N

utr
it
ioni
nTr
opi
calA
qua
cul
tur
e
E
ssen
tia
lso
ffishnutr
it
ion
,feeds
,andf
eed
ing
o
ftropi
calaqua
ticsp
ecie
s
Ed
ite
dby
O
sen
iM.M
il
lam
ena
,Re
li
card
o M
.Co
los
o,andF
el
ic
ita
sP.P
ascu
al

Aqua
cul
tureDep
artmen
t
SOUTHEASTAS
IANFISHERIESDEVELOPMENTCENTER
T
igbauan
,I
lo
ilo
,Phi
lipp
ines
M
ay2
002
Nu
trit
ioninTrop
ica
l
Aquacu
ltu
re
Essen
tia
lsoff
ishnut
rit
ion
,f e
eds
,andfeed
ing
o
ftrop
ica
laquaticspe
cie
s

Ed
itedby
O
sen
iM. M
il
lam
ena
,Re
lica
rdo M. Co
loso
, and F
elic
ita
sP. Pa
scua
l

Aquacu
lture Depar
tmen
t
SOUTHEAST AS
IAN F
ISHER
IES DEVELOPMENT CENTER
T
igbauan
,Ilo
ilo
, Ph
ilipp
ines

May 2002
i
i

Nutri
tioninT rop
ica
lAquaculture
Essent
ialso
ffishnut
rit
ion
,feeds
,andfeed
ingo
ftrop
ica
laqua
ticspec
ies

ISBN971
-8511
-58
-X

Copyright 2002
Aquaculture Depa
rtment
SoutheastAsianFisher
ies Deve
lopmen
t Cen
ter
Tigbauan,Ilo
ilo
,Phil
ippines

ALLRIGHTSRESERVED
Nopartofthispubli
cation mayberep roducedortransm i
ttedinanyfo rmorby
any means,elec
tronicor mechani
cal
,inc ludingphotocopy,record
ingo rany
info
rma t
ions to
rageo rre
trieva
lsyst
em ,w i
thoutthepe rmissionin w
rit
ingfrom
thepublishe
r.

Fo
rcommen
tsandinqu
ir
ies
:

Tra
iningandInfo
rma tion D
ivision
SEAFDECAquacu lture Department
T
igbauan,I
loi
lo,Phil
ippines

Fax :(63-33)3351008,3362891
Emai
l :t [email protected]
.org.ph
devcom@aqd .
seafdec.org
.ph
webs
ite :h
ttp://www.seafdec.o
rg.ph

Cove
rdes
ignby Ne
lsonV
. Go
lez
 i
ii

Fo
rewo
rd
Aquacu lture has been recogn ized asthefas tes tg rowingfood p roduc ing
secto r, con tribu ting sign if
ican tlyto na tiona l econom ic deve lopmen t,food
supp ly, andfood secur ity. A longs ide aquacu l
ture deve lopmen ta rethe
fishe ries schoo lstha tsupp lythe know ledge needs o ftheindus try. These
fisher ies schoo ls need current educat iona l mater ials re levant to
aquacu ltureinthet ropics. Th is need promp ted SEAFDEC Aquacu lture
Depar tmen t(SEAFDEC AQD ) to es tablishthetex tbook wr iting p rog ram .
It wasa lsoes tab l
ishedtofu lf
ill SEAFDEC AQD ’s manda teo fd issem ina ting
up- to-da te know ledge in aquacu lture techno logy . Th is tex tbook on
Nu t
r itionin Trop ica l Aquacu l
tureis nowthe secondin a se ries.
In deve lop ing coun t
ries,the g row th o f aquacu ltureis o ften hampe red
by alack o fexpe rtisein aquacu lture nu t
ri t
ion and knowhow on aqua feed
formu lation and prepa ra t
ion as we ll as eva lua tion o fthe nu tritiona l and
econom ic va lue o ffeeds . Th is book isin tendedto p rovide afounda tion
ofsc ien tific know ledge and unde rstand ingon nu trit
ion ,feeds , andfeed ing
with emphas is on trop ical aquacu lture spec ies. I tisimpor tan ttha t
studen tsin aquacu lture rea lize the cruc ial ro le tha t proper nu tr i
tion ,
managemen t, and econom ics offeed ing p laysin sus tainab le aquacu ltu re
deve lopmen t. Theresea rch-basedin forma tion con tained he re a re mos tly
produc ts of severa l years o fresearch exper ienceinf ish nu trition and
feed deve lopmen ta t SEAFDEC AQD .
Wh ilethetex tbookisin tendedfo r use p rima r
ily byf ishe ry s tuden ts,
other aquacu l
ture enthus ias ts,l ike researchers ,f ish farmers , feed
manu fac ture rs, fa rm manage rs , andtechn icians may a lsof indi t use ful
as are fe rence book . We we lcome sugges tionsf rom s tuden ts ,ins t
ruc to rs,
andlea rners on howtoimp roveth istex tbook .
Itis our hope tha t, through th is book , SEAFDEC AQD wou ld be
ins trumen talin advanc ing thef ron tiers o f know ledgein aquacu lture
through educa tion .

ROLANDO R
. PLATON
Ch
ief
Tab
leo
fcon
ten
ts
Foreword i
ii
Abouttheautho
rsanded
ito
rs xv
Pre
facex ix

Chap
ter1
:INTRODUCT
IONTONUTR
ITIONINTROP
ICALAQUACULTURE 1

Feedsinaquaculture1
Feedsandtheen vi
ronm en
t2
Susta
inableapproache
stoaqua
cul
tur
e3
Summa ry 5
Suggest
edreadings5

Chap
ter2
:THEESSENT
IALNUTR
IENTS 7

Proteinsandam inoa cids7

Introdu ction7
Aminoa cids8
C
la ssif
icationo fam inoa c
id s9
Essentialam inoa cids 10
Non-e ssent
ialam inoa cids 10
C
la ssif
icationo fp roteins 11
Proteins tructure 12
Fateo fab sorbedam inoa cids 14
Impo r
tan ceo fam inoa c
idp rof
ilesinfishnu
tri
tion 14
Qualitat
iveam inoa cidrequ i
rem ents 14
Quan ti
tativeam inoa c
idr equirem en
ts 15
Def
icien c
ie sandex ce sse
so fdietaryam inoa
cids 17
Evalua t
iono fproteinqua lity 17
Proteinr equirem ent 18
Guid equ est
ion s 20

Lip
id sandfa ttya cid s 21
Introdu ct
ion21
Typ eso flip
id s 21
Gen eralfun c
tiono fl ip
id s 23
Fattya c
id s 23
Stru ctureandc lassi
ficat
ion 23
Nom enclatu r
e 24
Fattya c
idcompo sitiono ff
ish 26
Biosynth es
iso ffattya c
id s 28
Oxida t
iono ffattya cid s 29
Lipidp e rox
ida tion 30
Impo rtan ceo ffattya cidp ro
fi
lesinf
ishnut
rit
ion30
Ess ent
ia lFa ttyA c
idR equirementso
fFish 31
Guid equ es
tion s 32
vi Con
ten
ts

Carbohyd rates 33
Introduc tion33
Classif
icationo fcarbohydrates 33
Monosaccha rides 34
Disaccha r
idesando ligosaccha
rides 35
Polysaccha r
ides 35
Uti
lizationo fcarbohydrates 37
Guideques t
ions 39

Energy 41
Introduct
ion41
Ut
ili
zat
iono fenergy 41
Energy metabol
ism 42
Energybalanceandd ie
taryrequ
iremen
t 43
D
ietaryene rgyrequ
irement 43
Guidequestions 44

V
itam ins 45
Introduc t
ion45
C
lassificat
iono fvi
tamins 45
Wate r
-solublevi
tamins 45
L
ipid-solublev
itamins 51
V
itam inrequ irementsoff
ish 53
Guideques tions 56

Mine rals 57
Introduc t
ion57
Class i
ficationo fm inerals 57
Gene ralfunc tionso fm inerals 57
Mine ralava ilabili
ty 58
Mac rom ine rals 58
Microm inerals 60
Mine ralsupp lemen tationo fprac
tica
lfishd
iets 61
Mine ralrequ i remen tsoffish 61
Guideques tions 63
Summa ry 63
Sugges tedread ings 66

Chap
ter3
:FEED
ING HAB
ITSAND D
IGEST
IVEPHYS
IOLOGY OFF
ISHES 77

Introduc t
ion77
Feedinghab itsandbehav io
r 77
Anatomyandphys iologyo fth edigestivesys tem 79
Fishes 79
Crus taceans 84
Digestionandabso rption 86
Digestionandabso rpt
iono fprote
ins 86
Digestionandabso rpt
iono fcarbohyd rates 88
Digestionandabso rpt
iono fl
ipids 89
Measu remen tsandana lysis usedind iges
tionstudies 89
Measu remen tso fstomachcon tents 89
Measu remen tofd igestib
il
i ty 90
Facto rsa f
fectingdiges t
ionandabso rp t
ion 92
Feedingp rocessinf ish 92
 Con
ten
ts v
i
i
Appe t
iteandsa tiat
ion 92
Arou sa
lands earch 94
Locationandid ent
if
ica
tion 94
Captu re 94
Tastete st
ing 94
Swallowingo rreject
ion 94
Summa ry 95
Guidequ est
ion s 95
Suggestedr eadings 96

Chap
ter4
:FORMULAT
ION OFAQUAFEEDS 99

Introdu ct
ion99
Feeding redientsforaqua cul
ture 99
Feedformu la t
ion 104
Pearson’ssqua r
eanda lgeb
raicequa
tion m
ethod
s 108
Trialande rrorm ethod 113
L
inea rp rogramforleastcostformu
lat
ion 119
Puri
fiedd ietformulation 119
Summa ry 120
Gu
id equ es
tions 121
Sugge s
t edr ead
ings 121

Chap
ter5
:PROCESS
ING OFFEEDSTUFFSANDAQUAFEEDS 125

Introdu ct
ion125
Feeds tuffpro cessing 126
Di
fferen tm ethod so ffeed s tuffprocessing 127
Soak ing 127
Hea t
ingandcook ing 127
Dehu ll
ing 128
Extra c
tion w itho rgan icso lventandch emica
ltr
eatm
ent 128
Feedp reparationte chn iqu es 129
Grinding 129
Sizeg radingo rs ie v
ing 130
Weighing 130
Mixing 130
Cond it
ioning 132
Pellet
ingandex tru s
ion 132
Pelletcoolingandd ry
ing 135
Pelletcrumb ier 136
Pelletandc rumb ledfe edclean e
r 136
Produ ctpa ckag ingands torage 137
Stepsinla rge-s calefe edp reparat
ion 137
Step sinsma ll-sca lefe edp repa ra
tion 140
Step sinla rvalfe edp repa ration 142
Qualitycon t
ro l 144
Feed m i
llsan itationand ma intenan ce 145
Summa ry 145
Guid equ es
tion s 145
Sugg e st
edr ead ing s 146
vi
i
i Con
ten
ts

Chap
ter6
:EVALUAT
ION OFFEEDSTUFFSANDAQUAFEEDS 149

Int roduc t
ion 149
Phys ica leva lua tion 150
Useo fth esenses 150
Feed m ic roscopy 150
Measu rem en tso ffeedstuffsbulkdensi
ty 150
A t
trac tab ili
ty 151
Wa ters tab il
ity 151
Chem ica leva lua tion 152
Proxima teana lysis 152
Me thodso fp roteinevalua tion 155
Me thodso flipideva luation 156
Me thodo fv itam ineva luation 158
M ethodso fm ine ralevalua t
ion 159
Me thodso fene rgyde term inat
ion 160
Ana lysiso ftox insinfeeds 161
M
ic rob iologica leva lua t
ion 162
B
iolog ica leva lua t
ion 162
Pa rame terstobe mon itoredinafeedingexpe
rimen
t 175
Summa ry 165
Guideques tions 165
Sugges tedread ings 166

Chap
ter7
: MANAGEMENT OFFEED
INGAQUACULTURESPEC
IES 169

Introduc t
ion 169
Feedings trateg iesinpondcu lture 169
Produc tiono fna tu ra
laqua ticfood 170
Feed ingasupp lemen tarydiet 170
Feed ingacomp leted iet 171
Feeding managemen t 171
Samp l
ingandreco rdkeep ing 171
Feed ingra tion 172
Feedpa rticles ize 174
Feedapp lica tion me thods 174
Feeding , wate rqua li
ty,andtheenv ironment 175
Feeding ,oxygenrequ irements,and wate
rqual
ity 175
Fishfa rm was tes 177
Per
fo rmance measu res 177
Biomass 177
Feedconve rsionra tio 177
Feedingschemes 178
Milkf
ish 178
Tilap ias 181
Rabb it
fish 184
Bigheadca rp 185
Na t
iveca tfish 186
As ianseabass 188
Orange -spo ttedg roupe r 190
Mang roveredsnappe r 192
Tige rsh rimp 193
Mudc rabs 196
 Con
ten
ts iv
OtherspeciesforSto
ckenhancemen
t 199
Donkey’searabalone 199
Seahorses 200
Windowpaneoys ter 201
Summa ry 202
Guideques t
ions 203
Suggestedreadings 203

Chap
ter8
:ECONOM
ICS OFFEED
ING 209

Introduc tion 209

Costo fp roduc ingfeeds 209
Singleinpu tandsingleoutputproduct
ionfunction 211
Thep roduc t
ionfunc t
ionandthecostofp roduc
tion 213
Econom ice f
ficiencyoffeeds 214
Leas t
-cos tcomb inat
ionoffeeds 217
Minimumcos toffeedformulat
ionusinglinearprog
ramm ing 219
Summa ry 220
Guideques tions 220
Sugges tedread ings 220

