteinconcentrate

83

110 118 63* 127 109 85

Fishsilage

10

98

122 72

101 129 120 94

Wholeshrimpmeal

83

97

109 85

112 106 95

105 86

73

Meatandbonemeal

77

128 59* 89

109 113 88

60* 86

100 150 88

Bloodmeal

69* 158 33* 52* 24* 162 124 69* 89

214 62* 123 Ils

Livermeal

76

135 72

89

105 121 109 106 71

98

105 153 Lys

Poultrybyproductmeal

76

125 81

141

132 123 80

60*

71

87

134 112

Hydrolysedfeathermeal

91

164 24* 289

131 124 78

86

33* 50* 147 76

Wormmeal

11

107 99

Houseflylarvae

12

75

100
98
109
104

70

113 116 94

132
108
117
121

109
110
107
109

108 86

106 89

141 Met

137 117 84

104 86

118

Met

138 71

83

84

118

Cys

98
117
95
94

92
121
96
94

96
97
111
116

135
135
123
129

Thr
Phe
Thr
Thr/Phe

90

95

153 Cys

97
80
80
81

106 52* 112 124 84

103 72

52* 96

90

Met

78
101
89
90

103 92
98

121 108 59* Trp

134 106 His
Met

Tyr
Met

108 79

125 98

82

128 218 77

127 82

147 Cys

Cys

1Scoresbasedoncomparisonwiththemeanessentialaminoacidrequirementsofrainbowtroutandcarp(Ogino,1980).MeanEAArequirement(expressedas
%oftotalEAA)being:threonine10.6valine9.5methionine5.4cystine2.7isoleucine7.5leucine13.5phenylalanine9.5tyrosine6.5lysine16.8histidine4.8
arginine11.6andtryptophan1.7
2Source:1Kay(1979)2Gohl(1980)3BoltonandBlair(1977)4NationalResearchCouncil(1983)5TunnelAVEBEStarchesLtd.,UK6Coweyetal.
(1971)7Unpublisheddata8CoweyandSargent(1972)9ConnellandHowgate(1959)10Jackson,KerrandCowey(1984)11Tacon,Staffordand
Edwards(1983)12Spinelli(1980)
*Limitingessentialaminoacids(presentbelow30%meanfishrequirement)

2.7Evaluationofproteinquality
Apartfromchemicallymeasuringaminoacidsandtheiravailabilitywithinfeedproteins,therearemanybiological
methodsofevaluatingproteinquality:
Specificgrowthrate(SGR)Therateofgrowthofananimalisafairlysensitiveindexofproteinqualityundercontrolled
conditionsweightgainbeingproportionaltothesupplyofessentialaminoacids.DailySGRcanbecalculatedbyusing
theformula:

Foodconversionratio(FCR)Definedasthegramsoffeedconsumedpergramofbodyweightgain.
*Asfedbasisie.dryweight
**Wetorfreshweightgain

Foodefficiency(FE)Definedasthegramsofweightgainedpergramoffeedconsumed.Unitsofexpressionasabove.

Proteinefficiencyratio(PER)Definedasthegramsofweightgainedpergramofproteinconsumed.
*Withthismethodnoallowanceismadeformaintenance:ie.method
assumesthatallproteinisusedforgrowth.

Apparentnetproteinutilization(ApparentNPU)Definedasthepercentageofingestedproteinwhichisdepositedas
tissueprotein.
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wherePbisthetotalbodyproteinattheendofthefeedingtrial,Paisthetotalbodyproteinatthebeginningofthe
feedingtrial,andPiistheamountofproteinconsumedoverthefeedingtrial.Inthiscalculationnoallowanceismadefor
endogenousproteinlosses.Incontrasttothepreviousmethodsofevaluatingproteinquality,thismethodrequiresthata
representativesampleofanimalsbesacrificedatthebeginningandendofthefeedingtrialforcarcassproteinanalysis.
Themaindrawbackofthesemethodsofpredictingdietorproteinqualityisthattheyhavetobeperformedunder
controlledexperimentalconditionsintheabsenceofnaturalfoodorganisms.Consequently,thesemethodscanonlybe
usedwithinintensiveorclearwaterculturesystems.

2.8Nonproteinnitrogenousconstituents
Aminoacidsareimportantnotonlyasbuildingblocksofproteinbutastheprimaryconstituentsornitrogenprecursors
formanynonproteinnitrogencontainingcompounds.Table6listssomeofthemorebiologicallyimportantnonprotein
nitrogenouscompoundsthatoriginatefromaminoacids.

Table6.Nonproteinnitrogenousconstituentsderivedfromaminoacidsinanimals1
Nitrogenouscompound

Aminoacidprecursor

Physiologicalfunctionofcompound

Purines&pyrimidines2
Creatine
Bileacids(glycoholic&taurocholicacids)
Thyroxine,epinephrine&norepinephrine
Ethanolamine&choline
Histamine
Serotonin
Porphyrins
Niacin
Melanin

Glycine&asparticacid

Constituentsofnucleotidesandnucleicacids

Glycine&arginine
Glycine&cysteine
Tyrosine
Serine
Histidine
Tryptophan
Glycine
Tryptophan
Tyrosine

Energystorageascreatinephosphateinmuscle
Bileacids,aidinfatdigestionandabsorption
Hormones
Constituentsofphospholipids
Avasodepressor
Transmissionofnerveimpulses
Constituentsofhaemoglobinandcytochromes
Vitamin
Pigmentofskinandeyes

1Lloyd,McDonald&Crampton(1978)
2Pyrimidineandpurinehavebeensuggestedtobeessentialdietarynutrientsfornewlyhatchedfishlarvae(Dabrowski&Kaushik,1982)andArtemiasalina
(Hernandorena,1983)respectively.