L
ISTOFTABLESANDF
IGURES

TABLES

Chap
ter2

P
rote
ins

Tab
le2
.1 Namesandabb reviat
ionsofthecommonam inoacids 8
Tab
le2
.2 Aminoacidrequi
remen tsofsomefishesandsh r
impinpe rcentof
pro
tein 16
Tab
le2
.3 Opt
imaldieta
ryp rote
inleve
ls(%ofd rydie
t)fo
rsom eaquaculture
spec
ies 19

L
ipidsandFa
ttyAc
ids

Tab
le2
.4 Thecommonfa ttyacids 24
Tab
le2
.5 Unsaturatedfattyac idsfami
lies 25
Tab
le2
.6 Majo
rfattyac idsinlipidsof marineandfreshwa
terf
ishes 26
Tab
le2
.7 Ef
fectofdietonfa ttyacidcomposit
ionofshrimpPenaeusset
ife
rus 27
Tab
le2
.8 Essent
ialfattyac idcomposit
iono fvar
iouslip
idsources 30
Tab
le2
.9 Essent
ialfattyac idrequirementsoff
ishandsh r
imp 31

Ca
rbohyd
rates

Tab
le2
.10 Classi
fica
tiono fcarbohydra
tes 33
Tab
le2
.11 Meansfo rwe igh
tgain,feedef
fic
iencyra
tio(FER
),p
roteinef
fic
iency
ratio(PER )
,andsu rviva
lrateoftige
rshrimpfedd
iets w
ithdi
fferen
t
carbohyd ratesourcesandlevels 38

V
itam
ins

Tab
le2
.12 V
itaminde
fic
iencysymptomsinf
ishes 54
Tab
le2
.13 Summaryofthevi
taminrequi
rementsofva
riousspec
ieso
ffishand
shr
imp55
x Con
ten
ts

M
ine
rals

Tab
le2
.14 Summa ryofm inera
lfunct
ions 58
Tab
le2
.15 Summa ryofth eminera
lrequi
rementsofva
riousf
ishandsh
rimp
spec
ies 62
Tab
le2
.16 M
ineraldef
iciencysymptomsinfishandshr
imp 62

Chap
ter3

Tab
le3
.1 Feedinghabitsandnatura
lfoodofsom e
juven
ilefishandshr
imp 78
Tab
le3
.2 Generalobserva
tionsonfeed
inghabitsandrelat
ivegutleng
ths
(rat
ioo fin
test
inetobodylength
)infish82

Chap
ter4

Tab
le4
.1 Somesou rceso fpro te
in ,lipid,andca rbohyd rateinaquacu l
tu re
feeds 100
Tab
le4
.2 Am inoac idcompos i
tiono fsom efish m eals
,lea fm eals,ando ther
proteinsou rces 101
Tab
le4
.3 Proxima tecompos itiono fsom efeeding red
ien tsana ly zedbyth e
Cen trali
zedAna lyt
ica lLabo ratorya tSou theas tAsianF isher
ies
Developmen tC enter,Aquacu lture Depa rtmen t 102
Tab
le4
.4 Appa ren tp rote ind iges t
ibili
tycoe fficients(APDC )in %o fsom e
feeds tuffsfo raquacu ltu respec ies 103
Tab
le4
.5 Vitam inand m ine ralm ixtu resforc rustaceansandt igersh r
imp
juven ile 104
Tab
le4
.6 Recommendedv itam in m ixtu reforwa rmwa terf ishessucha s
mi
lkfish ,seabass , andca t
fish 105
Tab
le4
.7 Mineralm ixtu resfo rpu rif
iedandp ra ct
icalwa rmwa terfishdiets 105
Tab
le4
.8 Othe rfeedadd itives 105
Tab
le4
.9 Recommended max imuminc lusionleve ls(% )o fsome ma jorfeed
ing redien tsinap racticald ietforf ishandsh rimp 107

Chap
ter5

Tab
le5
.1 Var
iousan t
inut
rit
ionalsubstancesinsomefeeds
tuf
fsandtheir
removalorinact
ivat
ion 126
Tab
le5
.2 Guidetotypesandkindsoffeedforaquat
icanimalsatva
rioussizes
andages 143

Chap
ter6

Tab
le6
.1 Essen
tia
laminoac
idind
ices(EAA
I)o
fsom
ecommonfeeds
tuf
fsfo
r
shrimp 156

Chap
ter7

Tab
le7
.1 Some use
fulparame
tersthat mustberecordedinthefarm 172
Tab
le7
.2 Wate
rquali
typarame
ters, methodof measuremen
t, wa
terquali
ty
prob
lemsandpossib
lecau ses,scheme/methodof management,
andtarge
t wa
tercondi
tionsinb rack
ishwaterpondculture 176
 Con
ten
ts x
i
Tab
le7
.3 Pract
icaldietformulas(g/kgdryd iet
)form ilkf
isha tvariousstages
ofculture 179
Tab
le7
.4 Pract
icaldietformulas(g/kgdryd iet
)fortilapiaatva r
iouss tageso
f
culture 183
Tab
le 7
.5 Pract
ica
ldietformu
la s(g/k
gdrydi
et)forrabb
itfishbroodstockandfry
184
Tab
le7
.6 Pract
icaldietformulas(g/kgdryd iet
)forbigheadca rpa tvar
ious
stagesofcu l
ture 184
Tab
le7
.7 Pract
icaldietformulas(g/kgdryd iet
)forAs ianca t
fisha tvar
ious
stagesofcu l
ture 187
Tab
le7
.8 Pract
icaldietformula(g/kgdrydie t
)forjuven i
leseabass 189
Tab
le7
.9 Pract
icaldietformulas(g/kgdrydiet)forgrow -outcultureof
grouper 191
Tab
le7
.10 Pract
icaldietformula(g/kgdrydie t
)forredsnappe r 193
Tab
le7
.11 Pract
icaldietformulas(g/kgdryd iet
)fortigersh r
impa tva r
ious
stagesofcu l
ture 194
Tab
le7
.12 Pract
icaldietformulas(g/kgdrydiet)for mudc rabb roodstockand
grow-out 197
Tab
le7
.13 Pract
icaldietformula(g/kgdrydie t
)foraba lonebroodstock 199

Chap
ter8

Tab
le8
.1 Productioncos to fshrimpd ie
t 210
Tab
le8
.2 Relat
ionsh ipamongto talphysica
lp roduct
,ave ragephysical
product,and ma rginalp roduct 212
Tab
le8
.3 Relat
ionsh ipamongto talphysica
lp roduct
,ave ragephysical
product, ma rginalphys ica
lp roduct
,valueo ftota
lphysicalp rodu c
t,
valueo fave ragephys icalproduct,valueof ma rg
inalphysical
product,to talvariableinpu tcostandp ro
fit 215
Tab
le8
.4 Costandre turnso fthepencu ltu
reo fClar
ias ma cro
cephalus a t
stockingdens ityo f10f ish/m2andfedth reed i
fferentd
ietsfo r 120
days 217
Tab
le8
.5 Hypotheticalre lat
ionsh ipfo rcombiningfeedstop roduceag iven
levelofou tpu t(P1 =P9,P2 =P6)218
Tab
le8
.6 Dataonnu tr
ien tava i
lab i
li
tyandrequ iremen ts
,feedco s
t,objective,
andcons t
ra intsinl inea rprogramm ing 219

F
IGURES

Chap
ter1

F
igu
re 1
.1 Fishnu trition,feeds,andfeed ing managemen tplayimpo rtantroles
ininc reas ingthep roduc t
ivityofaquaculturefarms 1
F
igu
re 1
.2 Nutr
itiona lly-balancedfeedandadequa tefeeding arefactorstha t
maximizef ishp roductionandp ro
fi
tabil
ity 1
F
igu
re1
.3 Productiono fhighqua l
ityfeedsstar
ts wi
thgoodqua lityingred
ients
andp rope rqua litycontrolthroughoutprocessingunti
lu seinfish
farms2
F
igu
re 1
.4 Approp r
ia tefeed ing managemen tandgoodqua l
ityfeed maximize
feede f
ficiency ,lowe rcosto fproduct
ion,andreduceenv ironmen tal
degrada t
ion 4
xi
i Con
ten
ts

Chap
ter2

P
rot
eins

F
igu
re2
.1 Chem icals t
ruc tureso fthetenam inoacids 9
F
igu
re2
.2A Prima rystruc ture 12
F
igu
re2
.2B Seconda rys tructu re 12
F
igu
re2
.2C Thea lpha -helixs truc ture 12
F
igu
re2
.2D Thebe ta
-shee ts truc ture 12
F
igu
re2
.2E Tert
iarys truc ture 13
F
igu
re2
.2F Quaterna rys tructu re 13
F
igu
re2
.3A Growthresponseo ftigersh r
impf edg radedleve
lsofphenyla
lanine
for8weeksasdesc r
ibedbythequad rat
icregress
ion model 15
F
igu
re2
.3B Growthresponseo ftigersh r
impf edgradedlevelsof me
thion
inefor
8weeksa sdesc ribedbytheb rokenlineregress
ion model 15

Lip
idsandFa
ttyAc
ids

F
igu
re2
.4 Denovo synthes
iso
fsaturated monoeno
icfa
ttyac
ids28
F
igu
re2
.5 Biosyn
thes
isofunsatu
ratedfat
tyacids 29

Ca
rbohyd
rates

F
igu
re2
.6 Summa ryofhydro
lysisofcarbohyd
rates 36
F
igu
re2
.7 Growthandfeedef
fic
iencyofcarpfeddie
tsconta
ining42%o
fα-
starch
,dext
rinorglucose 38

Ene
rgy

F
igu
re2
.8 U t
il
iza
tionofenergy 41
F
igu
re2
.9 Chem icalst
ruc
tu reo
fadenosinetriphospha
te (ATP
) 42
F
igu
re2
.10 Thero
l eofATPincel
lularenerge
tics 42

Vi
tam
ins

F
igu
re2
.11 Thiamin(vitamin B1)def
ici
ency 46
F
igu
re2
.12 Ascorbicacid(vi
tamin C)defic
iency 50
F
igu
re2
.13 Retino
l(vi
tam inA) defic
iency 51
F
igu
re2
.14 Tocopherol(vi
tam in E
)defic
iency 53

Mine
ral
s

F
igu
re2
.15 Ca
lciumandphospho rusdef
ici
enc
ies 58
F
igu
re2
.16 Z
incdef
iciencyca
taract 61

Chap
ter3

F
igu
re3
.1 Diagramma ti
crep resen tationofthed igestivesystemsoffourf
ishes
arrangedino rde rofinc reasinggu tlength 79
F
igu
re3
.2A Variat
ionsofthe mou ths tructureinf ishes 80
F
igu
re3
.2B Some ma jorkindso fjawtee th 80
F
igu
re3
.3 Reg ionsofthed igest
ivet rac tofmilkfi
sh Chanoschanos 81
F
igu
re3
.4 Schema t
icrep resentationo fapo rtiono fanabso rpti
vecel
lfromthe
intest
ine 83
F
igu
re3
.5 D iagramo fthed igest
ivesys temofsh rimp w i
thg i
llsand
musculatureremovedtoshow ma jo rorgansys tems 84
 Con
ten
ts x
ii
i
F
igu
re3
.6 Schemeo fdif
feren t
iationandfunctiono fthedigest
iveglandtubu
le
85
F
igu
re3
.7 Sequenceo fproteind iges
tion 88
F
igu
re3
.8 Represen
tationo fdigestionandabso rpt
ionofca rbohydra
tes 88
F
igu
re3
.9 Diagramma t
icrep resentationofd
igestionandabso rpt
ionoflip
ids
89
F
igu
re3
.10 Experimentalset-upfo r invivod
igestibi
li
ty measurement91
F
igu
re3
.11 Feedingprocessinf ish 93

Chap
ter4

F
igu
re4
.1 Somefeeds
tuffsfo
raquafeeds
,fishof
fal
,shr
imp mea
l,an
imal mea
t
was
te,animal mea
tandbone m eal
,soybean m
eal
,andyeas
t 100
F
igu
re4
.2 Somesourcesofdieta
ryenergy 101

Chap
ter6

F
igu
re6
.1 Flowd iag ramfo rthep roximateana lysiso ffeedstu f
fsandfeeds 152
F
igu
re6
.2 A moistu reba lanceusedtode term ine mo isturecon tentoffeedstuf
fs
andaqua feeds 153
F
igu
re6
.3 The K jeldah ld isti
llation-ti
tration(K jeltec™ )appa ratususedto
analyzec rudep rotein 153
F
igu
re6
.4 The Sox tec ™appa ratus usedfo rcrudefa tana lysis 154
F
igu
re6
.5 A mu fflefu rnaceusedtoana ly zetheashcon tento ffeedstuf
fsand
aquafeeds 154
F
igu
re6
.6 TheF ibe r
tec™usedfo rc rudef iberde termina tion 154
F
igu
re6
.7 The H ighPe rformanceL iquid Ch roma tog
raph(HPLC )foranalyzing
theam inoac idcompos it
iono fap ro tein 155
F
igu
re6
.8 The Gas Ch roma tograph(GC )
,anins trumen tusedfo ranalysisof
fattyac idcompos it
ion 156
F
igu
re6
.9 Asamp lech roma tog raphic ana lys
isshow ingre tentiontimeo f
var
iousfa ttyac idsinasamp le 157
F
igu
re6
.10 Asamp le HPLCch roma togramo fav itamin m ixture 158
F
igu
re6
.11 Alabo rato ryse t
-upfo rafeed ingexpe rimen t 163

Chap
ter7

F
igu
re7
.1 Rangeo faquacu lturep racticesinre la
tiontoinpu ts 170
F
igu
re7
.2 Na tura laqua t
icfoodinponds :lablabandlumu t 170
F
igu
re7
.3 Samp lingbycas tne tandl if
tne t 171
F
igu
re7
.4 Useo ffeedingt rayto mon itorfeeding 174
F
igu
re7
.5 Ademandfeede r 174
F
igu
re7
.6 In terlinkedfac torstha tarec rit
icalforth esuccessofanaqua feed
175
F
igu
re7
.7 Milkfish Chanoschanos 178
F
igu
re7
.8 Floa ting ma r
inecagesandconc retetanksfo rb roods
tock 178
F
igu
re7
.9 In tens ivelarvalrea r
ingtanks 179
F
igu
re7
.10 Feed ing managemen tschemefo rlarvalrearingofm i
lkf
ish 180
F
igu
re7
.11 Feed inginthesam ea reaofthecageo rpond 181
F
igu
re7
.12 Tilap ian i
loti
ca, mossamb ica,aurea, redtilapia181
F
igu
re7
.13 La rvaere leasedf rom mou th -broodingfema le 182
F
igu
re7
.14 Grow -outfloatingne tcagesinalake 183
F
igu
re7
.15 Rabb it
fish Siganusgu t
tatusjuven iles184
F
igu
re7
.16 Feed ingschemefo rrabb it
fishla rvae 185
F
igu
re7
.17 Bigheadca rpA r
istichthysnob il
isinducedtospawnus inghormonal
in jectionandindoo rlarvalrea r
ingfac i
lit
ies 185
x
iv Con
ten
ts

F
igure7
.18 Nativeca t
fish C lar
ias ma c
ro cepha lus rea redinha tche ryandinne t
cagesinpond 187
F
igure7
.19 AsianseabassLa tescalcarifer 188
F
igure7
.20 Feeding managemen tschemefo rthela rvalrea r
ingo fseabass 189
F
igure7
.21 Orange -spottedg rouper Epinephe lusco ioides 190
F
igure7
.22 Feedingand wa ter managemen tschemefo rin tensiverea ringof
groupe rlarvae 191
F
igu
re7.23 Mang roveredsnappe r Lu t
janusa rgentima cula tus 192
F
igu
re7.24 Feedingand wa ter managemen tdu ringla rvalrea ringo fthe
mang roveredsnappe r 192
F
igu
r e7
.25 Tigersh r
impPenaeus monodon ands tag e
so fova r
ian ma tura
tion193
F
igu
re7.26 Feedingschem efo rt
igersh rimpla rvaeZ -zoea , M -mys is,
PL-pos t
larvaea tdays 1to20 195
F
igu
re7.27 Afeed ingtrayfo rshr
imp 196
F
igu
re7.28 Mang rovec rabs S cyl
la sp 196
F
igu
re7.29 Feedingscheme usedinthecu ltu reo f mudc rabla rvae 197
F
igu
re7.30 Grow -outcu ltureinpondsandinpensins talledinre forested
mang roves 198
F
igu
re7.31 Donkey ’sea raba lone Hal
iotisas inina feed ingonreda lga199
F
igu
re7.32 Platesubs tratesfo repiphyticdia tom s
,livefoodfo rla rvaeins ta
lled
intanks 200
F
igu
re7.33 Seaho rsesinb reedingandrea ringtanks 201
F
igu
re7.34 W
indow -paneoys terandhand icra fts using kap is she l
ls201
F
igu
re7.35 Rehab il
itat
iono ftheIloi
locoas tl
ine 202

Chap
ter8

F
igu
re8
.1 Re
lat
ionsh
ipbe
tweenTPP
,APP,and MPP 213
F
igu
re8
.2 Re
lat
ionsh
ipbe
tweenTVP
,TVIC,andpro
fi
t 216

APPEND
IX AP
-1

Ap
-A Samp
lewo
rksh
eetfo
rca
lcu
lat
ingth
enu
trien
tcompo
sit
iono
ffe
ed AP
-1

Ap
-B M e
thodso fp roteinana lys
is(K jeldahlM e
thod ) AP
-2
Crudefatana lysis(Soxhtec Me thod) AP-2
Lip
idex tract
ion(B l
ighand Dye rM ethod) AP -3
Saponif
ica t
ionandt ransester
ificat
ion AP -4
Methodofpe rox idevaluede term inat
ion AP -5
Procedureo ffa ttyacidva luede terminat
ion AP -5
Thiobarbituricac id(TBA)va luede termination AP
-5

GLOSSARY G
-l

ILLUSTRAT
IONANDPHOTOCRED
ITS CR
-1

INDEXI
-1
 x
v

Abou
ttheAu
tho
rsandEd
ito
rs
Rena to F. Agbayan i is consu l
tan t
-p rojectleade r, Ilda G . Bo rlonganis Sc ientistIIa t SEAFDEC AQD .
Coas tal Resou rce Managemen ta t Southeas t Asian Sheob tainedhe rdeg reein BS Chem is
t ry(cumlaude )
Ministerso fEduca t
ion O rgan iza t
ion Regiona l Cen te
r fromthe Cen tralPh il
ipp ine Un iversity andhe r MSin
forG radua te Studyand Resea rchinAg ricultureand Chem is tryth roughPCAMRD -SEAFDEC Scho larsh ip
forme r Associate Sc ien tista tthe Sou theas t Asian Grantf romthe UP D iliman . Sheis p resen t
ly a PhD
Fisher ies Deve lopmen t Cen ter Aquacu lture candida teonascho larshipf romtheJapanSoc ietyfo r
Depa rtmen t(SEAFDEC AQD )
. He ho lds deg reesin theP romo t
iono fScience .
Maste rsin Bus inessAdm in istrationand Bache loro f Herwo rkhasbeenfocusedones tabl
ish ingnu trient
Sciencef romthe Un ive rsity o fthe Ph i
lipp ines(UP ), requiremen tso fsome ma rinefishesanddeve lopingfish
Di
liman . feeds.Sheisa lsoinvolvedinJapanese -fundedp rojects
Hewasp rojectleade ro fthesuccess fullycomp le
ted onfishhea lth managemen tandb iotechno logyandis
Commun ity F
ishe ry Resou rces Managemen tp roject act
iveinthet rainingp rog ramso fSEAFDEC . Shehas
ofSEAFDECinco l
labo rationw i ththeloca lgove rnmen t autho redo rco -autho redove r25sc ien ti
ficpape rsin
units(LGUs ) and non -gove rnmen to rgan iza t
ions interna tional jou rnalsandp roceedings . Currently,she
(NGOs )a t Malal
ison , Antique ,Ph il
ippines(1998 ).H is isa membe ro ftheboa rdo fdirectorso fthePh ilipp ine
pub licationsinin terna tiona ljourna ls cove rthe Societyfor B iochem istry and Mo lecu lar Bio logy,
econom ics ofcomme rcia l
ly-impo rtant aquacu lture Integra ted Chem istofthePh i
lippines -Panay Chap ter,
speciesandcommod i
ties . andisa membe ro ftheed itorialboa rdo fthePh il
ipp ine
Journa lo fSc ience .
Ve ron ica R .A lava is Sc ientistI a t SEAFDEC AQD
whe reshespec ializesinaquacu lturenu tri
tionandfeed Mae R . Ca tacu tan ho ldsapos i
tionofSc ient
is tI at
deve lopmen tresea rch. She ob tained he r BS Ma rine SEAFDEC AQD . She gradua ted from S il
liman
Biologydeg reef romXav ier University, Cagayande O ro Univers i
tyw i thaBS Chem istrydeg ree (cumlaude )as
City,Ph il
ipp ines , MSF ishe r
ies(ma jorinaquacu lture) ana tiona ls tatescho lar. Sheob tained her MSdeg ree
atthe UP V isayas , anda PhDin F ishe r
ies(ma jo rin from Kagosh ima Un ive rsi
ty whe resheiscu rrentlya
nu tri
tionalchem istry )atKagosh imaUn iversity,Japan . PhDcand ida teonascho larshipf romJapan Soc iety
At SEAFDEC AQD ,she wo rksonva riousspec ies fortheP romo tiono fSc ience.
suchas m ilkfish,N ileti
lapia,seabass ,anemonef ish, She has conduc ted s tudies onthe nu tr
ition of
groupe r
,t ige rsh rimp , wh i
te sh r
imp , and mudc rab impo r tan taquacu l
tu respec ies;sh rimp
,c rab,seabass ,
eithe rintheha tche ry,nu rsery,grow -ou to rb roodstock andsnappe r,onp rotein ,vitam in Crequ i
remen t,and
stages . She has pub l
ished a numbe ro f pape rsin nutrien td iges tibil
ity . He r research resu l
ts a re
in terna tiona ljourna ls and p resen ted pape rs in publishedinre fereedsc ien t
if
icin terna
tionaljournals.
in ternationa lcon ferences,inc ludingtheAs ianF isheries She has a lso presen ted her work inloca l and
Soc iety(AFS ) and Wo r
ld Aquacu ltu re Soc iety(WAS )
, interna tiona l con ferences . Sheis a membe ro fthe
of wh ich sheis a membe r
. Sheisa lso alec tureron Integ rated Chem istso fthe Ph il
ippines,the AFS and
top ics pe r
ta iningto aquacu lture nu tr
ition andfeed anassoc iate membe ro ftheNa t
iona lResearchCounc i
l
deve lopmen tinSEAFDECAQDt rain ingcou rses. ofthePh ilipp ines(NRCP ).
xv
i
Relica rdo M . Coloso isSc ien tistIIa tSEAFDECAQD . Osen iM .M illamena issen io rSc ien tis ta tSEAFDEC
He ob ta ined h is PhDin Nu tritiona l Sc iencesf rom AQD . Shecomp le ted he r BS Chem Eng inee ring ( cum
Corne ll Un ivers i ty as a Fu lbrigh t-Hays Mu tua l laude )a t Cen tralPh ilipp ine Un ive rsity;ob ta ineda M .
Educa tiona l Exchange G ran tee. Hef inished h is MS Env ironmen tal Eng inee ringf rom As ianIns titu te o f
degreein B iochem is t
rya t UP Co llegeo f Med icineona Techno logy , Tha iland , as Sou theas t As ian Trea ty
PCAMRD-SEAFDEC scho larsh ip and h is BS Organ iza tion scho la r: a Pos t-g radua te D iplomain
Chem is t ry(cum laude )a t UP D il
iman , as a NSDB Chem Eng inee r
ingf romTokyoIns titu teo fTechno logy
scho la r
. He wasapos tdoc to ralfe l
lowa tthe Un iversity as a Un ited Na tions Educa tiona l, Sc ien t
ific and
of Med icineand Den tistryo fNewJe rsey . NewJe rsey Cultu ralO rgan iza tionfe l
low ; andaPhDin F ishe ries
Medica lSchoo l
. He wasa lsoafe l
lowinf ishnu tri
tion Science(ma jorinAqua tic Nu tritiona l Chem istry),f rom
attheIns t
itu te o f Ma rine B iochem istry , Abe rdeen , Kagosh ima Un ivers ity , Japan as rec ip ien to fa
Scotland . scho larsh ipf romJapan Soc ietyfo rthe P romo tiono f
His a reas o fspec ia l
iza tion a re onf ish nu tr
ition: Science . He r researchin teres ts a re ma inly on :
nutritiona lb iochem istry,am inoac idandphospho rus nut r
itiona lrequ iremen tsanddeve lop ingcos t-efficien t
metabo lism , nu t
rition o fm i
lk f
ish ,t igersh rimp , and feedsfo rt ige rsh rimp , mudc rabs ,g roupe r, aba lone ;
Asian sea bass , mo llusc icides and env ironmen tal andenv i ronmen taleng inee r
ing .
con tam ina tion ,andtox ico logy . He wasarec ipiento f Sing lyandinco l
labo ra t
ion w i
tho the rsc ien tis ts,
aresea rchg rantf romtheIn terna tiona l Founda t
ion she has unde r
takenseve ralresea rchs tud ies, 40o f
forSc ience( IFS )in 1991 . Heau thoredo rco-au tho red which a re pub lishedinin te rna tiona ljourna ls and
over30sc ien ti
ficpape rsinin te rna t
iona l journa lsand scien ti
ficp roceed ings . Sheisarec ip ien to fna tiona l
proceed ings . Heis a membe ro fS igma -Xi Hono r research awards for bes t pub lished papers in
Socie ty,AFS ,Ph ilipp ineSoc ietyfo rB iochem is tryand Aquacu l ture , Aquacu l
ture Eng inee ring , and the
Molecu la rB iology, NRCP ,In teg rated Chem istso fthe “Fishfo rthePeop le”2001awa rd ,g ivenbyPCAMRD ,
Philipp ines- Panay Chap te r,andPh ilippineFu lbrigh t DA -BAR ,and Ma rineTechno log ica lFounda tionInc .,
Scho la rs ’Assoc iation . respec tive ly. He rinvo lvemen tino the rac tivit
iesa re
nume rous : aslec tu re r
, thes is adv ise r and c ritic,
Nelson V . Golez is Resea rche rIIa t SEAFDEC AQD . resea rcheva lua to r,ed ito randcon tribu to r. Shed id
HeisalsotheFeed M illSupe rvisoro fthesamesec tion. collabo ra tiveresea rch p rojec ts w iththe Aus t ra lian
He holdsa BS deg reein Chem ical Eng ineeringf rom Cen terforIn terna tiona l Agr icu ltura l Research
Adamson Un iversity, Man ila, Ph i
lippines , andthen (AC IAR ), and is a member o f the AFS , the
wentto Kyo to Un ive rsity
, Kyo to,Japanfo rhis MSin Internat iona l Work ing Group in Crus tacean
Soi
l Chem istryasa MonbushoScho larin 1988 . Nu trit
ion ,the Soc ie ty o f Aquacu l ture Eng inee rsin
H
isa reaso fspec ializationa reva ried:fishnu trition thePh ilipp ines , Ph i
lipp ineAquacu ltu re Soc iety ,and
andfeeddeve lopmen t, waterqua l
ityandso ilchem istry, assoc iate membe ro fthe NRCP . She wasse lec tedfo r
pond cu l
tu re,feed m i
ll
ing ,feed p rocess ing, and inc lusioninthe 6 thEd itiono f Who ’s WhoinSc ience
storage. Heisa membe ro fthePh ilippineIns ti
tu teo f and Eng inee ring .
Chem icalEng inee rs,thePo llutionCon trolAssoc iation
ofthe Ph i
lipp ines -Reg ion V I chap ter, and the Felici
tas Piedad -Pascua l is one ofthetechn ica
l
Philippine Assoc ia tion o f Japanese Monbusho advisers in fish nutr ition o fthe Interna t
ional
ExchangeScho la
rs . Hewasarec ipiento fthe Dr.E lvi
ra Founda t
ion for Sc ience, and consu ltant at
O. Tan Awa rdfo r bes t pub lishedresea rch pape rin Aquacu l
ture Spec ia
list
, Gu imbal,Iloi
lo. She was
Aquacu l
tu rein 1998 . Heisau tho randco -autho ro f Scientist 1 when she re tired f
rom the Feed
several pape rs pub lishedinre fereedin ternationa l Developmen t Sect
ion , SEAFDEC AQDin 1990 . She
journals. wasaf ish nutrit
ionconsu ltantatthe Un ivers
ityof
 xv
ii
Diponego ro, Sema rang,Indones ia and o fthe Cuban Myrna N . Baut ista -Terue l is Sc ien tistI Ia t
governmen t
. Sheob tained he rPhDin Nu tr
itionf rom SEAFDEC AQD . She ob tained he r MSin Food
Iowa S tate Un iversity, and her MSin Food and Scienceth roughaPCAMRD -SEAFDECScho larship
Nutrit
ionf rom M ichiganS tate University
. Shef inished Gran tfromUP D i
liman .Shef inishedhe rBSF ishe r
ies
her BS Pha rmacy deg reein UP D i
liman . Sheis an degreea lsoin UP D i
liman .
associa te membe ro fthe NRCP and a membe ro fthe Shehaspub lishedseve rala r
ticlesonva r
iousaspec ts
Phil
ipp ine Assoc iat
iono f Nutrit
ion , AFS,ando thers. ofaquacu ltu re nu tri
tioninin terna t
iona landloca l
Sheisarec ipiento fnume rousawa rdsandhono rs, journa ls.
amongthemthe E lvira O . Tan Memo r
ial Awa rd,F ive She wasarec ipiento faresea rchg ran tf romthe
Thousand Pe rsona li
ties o fthe Wo rld, and resea rch Interna tiona lFounda t
ionfo rSc ience( IFS )from 1991
grantsf romtheIn terna tionalA tom ic Ene rgyAgency , to 1997 . She wasarec ipiento fthe D r
.E lvi
ra O .Tan
Amer ican Soybean Assoc iat ion , San M igue l Awa rdfo rbes tpub lishedpape rinaquacu lturein1991 .
Corpora tion, and Te tra, among o the rs. She a lso Herresea rchin te rests a re ma inly on aquacu lture
receivedt raining and s tudyg rants. Sheis au thoro r nutr
i t
ion ,feedp roc e s
sing,andfeedqua litycon trol
.She
co-au tho ro f mo rethan 40 pape rsinin terna tional is presen tlyoneo fthetechn ica ladv ise rsonAn imal
journa ls,p roceedings,books ,andex tension manua ls
. Produc tionandAqua t
icResou rcesfo rth eIFS.Sheisa
Sheisco -authoro ftherecen tlypub lished Handbook membe ro f
th eAsianF ish e
riesSo cie
ty ,theIn ternational
ofIngred ientsforAquacu lture Feeds . Working G roupon C rustacean Nu trit
ion , NRCP ,and
Phil
ipp ineAquacu l tureSoc ie
ty.
 xix