2.9Proteinandaminoacidpathology
2.9.1DietaryEssentialAminoAcidDeficiency
AlthoughallfishexaminedtodatedisplayreducedgrowthwhenfedEAAdeficientdiets,thefollowingadditionalgross
anatomicaldeficiencysignshavebeenobservedunderexperimentalconditionswithjuvenilefishfedsyntheticrations
deficientinoneormoreEAAs:
LimitingEAA

Fish

Lysine

Tryptophan

Salmogairdneri
Cyprinuscarpio
S.gairdneri
Salmosalar
S.gairdneri

Miscellaneous

Oncorhynchusnerka
O.keta

Methionine

C.carpio

Deficiencysigns1
Dorsal/caudalfinerosions(1,2)increasedmortality(2)
Increasedmortality(3)
Cataract(4,5)
Cataract(6)
Scoliosis2(710)lordosis2(7,10)renalcalcinosis(8)cataract(7,9)caudal
finerosiondecreasedcarcasslipidcontent(9)elevatedCa,Mg,NaandK
carcassconcentration(7)
Scoliosis(11)
Scoliosis/lordosis(12)
Increasedmortalityandincidenceoflordosisobservedwithdietarydefi
cienciesofleucine,isoleucine,lysine,arginineandhistidine(3)

11Walton,CoweyandAdron(1984)2Ketola(1983)3Mazidetal.(1978)4Walton,CoweyandAdron(1982)5Postonetal.(1977)6Barash,Poston
andRumsey(1982)7Waltonetal.(1984)8KloppelandPost(1975)9PostonandRumsey(1983)10Shanks,GahimerandHalver(1962)11Halverand
Shanks(1960)Akiyamaetal.(1985a).
2Curvatureofthevertebralcolumn

UnderintensivefarmingconditionsdietaryEAAdeficienciesmayarisefromoneoffourpossibleroutes:
PoorfeedformulationarisingfromtheuseofdisproportionateamountsoffeedproteinswithnaturalspecificEAA
deficiencies(Table5).

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Dietaryimbalancesmayalsoarisefromthepresenceofdisproportionatelevelsofspecificaminoacidsincluding
leucine/isoleucineantagonisms,andtoalesserextentarginine/lysineandcystine/methionineantagonisms.For
example,bloodmealisarichsourceofvaline,leucineandhistidine,butisaverypoorsourceofmethionineand
isoleucine.However,inviewoftheantagonisticeffectofexcessleucineonisoleucine,animalsfedhighdietary
levelsofbloodmealsufferfromanisoleucinedeficiencycausedbyanexcessofdietaryleucine(Taylor,Coleand
Lewis,1977).Althoughsimilarantagonismshavealsobeenreportedforcystine/methionine(useofhydrolysed
feathermealIchhponaniandLodhi,1976)andarginine/lysine(Harper,BenevengaandWohlhueter,1970)in
terrestrialfarmanimals,theyhavenotbeenfoundtooccurinfishfedsyntheticaminoaciddietcombinations
(Robinson,WilsonandPoe,1981).
DietaryEAAdeficienciesmayarisefromexcessiveheattreatmentoffeedproteinsduringfeedmanufacture.
DietaryEAAdeficienciesmayarisefromthechemicaltreatmentoffeedproteinswithacids(silageproduction)or
alkalies,duetothelossoffreetryptophanandlysine/cystinerespectively(Kies,1981).
DietaryEAAdeficienciesmayarisefromtheleachingoffreeandproteinboundaminoacidsintothewater.For
example,Grabner,WieserandLackner(1981)reportedtheloss,throughleaching,ofalmostallthefreeand
aboutonethirdofthefreeplusproteinboundaminoacidsfromfrozenorfreezedriedzooplankton(Artemiasalina
andMoinaspp.)aftera10minutewaterimmersionperiodat9C.Considerablelossesofwatersolubleamino
acidshavealsobeenobservedincarpduringmastication(YamadaandYone,1986).However,theproblemof
nutrientleachingofwatersolublematerialsisprobablygreatestforcrustaceansduetotheirveryslowdemersal
feedinghabitandnecessitytomasticatetheirfoodexternallypriortoingestion(Farmanfarmaian,Lauterioand
Ibe,1982).Forexample,BagesandSloane(1981)reporteda28%lossofdietaryproteinduringthepreparation
andrehydrationofadryalginateboundshrimpdietpriortofeeding,andatotalproteinlossof3947%afterasix
hourimmersionperiodinseawater.Ingeneralnutrientlossesaregreaterinfreshwaterthaninseawater(Balazs,
RossandBrooks,1973).However,problemsofnutrientleachingcanbeminimisedbyusinganappropriate
feedingregime(ie.regularratherthaninfrequentfeedingSedgwick,1979)andasuitabledietbindingormicro
encapsulationtechnique(Goldblatt,ConklinandDuaneBrown,1980Jonesetal.,1976).

2.9.1Toxicnonessentialaminoacids
Nutritionalpathologiesmayalsoarisefromtheingestionoffeedproteinscontainingtoxicaminoacids.Commonlyused
feedproteinswhichareknowntocontaintoxicaminoacidsincludealkalitreatedsoybean(toxicaminoacid
lysinoalanine),thelegumeLeucaenaleucocephalaoripilipil(toxicaminoacidmimosine),andthefababeanVicia
faba(toxicaminoaciddihydroxyphenylalanine).

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