P
reface
Increas ingf ish produc tionthrough aquacu ltureto supp lythe demands
of an eve rinc reas ing popu la t
ionisimpo rtan t becausef ish y ieldf romthe
seais unpred ic tab le and manyf ish s tocks a refas t becom ing dep leted .
Aquacu l ture is present ly cons idered as the fas tes t expand ing food
produc ingsec torinthe wo rld andisexpec tedto assume ag rea te rro lein
Phil
ipp inef ish produc t
ioninthe new m i
llenn ium . The recen t advances
in aquacu l
ture techno logy are geared towardsfas terf ish grow th and
higher y ie lds. Properf ish nu trition and adequa tefeed ing are cruc ia l
componen ts ofth istechno logy . Feeds mus t be nu tritious and econom ica l
for ag ivenfarm ing sys temto be compe tit
ive and sus ta inab le andfeed ing
must be prope rly managedto be e ffect
ive .
The nu tr
itiveva lue o fan aquacu lture d iet depends no t on ly uponthe
proper ba lance o f nu trien ts accord ingtothe nu t rien t requ iremen ts o f
the cu l
tured spec ies and nu t r
ien td iges tib i
lity bu ta lso onthe use o f
propertechn iquesin the process ing o ffeeds tu ffs andfeedsto remove
antinu tritiona lfac to rs. The op tima lcond i t
ions dur ingfeed s to rage shou ld
be known sotha tfeed nu trien ts do no t de terio ra te. Qua li ty con trol and
carefu linspec tion o fthefeed dur ing p rocess ing a re necessa ryto ensu re
the produc tion o fh igh qua lityfeeds . Goodfeedformu la tion and prope r
feeding managemen ta re necessary no t on lyfo r mak ing an aquacu lture
enterpr ise pro f i
tab le but a lso for reduc ing me tabo lic was tes and
preven ting aqua tic po llu tion . Env ironmen t-friend lyfeeds a re c riticalin
makingf ishfarm ing sus tainab le.
Th is bookisthe produc to f mo rethan 25yea rs o fresea rch expe r
ience
inf ish nu trit
ion andfeed deve lopmen ta tthe Sou theas t As ian F ishe ries
Deve lopment Center (SEAFDEC ) Aquacu lture Depar tment (AQD ).
SEAFDEC AQD sc ien t
is ts have endeavo redto deve lop cos t-effec tivefish
and c rus tacean d ie tsforthe aquacu ltureindus tryinthereg ion . Mos to f
thein forma tion presen ted a re based on pub lished papers on nu tr
ition
andfeed ing as we ll as unpub lished researchesf rom SEAFDEC AQD ,
those ga the redf rom o the rresearchins titu tions , andf romthel iterature
pub lished wor ldw ide . Feeds andfeed ing managemen to ftrop ica lfish
spec iesl ike m ilkfish ,N ilet i
lap ia,rabb itf
ish , As ian sea bass ;c rus taceans
liket ige r shrimp and mud c rab ; and o the rloca l spec ies a re h igh l
ighted .
This bookis a lso a produc to f mo rethanten years o fexpe rienceinthe
conduc to ftheIn te rnationa lF ish Nu t
rition Tra in ing Cou rse a t tended by
interna tiona l andloca l par ticipan ts. A SEAFDEC AQD aquacu lture
extens ion manua l en t
itled Feeds and Feed ing of M ilkfish ,N ile T ilapia ,
Asian Sea Bass , and T iger Shr imp preceded the pub lica tion o fth is
textbook . In add ition, our cons tan tin terac tion w ith the tra inees and
theirsugges tions andcommen ts we re mos t bene fic ialintheimp rovemen t
and upg rading o fthe ex tens ion manua lin toth istex tbook .
xx Pre
face

Th is bookisin tendedtoteach unde rg radua te s tuden tsthe essen tials

of aquacu l ture nu tri t
ion ,feedfo rmu la t
ion , andfeed ing managemen t.I t
serves as are ference bookfo rresea rche rsin aquacu ltu re, aquacu lturists,
fishfa rme rs, and aquacu ltu re nu trition ists. Fo rthe bas ic unde rstand ing
ofthe ma terials presen ted ,i tis essen tialtha tthe s tuden ts, teache rs,
and researchers have a good backgroundin chem is try,f ish b iology,o r
fisher ies. The book covers the sub ject areas o f known nutr ient
requ i remen ts,e ffec ts o f nu t rien t defic ienc ies on va rious aqua tic spec ies,
nu t
rien tsou rces,d iges t ibili
ty , and d iges tive phys iology . Feedfo rmu lation ,
process ing and s torage , eva luat ion and qua lity contro l, feed ing
managemen t, as we ll asthe econom ics o ffeed ing a reinc luded . Aspec ts
on feeds and feed ing re lated to the conservat ion o f the aquat ic
envi ronmen ta re a lso dea l
tw ith.
Each chapter o f th is book has com mon features such as an
introduc tion , bas ic concep ts, and a summary . Bo th the bas ic and
prac tical aspec ts o ff ish nu trition a reinc ludedto g ivethe s tuden ts and
allow the readers who are un fami liar w ith the top ics a c lear
unde rstand ing and know ledge o fthese concep ts .S tudy ques tions a tthe
end o feach chap te r se rve as a gu ideto summa rize andimp ress onthe
studen ts the sa lien t po in ts o fthe sub jec t ma tterin each chap ter. To
easily comprehendthe sub jec t ma t te r
,the reis an append ix con tain ing
the ana lytical me thods and a g lossary o ftechn ica lterms . The users
particu larlythe s tuden ts a re encouragedto broadenthe ir know ledge by
refer ringtothel is to fre ferences and sugges ted read ings a tthe end o f
each chap te r
.
We w ishto acknow ledgethe suppo rto fthefo llow ing: the SEAFDEC
Aquacu l
ture Depar tmen t andi ts Ch ief, D r. Ro lando R .P laton; Head o f
Train ing andIn forma tion D ivision, Eng r. Pastor L . Tor res ,J r.
; andthe
Pres iden to fIloilo S tate Co llege o fF ishe ries, D r.E lpidio Locs in Jr. We
also thank Mar ilyn Sur tida , Edgardo Ledesma , Er ic Gasa taya , Jose tte
Bangcaya ,E lme r M i raso l
, and Na than iel Ma r
tefo rtechn ical and ed iting
assis tance .

The Ed
ito
rs
 CHAPTER

In
trodu
inT
c
t
rop
i
ionto Nu
c
alAqua
tr
i
cu
l
t
ion
tu
re
OSEN
I M
. M
ILLAMENA
1
In
troduc
tion
Fishis av ital componen to ffood secu ri
ty espec ia
llyin
developingcoun tr
ies o fthe world. Asth e wo r
ld popu lation
grows , the need fo r mo re food and mo re fish has
correspond inglyinc reased. Aquacu l
tu re,thefa rming and
husband ry o ffish and o ther aqua t
ic o rganisms ,is now a
well
-es tablishedindus try wo rldwide andisthefas tes t
growingfood p roduc t
ion sector
. How ever, as aquacu ltu re
operations expand ,th erisktoth eenv ironmen tg rows.
Fish nu t r
ition andfeed ing playimpo r
tan tro lesinthe
sustainab le deve lopmen to f aquacu lture. The e ff
icien t
conve rsion o ffeedtof ishi simpo rtan ttof ishfa rme rs
becausef eedi sthela rgestcomponen to ftheto talcos to f
produc tion. Imp rovedf eed compo sition and be tterf eed
eff
iciency w i
llre su l
tin h igherf i
sh p rodu ct
ion ,lowe rf eed F
igure1.1
F
ishnu t
ri
tion
,feeds,andfeeding managementp
lay
cost, andlow wa ste produ c
tion hence , de c
rea sed nu trient importan
trolesininc
reasingthep roduc
t
ivi
tyof
loadf romf ishfa rm ing . aquacul
turefarms.

FeedsinAquacu
ltu
re

Aqua tic an ima ls,like any o the rliving o rgan isms, need
essen t
ia l nu tr
ients orsubs tancesfo rg rowth,t issuerepa i r
and ma intenance , regu lation o f bodyfunc t
ions , andto
maintain hea lth. Asfish cu lture becomes mo rein tensive ,
ita lso becomesless dependen t on na tura
lfood and mo re
on p repa redfeeds . A nu trit
iona lly -balancedfeed and
adequa tefeed ing areimpo rtantfac to rsthat help max im ize
fish p roduc tionand p ro
fitabil
ity
. Inapp ropr
iatefeedscou ld
resultind iseaseou tbreaks, poorg row th
, and high mo rta l
ity
offishesinthefa rm . Good qua l
ityfeed coup led w ith
approp r
iatefeed ing managemen t has been showntoresu l t
inimp rovedfeed conve rsion efficiency ,lowe r costs o f
produ ction,andreducedleve lso fenv ironmen tald egradation .
An e ffectivefeed ing p rogram shou ld cons ide rthe bas ic Figu
re1 .2
Anutr
itionally-ba
lancedfeedandadequatefeeding
principles o ffish nu trit
ion andfeed ing . Th isrequ ires an areimpo rtantfacto
rsthat max
imizef
ishproduc
t
ion
unde rs tand ing ofthe nu tr
ien trequ iremen ts ofcu ltu red andp rof
itabi
li
t y
.
2 CHAP
TER1 NUTR
ITIONIN TROP
ICAL AQUACULTURE
spec ies ,the irfeed ing hab itsandbeha vio r
, andth eab ilityo ff ishtod ig est
and u tilize essen tial nu trients .
Aqua feeds mus t sa tisfythe nu trien trequ iremen ts o fthe cu l
tu red
spec iesinte rm so fp roteinandessen t
ia l am ino a c
id s,l ipidandessen t
ia l
fa tty a cids, ene rgy ,v itam ins, and m ine rals
. The qua lity o fth efeed w il
l
ultima tely depend ontheleve lo f ava ilable nu tr
ien tsfo rf ish . Because
fishea ttosa tisfythe irene rgyrequ irem ent
,th eene rgyva lue o fth efeed
willa ffec tits e f
fic iency .
The nu trien trequ iremen tsshou ld b e knownfo raspec ificf ishspec ies
in tendedfo r cu ltu re. Fo r examp le, the p ro tein requ iremen ts o f an
omn ivo rel ike m ilk f
ish and a he rbivo rel iket ilap ia a re gene ra llylowe r
thanthoseo fca rn ivo resl ikeg roup er,seaba ss,andsnapp er.F ishspec ie s
also d ifferinthe irrequ iremen tsfo r essen tia l am ino ac ids . M i
lkfish
requ i
resn-3fa ttyac ids wh ile As ianseabassandt ige rsh rimp needbo th
n-3 andn-6fa tty ac ids . In con t
ras t,t ilap iarequ ires n-6fa tty ac ids .
Thus ,feedfo rmu la tionsshou ld be based onth erequ iremen tand leve ls
of essen tial nu trien tstha ta re op tima lfo r th ecu ltu red sp ecies.
In feed deve lopmen t
, the re shou ld be a con tinuous
inves tiga tionon me thodstoimp roveth equa lityo fraw ma terials,
reducefeedco st, andimp rovefeed ing managemen tinth efa rm .
The nu trien tsinfeeds haveto bee f
fic iently u tili
zed byth ef ish .
The re a re nume rous new p roduc tsinth e ma rket such a sfeed
attrac tan ts,b inde rs, andg row th p romo ters . How eve r, newfeed
add i tives a re cons tan tly be ingin troduced w ithou tp rope r
evalua tion o fthe i refficiency ,thu scau tion mus t beexe rcisedin
usingthemin aqua feeds . Feed manu fac tu ring shou ld ensu re
tha tth e nu tr
ien t compos i tionis ma intained and an ti-nu trien t
fac to rs a re elim ina ted . Feed qua l
i ty con t rol mus ts tartf rom
ing red ien t se lection and con tinueth roughfeed p rocess ing ,
storag e, and useinf ishfa rm s.
Figur
e1 .3
Product
ionofhighqual
ityfeedssta
rts w
ith A good qua lity and nu tritiona lly -adequa te feed can be
goodqua l
it
yingred
ien
tsandp roperqual
ity ine ffec tive un less p rope rfeed ing p rac t
ices a re us ed . Emphas is
cont
rolthroughou
tprocess
ingunti
lu seinfish
farms.
m ust als o be gi ve n to g ood fee ding m ana g em e nt a n dim pr ove d
feed pe rformance . An e ffec tivefeed ing managemen trequ ires
answe rsto ques tions o f wha t, how mu ch, when , how o ften, and whe re ,
tofeedthef ish . Thefeed ing reg ime used shou ld ma tchthefeed ing
behav io r and d iges tive cyc le o fthef ishin o rde rto max im izefeed
util
i zat
ion . Anyreduc tioninfood was tage w ill have as ignifican timpac t
onf ish p roduc t
ioncos ts andth e qua lity ofth e cul tu reenv i ronmen t
.

FeedsandtheEnv
ironmen
t

Therapiddeve lopmen tofaquaculturefromex tensivetoin tensivesystem s

has posed some p roblems aboutits sus tainab i
lity
. Aqua feeds p rovide
nutr
ientsforfish, bu tcanbe ma jo
rsou rceso fpo l
lutantsinaquacu lture
production systems .N it
rogen, phospho rus,o rganic subs tances, and
hydrogen sul
fide a reth e mainfacto
rs a ffectingenv ironmen tal po
llution
off
ishfa rms
.Asf ishs tockingdensi
tyisinc reased,ap roport
ionateamoun t
of metabol
ic was tesis p roduced
. Was testha t accumu lateinthe pond
cans low downf ishg rowth and ar
etoxictof ish.
 CHAP
TER1 In
troduc
tionto Nu
tri
tionin T
rop
ica
l Aqua
cul
ture 3
Fishfa rm was tes cons isto f: a ) so l
id ma t
ter, ma inly a m ix tu re of
unea tenfood ,feces , andco lonising bac ter
ia ; b )d issolved ma t ter, such
as ammon ia,u rea, ca rbon d ioxide, phospho rus and hyd rogen su lfides.
The was tesinc lude am ino ac ids,p ro teins,fa ts
, ca rbohyd ra
tes ,f ibe r
,
mine ralsandbac teria(Boyd 1989 )
. Sinceth eamoun to f was tein creases
with poo rfeed conve rsion
,thelesse ffic
ien tfish and sh rimp p roduce rs
are a tg rea testrisk. The amoun to ffeedshou ld be ad justedto m in im ize
unconsumedfeed and p reven t pollution o fthe pond ,the su r round ing
area s,andth ecoas ta
lzon e
. Poo rfeeding managemen tcanb ringd is eases,
slowg row th,lowsu rvival
, env ironmen tal degradation, poo r harv est, and
la rgef inanc iallosses .
Aqua feeds haveto beenv ironmen t-friendly. Feed deve lopmen tshou ld
takein to accoun t new know ledgerega rding nu trientrequ iremen ts and
diges tibi
lity,imp rovedtechn iquesto make mo re wa te rs tablefeeds and
grea ter u t
ilization o fa lterna t
ive sources o fp rotein . The ma jor
cons ide rationsinfo rmu lation o f feedsfo r sus tainable aquacu l tu re a re
:

□ Feeds shou ld have a we ll

-ba lanced nu t rientp rof
ile;i .e.
, mo re precise
am ino ac id prof
ile,inc reased ene rgy and ba lanced p roteinto ene rgy
ratio, sotha t mo re n itrogen w il
l be ass imilated andle ss nitrogenous
wastes w il
l be exc re ted byf ish;
□ Total phospho rusleve lsinfeedsshou ld be based onth erequ iremen t
ofcu l
tu red spec ies andfeeding redien ts w ith h igh ly ava i
lablefo rms
ofphospho russhou ldbe u sed. Phospho rusd ischa rgeinponde ff
luen t
isin f
luenced bytheleve l and b ioava ilabil
i ty o f phospho rusinthe
feed . A be t
ter unde rstand ingo ffac to rsinvo lvedinth e pa r
tit
ioningo f
phospho rusinth eva riousf ractionso fth e ponde f
fluen tw il
lresu l
tin
bette r pond managemen t
;
□ Diges tiblefeeding red ien ts shou ld be usedtoreduce o rganic was tes
fromfeeds . G rea te r ass imilat
ion o ffeed nu trients byf ishimp roves
feed e fficiency andreduces aquacu ltu re was tes;
□ Pel
le t wa terstab i
lityshou ldbeimp rovedby us inge ff
ic ientd ietbinde rs
and p rope rtechn iquesfo r pelletingfeeds . Unea tenfeeds co llecta t
the bo t tom o fthe pond and con t ribu te to rap id wa te r qua li
ty
deg rada tion;
□ Grea te r use ofa lte rna tive p ro
tein sou rcesfo rf ish mea lin aqua feeds
shou ld be encou ragedth roughcon tinuous andimp rovedre search;
□ Exo t
icfeeding redien ts wh ich maycon tain unknowng row thinh ibitors
shou ld be avo ided un less me thods a re ava i
lab leto m inim ize or
el
im inatethesefac to rsinth efinishedfe ed.

Sus
tainab
le App
roachesto Aquacu
ltu
re

In mak
ingaquaculturesus
tainab
leforfu
turegene
rat
ions
, th
ekeyissu
es
re
lat
ingtofeed deve
lopment andfeed
ing a
re:

1
. the needtoreducefeed co st
.
Feedisth e mostcost
lysinglei
teminf ishfarmingandth eavai
lab i
li
ty
of a cos
t-ef
fectivefeed remains asthe bott
leneckto aquacu l
ture
deve
lopmen t
. Feedformulasshouldbere f
inedandfeeds madech eaper
.
Readi
ly avai
lableratherthan expensiveimportedingred
ients should
4 CHAP
TER1 NUTR
ITIONINTROP
ICAL AQUACULTURE

be used a sfeeding red ien ts. These a l te rna t

iveraw ma te r
ia ls may
dif
fe rf rom one coun tryto ano the r andthe i r ava ilab ility may be
loca l
i zed. Thus ,feedsu rveysshou ldbeconduc tedtoknowth eloca tion
oftheseresou rces andthe ir ava ilab ility , nu trien t compos ition , and
cost. Such an app roachis necessa ryto max im izethe iru tiliza t
ion
andreduceth ere liance onimpo rtedfeeding red ien ts.
2. the sea rchfo ra lte rna tives tof ish mea l andf ish b iomassfo r
aquacu ltu re .
Trad i
tiona lly,f ish m ea l has beenth e ma jo rcomponen to ff ishfeed s.
Inc reasedg loba laquacu ltu re p roduc t ionandareduc tioninf ish m ea
l
produc tioninc reasesthe needfo ra lte rna t
ive p ro te in sou rces .F ish
meal hasbecome mo reexpens iveandsca rcedu etoinc reaseddemand
ands t i
ffcompe tition w ith o the rfood p roduc tionsec to rs . Inc reas ing
demandfo rf ishe ry p roduc ts dueto arap idly g row ing popu lation ,
espec iallyf rom As iancoun t ries ,w illfu rthe r dec reaseth e ava ilab i
lity
off ish m eal. F ishshou ldbe usedtofeedpeop lera the rthan
be conve rtedin tof ish mea lo r be usedtofeed ca rn ivo rous
fish .
Agricu ltu ral by -p roduc ts , such a svege tab le and an ima l
mea ls, havebeen u tili zedtop roducecheapandcos t-effec tive
die tsfo r seve ral aquacu lture spec ies . Some o fthese
prom isinging red ien tsinc lude h igh p ro tein ,low ash mea t
mea ls, pou l try o ffalm eals, andlegumessuch a s beans and
peas , mod ified whea tg lu ten p roduc ts,o ilseeds , and o the r
gra ins . B io techno logy may a lso be usedto p roduce nove l
ing redien ts w iththe des ired nu tritiona l cha rac teris tics.
3. the greater use o f supp lementary feeds and
supp lemen ta lfeed ing sys tem s.
Comme rcia l aqua feeds a re usua lly expens ive becausethey
con tain nu t rien tsin excess o f wha tthef ish needs . They
are a lsofo rmu la ted as comp le te d ie tsi rrespec tive o fthe
in tendedf ishs tock ing dens ity and p resence o f na tu ra lfood
inthe pond . Supp lemen ta ryfeed ing p rac ticeis pa rt
ly
dependen ton na tu ra lp roduc tivitytosupp lysom e nu trien ts.
Feedison lyusedtosupp lemen tth eendogenousfoodsupp ly.
Thu s,th eimpo rtance o f na tu ra l p roduc tiv ityinth e ove rall
F
igure1.4
n utritio n of po n d- rai s ed f
i s h es sho ul d b e giv e n em ph asis.
Agoodquali
tyfeedandapp ropr
iatefeed
ing Sincefeedis used on ly as a supp lemen tto na tu ralfood ,
management max
imizefeede f
f
icienc
y,lowercostof th isfeed ing sys temis mo re cos t-e ffic ien ti fthe pond
produ
ct
ion
,andredu ceen v
ironmentaldegrada
t
ion.
ecosys temis we llunde rstood . Max im izingth e useo fna tu ral
food o rgan isms w illreduceth ecos to ff ish p rodu ction .
4. integrat ion o ffeed deve lopment and feed ing management
prac tices w ith env ironmen ta l awa reness .
Unea tenfeed and me tabo lic was tes o ffish a re ma jo r sou rces o f
pol
lu tan tsf rom aquacu ltu re p roduc tionsys tem s. The reis a needto
developfeedstha ta reenv ironmen t-friend ly. Thenu t rien tcompos i
tion ,
nutrien t ba lance ,d iges tibility , and wa te rs tab ility o ff ishfeeds have
as ign ifican timpac t on wa te r qua li ty . Imp roved d ie ts w ith be tter
feed e f
ficienc ies and use o feco logica lly -soundfeed ing managemen t
wi
llleadto s ign ifican treduc tion o f was tesf rom aquacu l tu re.
 CHAP
TER1 In
troduc
tionto Nu
tri
tionin T
rop
ica
l Aquacu
ltu
re 5
Summa
ry

Fish nu trition,feeds , andfeeding managemen ta rec ri

ticalfacto
rsinth e
sustainab le deve lopmen tofaquacu lture. Thecho iceo fasu itab
lefeedis
animpo rtan ttask o fan aquacu l
tu r
ist. Afeed ings trategythatu t
ili
zes
cost-ef
ficien t,lowf ish m ea
l,andlowpo llut
ionaqua feedsinasupp lemen ta
l
feed ing sys tem w ill help ensure thelong - term sus tainability of
aquacu ltu re.I tisa lsoth eresponsibi
lityoftheaquacu lturisttoimp lement
soundfeed ing strategiestoinc reasef ish production,p rofi
tabi
lity, and
preventth e degrada tion ofthe culture envi
ronmen t
.

Sugges
ted Read
ings

Ak
iyama D, Dominy WG,andLaw renceAL 1991
.Penaeidsh r
impnu tr
it
ion
forthe comme rc
ialfeedindustry
. Rev
ised.In: Akiyama DM and
Tan RKH(eds )
.P roceed
ings ofthe Aquacul
ture Feed P
rocessing
and Nu tri
tion Wo rkshop. Amer ican Soybean Assoc iat
ion ,
S
ingapore. 155 p
.

Boyd C
. 1989
. Waterqual
ity managemen
tandae ra
tioninsh
rimpfa
rming
.
Ame r
ican Soybean Assoc
iat
ion
.S ingapo
re. 70 p
.

F
inf
ish nutr
it
ionresearchin Asia
:P roceed
ings ofthe Second As
ian F
ish
Nut
rit
ion Networks M
eeting.1988.In:D eS i
lva S(ed)
. Heinemann
Publ
ishers As
ia Pt
c.Ltd.Singapore. 128 p
.

Feed Developmen t Sect

ion. 1994. Feeds andfeed ing o
f milk
fish,Nile
tilapia
, Asian sea bass andtiger shrimp. Aquacu
lture Extens
ion
Manua l No. 21. SEAFDEC Aquacu l
tu re Depar
tment
,T igbauan,
Iloi
lo, Phi
lippines
. 97 p
.

F
ish Nut
ritionand Ma r
icul
ture
. 1988. The Gene
ral Aquacul
ture Cou
rse
.
In: Wa tanabe T(ed)
. Departmentof Aquat
ic B iosc
ience
. Tokyo
Univers
ity o
fFish. 233 p
.

Tacon A
. 1988
. The nut
rit
ion andfeeding o
ffarmedfish and shr
imp.A
train
ing manua
l.3. Feeding me
thods
. FAOFie
ld Docum ent
,P ro
ject
GCP/RLA/075/ITA.F ield Document 7/E
, FAO,B rasi
lia,B ra
zil
.
208p.
 CHAPTER

The Essen
tia
l Nu
trien
ts
OSEN
IM.M
ILLAMENA
2
This chapter d
iscusses the essentia
lfood nu t
rients, theirc lassi
fication
ortypes, chemicalstructures, generalfunctions, andimpo r
tanceinthe
nutri
tion ofaquat
ic animals. Itis dividedinto six sect
ions:p ro
teins and
am ino ac
ids,lipids and fatty acids, carbohydra tes, energy ,vitam ins,
and m ine
rals. There are speci
ficlearn ing ob
jectivesfo r each section.

PROTE
INSANDAM
INOAC
IDS

In
troduc
tion
Pro teins are macromo lecu les made up o f carbon , hydrogen , oxygen ,
nitrogen , and may a lso con ta in su lfu r
. The n itrogen con ten to f protein
distingu ishesi tfromfa tsandca rbohyd ratesando the ro rgan iccompounds .
Pro teins occurin everyl iving ce ll as compounds o ft issues and o rgans
and a re ma jor componen ts o ff isht issues. They a re neededfo rg row th
andt issuerepa ir and ma in tenance . No o ther nu trient cantakethe p lace
of pro teinini ts ma jorro le o f bu ilding and repa iring wo rn ou t cel
ls and
tissues . In add ition,p roteins a re alsorespons iblefo r musc le contrac t
ion
and a re componen ts o f enzymes , ho rmones , and an tibod ies. Pro teins
may be comp lexed w ith heme , ca rbohyd rate,l ipid,o r nuc leic acids.
Aqua tic an ima ls mus t consume p roteinto p rovide acon tinuous supp ly
necessary for rep lac ing worn-ou tt issues (ma in tenance ) and for the
syn thes is of newt issues(g row th and rep roduc t
ion ). Inadequa te dietary
pro tein w i
llresu l
tinre tarda tion o rcessa t
ion o fg row th o r aloss o f weight
dueto w ithdrawa lo fp roteinf romless v i
talt issuesin o rde rto ma intain
thefunc tions o f mo re v i
ta l ones .
This sec t
ion a ims toteach the readertheten essen tial am ino ac ids
requ ired byf ish and the ir chem ica ls tructures ,d istingu ish be tween
essen tial and non-essen tial am ino ac ids; thefa te o f absorbed am ino
acidsinf ish; e f
fects o f defic ienc ies and excesses o fd ie tary am ino ac ids
inf ish d iets; the procedure on how to de term ine the qua li
ta t
ive and
quan tita t
ive am ino ac id requ i remen ts off ish ; me thods o f evalua ting
pro tein qua lity; and how to de term ine pro tein requ i remen ts o f some
aquacu lture spec ies.
8 CHAPT
ER2 NUTR
ITIONINTROP
ICALAQUACULTURE

Am
ino Ac
ids

Proteins can be b roken down o r hyd rolyzedin to a numbe rof basic un its
called am ino acids. These am ino ac ids a re ca l
ledthe bu i
lding blocks o f
pro te
ins . The te rm am ino comesf rom the -NH2or an am ino group
whichis“bas ic”in na ture andthe“ac id ” partcomesf romthe -COOH o r
a carboxy lg roup, hence the te rm am ino ac id. In pro tein mo lecu les,
am ino ac idsform pep t
ide bonds (bonds be tween am ino and carboxy l
groups )inlong s trands ca l
led po lypep tide cha ins. There are many am ino
acids in na ture bu t on ly 20 are natura lly
occu rring. These a re alsothe 20 am ino acids
spec i
fiedinthegene tic code commonto a lll
ife.
The componen ts o fthe gene ra
ls truc ture o f
anam inoac id are
:1 )aca rboxy l(-COOH )g roup .
2) an am ino group ( -NH2 ) on the a lpha (a )
carbon and 3 ) an a lkyl(R )g roup a ttached to
the α C a tom. The am ino ac ids d ifferinthe irR
group . The R group g ives an am ino ac idi ts
ind ividua l chem ical charac terist
ic . Theforma tiono f a peptide bond o r
am idel inkageinvo lves cova len t bond ing be tween an am ino g roup of one
am ino acid andthe ca rboxy lg roup o fthe ad joining am ino ac
id . Prote ins
maycons is tofoneo r mo re po lypep tidecha ins he ldtogetherinthe p rotein
molecu le.
Am ino ac ids are com mon ly re ferred to by three-or one- letter
abbrev iations asl istedin Tab le 2 .1.

T
able2
.1 Name
sandabb
rev
iat
ionso
fthecommonam
inoac
ids

Abb
rev
iat
ion
Essen
tia
l Am
ino Ac
ids One
-le
tte
r
Th
ree-
let
ter

Arg
inine A
rg R
H
ist
idine H
is H
Iso
leuc
ine I
le I
Leuc
ine Leu L
Lys
ine Lys K
Me
thion
ine Me
t M
Pheny
lalan
ine Phe F
Th
reon
ine Th
r T
Tryp
tophan T
rp W
Va
line Va
l V

Non
-essen
tia
l Am
ino Ac
ids

Alan
ine A
la A
Aspa
rag
ine Asn N
Aspar
tic ac
id Asp D
Cys
teine Cys C
G
lutam
ic ac
id G
lu E
G
lutam
ine G
ln Q
G
lyc
ine G
ly G
Pro
line P
ro P
Se
rine Se
r S
Ty
ros
ine Ty
r Y
 CHAP
TER2 P
rote
insand Am
ino Ac
ids 9

The chemica
ls t
ruc tures o
f each o
fthese essen
tia
l am
ino ac
ids a
re
shownin Figure 2
.1.

Figu
re2 .
1
Chemica
lstruc
ture
softhetenessen
t
ialam
inoa
cid
s.

C
lass
if
ica
tiono
fAm
inoAc
ids

Am ino acidsareclassi
fied asessentialorind ispensable, and non-essentia
l
or dispensable. Essen t
ialo rindispensable am ino acids cannot be made
or synthesized bythe an imal o r which are synthesizedin amoun ts not
enoughto suppo r
t max imum g row th and haveto be p resentinthe irdiet.
The capac ity o
fdiffe
ren tfeed proteinsto mee tthe am ino acid needs o f
fish differs cons
iderab ly. The essen t
iality of an am ino acid w i
lla lso
depend on the an ima l beingfed. Fo r examp le
,g lycineis requ ired by
chicken bu tis no
tessen t
ialforf
ish . Non -essentia
lo rd ispensable amino
10 CHA
PTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

acids can be adequa tely syn

thes
ized bythe animalo rformedf rom o
ther
am ino acidsin sufficient amoun tsintissues and does no t haveto be
presentinthe irdiet
. Am ino ac
ids can a
lso be c
lass
ified accord
ingtothe
chem ical composit
ion o ftheirside cha
in aslisted below
:

1
. A
lipha
tic am
ino ac
ids

a
. basic - arginine,lysine
b
. ac
id ic - aspa r
tic acid,glutamicac
id
c
. neu tra
l - leucine,isoleucine,val
ine
,alan
ine
,glyc
ine
methionine,cys te
ine
threon ine, ser
ine

2
. Aroma
tic am
ino ac
ids
phenyla
lan
ine
,ty
ros
ine

3
. He
terocyc
lic am
ino acids
h
ist
idine,t
ryp
tophan
,pro
line

Essen
tia
l Am
ino Ac
ids

The re areten essen tial amino ac ids(EAA ) required byf ishfo rgrow th
and ma in tenance o fli
fe:a rginine ,histid
ine,iso leucine,leuc ine ,lys ine
,
methionine, pheny lalanine ,threon ine,tryptophan , andva l
ine . Asidef rom
beingthebu ild
ing b locks o fp roteins,some am ino acids are precurso rs o
r
supp ly par to fthe s tructure o fo ther subs tances . Me thion ineis the
precursoro fcys teine and cys t
ine. Me thionine also supplies me thy
l(CH3 )
groupsfo rc reatine,cho line, and manyo thersubs tances. When a hyd roxyl
(OH ) groupis added to pheny lalanine, tyrosineisformed . Tyros ineis
neededtofo rmthe ho rmonesthy roxine, epineph r
ine and no rep ineph rine
,
and me lan in pigmen ts
. Arg inine yields ornithine when ureaisfo rmed
inthe ureacyc le . Theremova lofaca rboxyl(COOH )g roupf rom h istidine
forms h istam ine . Tryp tophan is the precursor o f seroton in and the
vi
tam in,n icot
in ic acid. A llfinfishes have arequ i
remen tforthe same
ten essen tial am ino ac ids.

Non
-essen
tia
lAm
ino Ac
ids

The non -essen t

ialam inoac idsforfisha re:a lanine,aspa ragine, aspa rt
ic
acid
,cysteine ,glutam icacid,glu tamine,glycine,p ro
line,se rine
,andty rosine.
These amino ac ids have nu tritionalsignificance becausethe i r presencein
the diet conse rves ene rgytha tisrequ iredfor pro tein syn thesis. Some
dispensableam inoac idscanpa rt
ial
lyrep laceo rspa reind ispensab leam ino
acids
. Pheny la
lan inecanbepa r
tia
llyrep laced byty rosineand me thionine
bycystine.Thus ,requ iremen tsfo rpheny lalanineand me th ionine dec rease
whenty ros ine and cys tine, respective
ly,a re presen tinthe d iet.
 CHAPTER2 P
rote
ins and Am
ino Ac
ids 11

C
lass
if
ica
tiono
fPro
teins

Pro
teins are broad
ly c
lass
ified accord
ingtothe
irfo
rm, so
lub
ili
ty, and
physica
l propert
ies
.

□ Globular proteins have compac

tsphe r
icalorg
lobula
rshape. They are
most
ly solublein aqueous systems. Theyinclude
: enzymes,p ro
tein
hormones,p ro
teinsofthese rumfrac
tionoftheb
loodandantibod
ies.

□ Fibrous prote ins cons

istof a long cha in of polypep
tides
. They are
highlyinsoluble andresis
tanttoac tiono fthe digest
iveenzymes. They
include collagen,foundin carti
lage o r soft bones,bloodvesse
ls, bone
matrix,tendon,finsandsk in
;elastins,tha tareacomponen tofarte
ries
andl igamen ts
; and kerat
ins, in protect
ivecove r
ingssuchassk inand
scales.

Basedonphysica
lprope
rties
,pro
teinsa
reg
roupedin
tos
imp
le,con
juga
ted
,
and der
ived p
rote
ins:

□ Simp le prote ins yield on ly am ino ac ids orthe ir der ivatives when
hydrolysed . Theyinc ludea lbum ins(egga lbum in, se rum a lbum infrom
blood, lac toalbum inf rom m i
lk,leucos inf rom whea t
); a lbum inoids
(kera tinf rom ha ir
,f ingerna ils
,fea thers, wool,silkfib roin,e last
inf rom
connec tivet issue,co l
lagenf romca rti
lageandbones );g lobu lins(edes t
in
from hempseed , serumg lobu l
inf rom blood, lac tog lobu linfrom m ilk,
legum inf rom peas )
;h istones(g lob infrom hemog lobin , scomb ronef rom
sperma tozoa o f mackere l)
; and pro tam ins (sa lminef rom sa lmon ,
scomb rinef rom macke rel
). Theseg roupsa red if
feren tiatedbyso lubili
ty
invar ious so lvents such as wa ter, saltsolution,a lcoho l, and by othe r
cha rac teristics.

□ Con jugated prote ins y ield am ino acids and non -p rote
in componen ts
when broken down by hyd rolysis. Theyinc ludethe nuc leoproteins,
glycopro teins , phosphopro teins, hemog lob ins, andlec ithoproteins.
Nucleopro te ins are compounds o fone or mo re pro tein mo lecules with
nucleic acid presen tina llce ll nucle
i. G lycop ro
teinsa recompoundso f
the prote in mo lecule and subs tancescon taining aca rbohyd rateg roup
otherthan nuc le
ic acid(e .g.
, muc in
). Phosphop ro teins arecompounds
ofthe pro te in molecu le with a phospho rus -contain ingsubs tance other
than nuc le icacido rlecithin(e .g.
,case in)
. Hemog lob insa recompounds
ofthe pro tein mo lecule w ith hema t
in o r as im ilar subs tance(e .g.
,
hemog lobin ). Lec ithop roteins a recompounds o fthe p rotein molecule
withlecith in (e.g.
,t issuef ibrinogen).

□ Derived proteins consisto f compounds representing a

ltered and
degradedproductsofnatura
llyoccurringpro
teins
,producedbytheaction
of heat
, enzymes, or chemical agents
.
1
2 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

P
rote
inS
truc
ture

The compos i tion,o rganiza

tion and shape o fp ro
teins a re di
rectly related
tothe irfunc tion. Because o fthe complicated struc ture of proteins,a
newte rm ino logy has deve loped as an aidin desc ribing the s truc tural
organization o f a prote
in mo lecule. These a re designa ted as prima ry
,
seconda ry, te r
tiary, and qua ternary s
tructure ofp roteins.

□ P
rima
ry s
truc
ture descr
ibes the spec
ific sequencein wh
ich the
am ino ac ids a rel inked toge ther by pep t
ide bondsin a po lypeptide
chain(F igu re 2.2A )
. Fo r examp le, the pep tide Leu -G
ly-Thr-His-Arg-
Asp-Va l has a d ifferent prima ry structuref romthe pep tide Val
-Asp -
His-Leu -G ly-Arg-Thr , eventhough bo th havethe same number and
kinds o f am ino acids.
The pr imary struc ture de termines the three-d imens iona l
con forma t ion ofthe pro tein mo lecu le andi ts cel
lular ro
le. Ita lso
showsthe sequence o f nuc leot
idesin DNA o r RNA andthus p rovides
informa tion onthe gene ticinpu tto pro te
in syn thesis and cellular
F
igu
re2
.2A poten tial
. Thus , a changein any one am ino acidin the pr imary
Pr
ima rys
tructurecons
istso
fam
ino
ac
idsjoinedinsequenceb y struc ture o fa p rotein may p roduce a d rast
ic effec
tinthe an ima l.
pept
idebonds .

□ Seconda
ry s
truc
ture descr
ibes the mannerin wh
ich the am
ino
ac
ids are arrangedinthe po lypept
ide chain. This developsthrough
theinterac
tions be
tween adjacentam ino ac
idresiduesin apo lypept
ide
chain(Figure 2.2B)
.
Thesecondarystruc
tures mayexistintheformo fco
iled he
lices(a
lpha-
he
lix
) (Figure 2.2C
)o rin an extendedform(be ta
-sheet)(Figure 2
.2D).

F
igu
re2
.2B F
igu
re2
.2C F
igu
re2
.2D
Seconda ryst
ru c
turedevelopsthrough Thealpha-he
lixstructu
reofa Thebe ta-shee
tst
ructu
reofal
lorpart
interact
ionsbe tweenneighbor
ing po
lypept
idecha in
,o rparto
fthecha in
, ofapo lypept
idecha
inischarac
ter
ized
residues,asb yhydrogenbond s isstab
il
izedb yhyd rogenbonds byhyd rogenbonds(dot
tedl
ines
)
betweenad jacent monome runi
tsinan (do
ttedl
ine s
)be tweentheo xygen betweeneve rypept
idebondandits
a
lpha -he
lixregion
. (co
lor)ofonepep tidebondandthe neighbor
.
hyd
rogeno fanother.
 CHAPTER2 P
rote
insand Am
ino Ac
ids 13

The alpha -helix structureis s tabilized by hydrogen bonds be tween

the oxygen ofone andthe hyd rogen o fano ther pept
ide bond whilethe
beta-sheets truc tureis charac terized by hydrogen bonds be tween
adjacent peptide bonds . The hydrogen bond ing betweenthe pep tide
chainsin the be ta-sheetg ives r ise to a repea ted z
igzag structure
hence,the name p leated sheet. F ib rous prote
insexh ib
ita high degree
ofseconda ry s t
ruc tureform ing shee ts o
f mo leculesthata reinvolved
in cel
lular cons t
ruc t
ion.

□ Te
rtia
ry s
truc
ture descr
ibesthethree
-dimens
iona
lar
rangemen
tor
the ac tual con forma t
ion o fallthe a tomsin the pro tein molecu le.
Thein teract
ions be tween dis
tan t amino acidres idues ofa polypep t
ide
chainleads tothefo lding and a more g lobu lar conformation o fthe
polypeptide cha in assum ing ath ree d
imensiona l shape,for examp le,
myoglobin(F igure 2.2E).G lobularpro te
ins usua l
ly haveacons ide rable
amoun tofuno rde redrandomco i
lregions,along w ithregionsofo rdered
alpha-he l
ix and o thertypes o f secondary structu re
. The seconda ry
andte rtiary st
ruc ture ofa protein can be determ ined simultaneous ly.

F
igu
re2
.2E F
igu
re2
.2F
Te
rt
iar
ys truc
ture dependsonthe Quaternaryst
ructu
reistheresu
lto
f
in
terac
tionbetween moredis
tant interact
ionsbetweentwoor more
res
idues,lead
ingtofold
ingandtoa po
lypep t
idechainsinap ro
tein
mo
reglobularconformat
iono
fthe mo
lecu le(fou
rchainsinhemoglob
in)
.
po
lypept
idecha in
.

□ Qua
terna
ry s
truc
ture re
ferstothe spa tialorgan izat
ion whentwo
or more po lypep tide chains are par tofas ingle protein mo lecule.
Com mon ly occurr ing examp les are dimers , trimers , tetramers ,
cons is
ting o ftwo , three, andfour po lypeptide cha ins, respect
ively.
The po lypep t
ides a re he
ld toge ther by weak chem ical bonds(F igure
2.2F). Theabso lute numbe rofpo lypept
idesandthenumbe ro fdi
fferent
kinds o f po
lypep tidesin a pro tein var
ies. Fo r examp le, hemog lobin
molecules cons isto ftwo alpha- chains and two be ta- chains. Each
globincha inin hemog lobinisbondedtoa hemeg roup , whichfunc tions
14 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

inoxygent ranspor
ttobodytissues
.D isrup
tionofanyleve
lof mo
lecula
r
organ
ization,including qua
ternary st
ructure
,leads to mal
function
ofa pro
tein.

Fa
teo
fAbso
rbed Am
inoAc
ids

Proteins are absorbedf rom thein testine, ma inly as am ino ac ids. The
absorbedam inoac idsa re usedinoneo fthefo l
lowing ways :a)inco rporated
into the me tabolic poo l
, m ixed w ithf ree am ino ac ids originatingf rom
varioust issues and syn thesizedin tot issue pro teins; b ) syn thesized
into n i
trogen-con taining tissue cons ti
tuen ts such as nuc leic ac ids
,
hormones and enzymes ; c ) deam inated( removalo fnitrogen), resulting
in a carbon cha in( -C-C-) and an am ino g roup( -NH2 )
.
The ca rbon chains a re oxidizedto p roduce ene rgy or syn thesizedin to
sugars andfa ts,o r againreac twith am ino groups ,forming am ino acids.
Aminog roupssepa ra tedfromam inoac idsa reexc re
tedinthefo rmo fn it
rogen
compounds such as urea and ammon iainthe u r
ine o r excretedthrough
theg i
lls
.

Impo
rtanceo
fAm
ino Ac
idP
rof
ilesinF
ish Nu
tr
it
ion

Fisht issuescon tain abou t 65-75% p rotein on a d ry weightbasis . D ietary

proteino ftencons titutesthep r
incipa land mos texpens iveiteminf ishd iets.
Thef ree am ino ac ids releasedf rom pro teins bythe ac t
ion o fd igestive
enzymes a re abso rbedinthein testina lt ract and used by var ious ce l
ls
to bu ild and repa ir wo rn ou ttissues . Excess am ino acids a re used as
energy sou rce o r conve r
tedtofa t.
Info rma tionong rossp rote
inrequ i remen toffishiso flimitedvalue w ithou t
data on essen t
ial am ino acidrequ iremen ts since pro te
in quality depends
largely oni ts am ino acid compos i
tion and d igestibi
lity. The determ ina t
ion
of essen tial am ino ac id pro fi
lesis he lp fulinthe des ign of am ino ac id
testd iets used for research s tud ies to de term ine the am ino ac id
requiremen to ffish. I tis also animpo rtant pa rame terinthe eva luation
ofthe p rotein qua l
ity offeeds tuf
fs.
The nu tritive value o fa d ieta
ry pro teinis dependen t onthe ex tentto
whichthecompos ition ofitsessen tia l amino ac idsfu l
fil
lstherequ iremen t
oftheo rgan ism . Thec loserthep rofiletotherequ iremen t,the highe risthe
nutrit
iona lva lue. The am ino acidlack inginthe pro teinis known asthe
lim i
ting am ino ac id
. For examp le, lysineisl im i
tingin corn wh ile
methion ineisl im itingin soybeans .

Qua
li
tat
iveAm
ino Ac
id Requ
iremen
ts

Thetwo me thods usedto determ ine whethe ran amino acidisessen t

ialor
non -essentialare: 1) g rowthresponse me thod, and 2)therad ioisotope
method .
The g row thresponse me thod as used by Ha lver(1957) uses a series o
f
am ino acidtes td iets con
taining c rysta
lline L-amino acids as source of
ni
trogen . The d ie
ts areformu lated based onthe am ino acid pat
te rn ofa
reference pro tein such as who le ch icken egg prote
in, ch inook sa lmon
egg p rote
in,o r chinook yolk-sacf ry protein
.
 CHAPTER2 P
rote
ins and Am
ino Ac
ids 1
5

For each o ftheten essen tial am ino ac ids, a 10 -weekfeed ingt rialis
conduc ted us ing a basa ld ie
t con taining a llthe am ino ac ids and a d iet
deficientinthe am ino ac id be ingtes ted . F ish are weighed eve rytwo
weeksto measure g rowth responsetothetes td iet
. F ishfedthe am ino
acid -def
ic ien td iet show poo rg row th bu t a subs tantia
lg row th response
is obse rved whenf ishisfedthe comp lete am ino acid diet. Subsequen tly
,
othe rinves tiga tors used as imilartes td ietfo r determiningtheessen tia l
ity
of some am ino ac idsin o therf ishes.
Intherad ioisotope me thod used by Coweye tal.(1970),f ish arein jected
int raperitonea lly withrad ioac tivelylabe led 14Cg lucose andfed on a na tura
l
dietfo r7days .F isha rethenk i
lled, homogen ized,and p rotein wasiso lated .A
samp le oftheiso lated pro teinisthen hyd rolyzed andthecons ti
tuent am ino
acidsa resepa ratedbych roma tographyandcoun tedforradioactivity
.S ignificant
rad ioactivityisinco rpora tedin tonon -essen t
ialam inoacids wh iletheessen tia
l
am inoac idshaveve ryli
ttleo rnorad ioac tiv
ity. Th is method wasa lso usedby
Colosoand C ruz(1980 )tode te rminethequa litativeam inoac idrequ i
remen ts
oftige rsh rimps .

Quan
ti
tat
ive Am
ino Ac
id Requ
iremen
ts

The quan t
itative am ino ac id requ iremen ts off ishis de termined us ing
eithe r puri
fied o r sem i
-purifiedtes td ie ts. A pu r
if
ied d ie tis made up o f
pure subs tances(case in, gela t
in,c rys tall
ine am ino ac ids ) wh i
le a semi-
pur i
fied diet may con tain othe ringred ien ts such asf ish mea lo rsoybean
mea l
. Thetes td ietisfo rmula tedsotha ttheam ino ac id pro f
ileisidentical
totha tof a reference pro tein usua l
lyf ish musc le p rotein. A se ries of
expe r
imen tald ietsisthen p repa red con tainingg radedleve ls ofone am ino
acidfo r whichtherequ iremen tisto be de termined. D iets arefedtof ishand
the ga inin we ighta re measu red at week ly orb iweek lyinte rvals
. Therequired
dietaryleve
lisd etermined us
ingthedo s
ere spon s
ecu rve(F igu res 2.3A,B).

F
igu
re2
.3A F
igu
re2
.3B
Growthresponseo ft
ige
rshr
impfedgradedleve
lsof G
rowthresponseoftige
rsh r
impfedg radedle
vel
s
phen y
la
laninefor8 weeksa
sdescr
ibedb ythequad
rat
ic o
f meth
ion
inefor8 weeksa sdescr
ibedb ythe
regression mode
l. b
roken-
l
ine regre
ssion mode
l.
Sou
rce
: M
il
lamena e
tal
. 1999 Sou
rce
: M
il
lamenae
tal
. 1996
16 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

A dose response curveind ica tes the grow th response o fthef ish to
gradedleve ls ofanessen tial nu trien tinthe ird iet. Th is me thod assumes
tha t we igh t ga inisl inear ly rela ted toincreas ing d ietaryleve ls ofthe
essen t
ia l am ino ac id a t or be low the requ iremen tleve l. When the
requ iremen tis me t, we igh t gains abrup t
ly p lateau andthen dec linei f
the d ietary concen tration o fthe nu trient exceedsthe an ima l’sto lerance .
Figure 2 .3 (A ) shows a quadrat ic curve w i
th the requ irement o f
pheny lalan ine a t1 .4 pe rcen to fd ieto r3 .7 percen to fd ie tary p rotein .
The b rokenl ine ana lys is has a lso been usedto es tima tethe essen tial
am ino ac id requ i remen t. Th is me thod a lso assumes al inear re lat ion
between g row th and d ie taryleve lo fthe essen tial am ino ac id a to r be low
the requ iremen t. A tthe requ i remen tleve l
, thel inear ly ascend ingl ine
ins tantly b reaksto ho rizon tal
. The po intis known as b reakpo int. F igu re
2.3(B ) shows al inea r cu rve withtherequ iremen tfo r me th ion ine at0 .89
percen to fd ieto r 2 .4 pe rcento fd ie tary p
ro te
in.
Inyoung g rowing an ima ls,the g rea testp ropor t
ion o fbody we igh tisin
thefo rm o f musc le. I tisreasonab letoin fe rtha tthe d ieta ry am ino ac id
requ iremen tw i
ll be c lose lyrela tedtothe am ino ac id p rofile o f musc le o r
who le body p rotein . Thus , whenth is hypo thesis was exam inedfo rp igs ,
chicken ,f ish and shr imp , the am ino ac id requ iremen t was c lose ly
correlatedtothe am ino ac id pro file o fthe musc le p ro tein .
There a re seve ral prob lemstha t occurinthe accura te de term ina t ion
of am ino ac id requ iremen ts o ff ish when based on g row th s tud ies: a )
grow th ra tes tha ta re common ly obse rvedinthe am ino ac idtes td ie ts
aref requen tlyin ferior orlower than those observed w ith na tura lo r
intac t pro teins. b ) some o fthe am ino acidsintes td ie ts mayleach ou t
dur ingfeed ing. c ) thein terpre ta tion o fthe breakpo in tin the g row th
response curve cou ld be sub jec tivei fthe appropr iate s tatis t
ica ltoo lis
not used . Therequ i
remen ts ofsomef ishesandsh r
impfo ressen tialam ino
acids are shownin Tab le 2 .2.

Tab
le2
.2 Am
ino ac
id requ
iremen
ts o
f somef
ishes and sh
rimpin pe
rcen
tofp
rote
in

Ra
inbow Ch
inook N
ile Sea T
ige
r
Ee
l Ca
rp M
ilk
fish
t
rou
t sa
lmon t
ilap
ia bass sh
rimp

Arg
inine 4
.5 4
.4 4
.0 6
.0 5
.2 4
.2 3
.6 5
.3

H
ist
idine 2
.1 1
.5 1
.8 1
.8 2
.0 1
.7 2.2

Iso
leuc
ine 4
.0 2
.6 2
.8 2
.2 4
.0 3
.1 2.7

Leuc
ine 5
.3 4
.8 5
.0 3
.9 5
.1 3
.4 4.3

Lys
ine 5
.3 6
.0 6
.0 5
.0 4
.0 5
.1 4
.5 5
.2

Me
thion
ine + 5
.0 2
.7 3
.3 4
.0 2
.5 3
.2 2
.9 2
.4
Cys
/2

Pheny
lalan
ine+ 5
.8 5
.7 6
.0 5
.1 4
.2 5
.5 3.7
Ty
r

Threon
ine 4
.0 3
.8 4
.1 2
.2 4
.5 3
.8 3.5

Tryp
tophan 1
.1 0
.8 0
.6 0
.5 0
.6 1
.0 0
.5 0
.5

Va
line 4
.0 3
.4 3
.6 3
.2 3
.6 2
.8 3.4

Sou
rce
: NRC 1993
; FDS Manua
l 1994
; San
tiago and Love
ll 1993
; Bo
rlongan and Co
loso 1998
; Co
loso e
tal
.1999
;
M
il
lamenaetal
. 1996
-99
 CHAPTER2 P
rote
ins and Am
ino Ac
ids 17

De
fic
ienc
ies and Excesses o
fDie
tary Am
ino Ac
ids

Essen tial am ino ac id de f

iciency may be caused by the use o ffeed
ing red ien ts usua lly protein sourcestha ta re sl
igh t
ly o rg rossly de fic
ien t
in a t leas t one essen t
ial am ino acid
. Am ino ac id de f
iciency may a lso
arisef romthepresenceo fchem ica
lstha t maya f
fect somefeeding redien ts,
excess ive hea tt rea tmen t dur ingfeed manu fac ture, andleach ingou t o f
nu tr
ien ts.
Am ino ac id imba lance due to am ino ac id an tagon ism o r am ino
acid tox icity may a lso cause reduced g row thin an ima ls. Am ino ac id
an tagon ism occurs when some am ino ac ids a re fed in excess o f the
requ ired leve ls, caus ing an inc rease in the requ iremen t fo r ano the r
am ino ac id o fs im ilars tructu re. Someexamp lesa retheleuc ine -isoleuc ine
an tagon ism and a rginine-lysine an tagonism obse rved in somef ish
spec ies. Am ino ac idtoxicityoccu rs when excess amoun ts o f ce rta in
am ino ac ids a re fed to the an imal and the nega tive e f
fec ts canno t be
improved by add ing am ino ac ids in the d iet
.
In prac tica ld ietformu lation, the recommended d ietaryleve ls o f
essen tial am ino ac ids can be me t by care fu lly se lec ting and
prope rly comb ining two o r mo re p rotein sou rces. The l im it
ing o r
deficien t am ino ac id in one p rote
in sou rce can be supp lemen ted by
ano the r p ro tein sou rce abundan tin the same am ino ac id and hence
make a be tterfeed . Ano ther way o f mee t
ingthe EAA needs o fan an ima l
is by supp lemen ting the prac ticald iet w ith crys ta l
line L -am ino
acids . Nu trien t leach ing can be m inim ized by us ing wa ter s tab le
diets throughthe use o fe f
ficientb inders and emp loy ing appropr ia te
feed ing prac t
ices .

Eva
lua
tiono
fPro
tein Qua
li
ty

Proteins a re sa idto be ofhigh qua lity whenthe i r amino acidcompos it

ion
closelyresemb lestheam inoac idpa tterno rapp roxima testheessen tialam ino
acid requ iremen ts ofthe an ima l unde r conside ration and whenthey a re
highly d igested bythean ima l
. The qua l
ityo fp roteinsis usua llyeva luated
byb iolog icalandchem ical methods . Thechem ica l method dete rm inesthe
quan tityo fp roteinoram inoac idinafeeds tuffwhe reastheb iological method
determ ineshowthef ishreactstothep roteininte rmso fgrow thandsu rv
ival.
Inthe b iological me thod
, body we igh tgain and n i
trogenre tention a re used
asc riteriaforpro te
inqua li
ty wh ichiscons ideredas mo reaccu ra tethanthe
chem ical me thod . Prote
ine ff
iciencyra tio,b io
log icalvalue, and ne tp rotein
uti
liza t
ion a re usedto measuretheb iologicalva lueo fap roteinandcanbe
calcula ted, by us ingthefo l
low ingfo rmu la:

1
. P
rote
ine
ffic
iencyra
tio(PER
)
L
ive we
ightgain(g
rams)
PER =
Amountofprote
infed(g
rams)
18 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

2
.Bio
log
ica
l va
lue(BV
)*

BV = True n
itrogenreta
ined
x 100
Ni
trogen abso
rbed

whe
re,R =true ni
trogenreta
ined
A = n
itrogen abso
rbed

and A = I -(F-Fo
),
R = A-(U- Uo
)

whe
re,I = nit
rogenintake
F = nitrogenexcretedinthefeces
Fo = metabol
icfaecaln i
trogen
U = nit
rogenexc re
tedinthe u rine
Uo= endogenous nitrogen

R I-(F-Fo)-(U- Uo
)
Thus
, BV= x 100 x 100
A I-(F-Fo
)

* Insu f
fic
ient da
taonthe bio
log
ica
lvalue a
re ava
ilab
leforfish dueto
di
fficult
iesin determ
iningthe metabo
licfaecal and endogenous
n
itrogensepa rate
ly

3
. Ne
tpro
teinu
til
iza
tion(NPU
)

True n
itrogenreta
ined
NPU = x100
Ni
trogenintake

whe
re, NPUis de
term
ined bythefo
llow
ingformu
la

N
itrogenincreaseinf ish Nit
rogen decreaseinf
ish
fedthetestproteindiet + fedthep ro
teinfreed
iet
NPU = x 100
Ni
trogenin takefromthetes
t protein die
t

A de
tai
ledp
rocedu
refo
rpro
teineva
lua
tionisd
iscussedin Chap
ter6
.

P
rote
in Requ
iremen
t

The op timum pro tein requiremen tis theleve lofh igh qua li
ty dietary
protein neededfo r maximumg row th.To determinethepro teinrequ i
remen t
of af ish spec ies,feed ingtria ls have to be conduc ted us ingtes td iets
con tain ing gradedleve ls ofp ro te
inf rom sou rces o
fh igh biolog
icalva lue.
Grow th response , usua l
ly weigh t gain,is measuredfor each d iet
. The
diettha tgivesthe h ighest weigh t ga in and surv
iva lis considered asthe
bestd iet
. Max imum t issue pro tein retent
ion may a lso be used asthe
criterionfor de term ining pro tein requ i
remen tins tead o f weigh t gain.
Thisis done by ana lyzingthe amoun to fnit
rogeninthet issues atce rtain
inte rva ls e
.g. everytwo weeks un ti
lthe reis no mo re decreasein n it
rogen
retainedinthet issues.
 CHAP
TER2 P
rote
insandAm
inoAc
ids 19

The minimum amoun tofd ie

tary prote
in neededfor opt
imum g row
th of
most cu
ltured aqua t
ic spec
ies rangesf rom 27%to 60% . A summa ry of
optimal proteinlevels and the prote
in sources used to de
term
ine these
values are presen
tedin Tab le 2.3.

T
abl
e2.
3Op
tima
ldie
tary p
rote
inleve
ls(% o
fdry d
iet
) fo
r some aquacu
ltu
re spec
ies

Op
tima
l
Spec
ies Pro
tein sou
rces p
rote
in Re
ferences
leve
l

As
ian sea bass f
ish mea
l, soybean mea
l 43 Ca
tacu
tan & Co
loso 1994

Common ca
rp f
ish mea
l, case
in 31
-38 Takeuch
i 1979

Groupe
r tuna
, musc
le mea
l 40
-50 Teng e
tal
.1978

f
ish mea
l, mea
t mea
l, sh
rimp mea
l 43 Sh
iau 1972

Japanese ee
l case
in + am
ino ac
ids
. 44 Nose & A
rai 1972

Ku
ruma sh
rimp squ
id mea
l 60 Desh
ima
ru & Sh
igeno 1972

case
in + egg a
lbum
in >55 Tesh
ima & Kanazawa 1984

M
ilk
fish f
ish mea
l, case
in 40 L
im e
tal
. 1979

case
in, ge
lat
in 30
-40 Pascua
l 1989

f
ish mea
l, soybean mea
l, 24* Sumagaysay & Bo
rlongan 1995

cassava mea
l

Red sea b
ream case
in 55 Yone 1976

Snakehead f
ish mea
l 52 Wee & Tacon 1982

Red Snappe
r f
ish mea
l, soybean mea
l, squ
id mea
l 44 Ca
tacu
tan & Pagado
r2001

T
igersh
rimp case
in 40 A
lava & L
im 1983

f
ish mea
l, soybean mea
l, 40 M
illamena & T
riño 1994

sh
rimp mea
l

N
ilet
ilap
ia f
ish mea
l, case
in 30 Wang e
tal
. 1985

f
ish mea
l 28 San
tiago e
tal
. 1986

Wh
ite sh
rimp musse
l mea
l,f
ish mea
l, co
llagen
, 34
-42 And
rew & S
ick 1972

squ
id mea
l 28
-32 Sedgw
ick 1979

Ye
llowta
il f
ish mea
l, case
in 55 Takeda e
tal
. 1975

Aba
lone soybean mea
l,r
iceb
ran 27 Bau
tis
ta-Te
rue
l& M
il
lamena 1999

f
ish mea
l, squ
id mea
l

*tes
tedunde
rpondcond
it
ions

Proteino fhighb iologica lvaluesuchas who leeggp ro

tein,casein ,combina t
ion
ofcase in and gelatin, andf ish mea lare often usedin de term ining protein
requ iremen ts. Severalfac torsinfluence proteinrequ irementsfo r max
imum
fish g rowth: species,f ish size or age
, wa tertempe rature,p rotein qua
lity as
reflected bythe am ino ac id prof
ile
,d ietarylevelo f non protein energy and
dai ly food a llowance . Sma l
ler or younger f ish have h igher pro tein
requ iremen tsthan o lde rfish ofthe same spec ies.F ishfed a tfeedingra tes
belowsa tiat
ion willrequ ire a h
ighe r proteinleve
l. F ish a
lsorespond be tter
to a h igherlevelo fd ietary prote
in whenfed poo r qua l
ity p
ro tein
.
20 CHA
PTER2 NUTR
ITIONINTROP
ICALAQUACULTURE

Gu
ideQues
tions

1
. Wha
tisa p
rote
in? Wha
tisthe mos
timpo
rtan
tfunc
tiono
fa p
rote
in?
2
. Howa
re p
rote
insc
lass
ified? D
ist
ingu
ish be
tweens
imp
le,con
juga
ted
,
and de
rived p
rote
ins
.
3
. Wha
tisthegene
rals
truc
ture o
fam
inoac
ids?
4
. Whya
re am
ino ac
idsca
lledthe bu
ild
ing b
locks o
fpro
teins?
5
. Namethetenessen
tia
lam
ino ac
idsrequ
ired byf
ish
.
6
. D
iffe
ren
tia
te be
tweenessen
tia
land non
-essen
tia
lam
ino ac
ids
.
7
. Desc
ribethefa
teo
fabso
rbed am
ino ac
ids
.
8
. Whyistheam
inoac
idp
rof
ileo
fafeedava
luab
leindexintheassessmen
t
o
fthe nu
tri
tiveva
lue o
fdie
tary p
rote
ins?
9
. Wha
tisthed
iffe
rencebe
tweenqua
lita
tiveandquan
tita
tiveam
inoac
id
requ
iremen
t? Desc
ribeeach b
rie
fly
.
10
. Wha
tarethetwo me
thods usedto de
term
ine qua
lita
tive am
ino ac
id
requ
iremen
ts?
11
. Desc
ribe a dose
-response cu
rve usedto de
term
inethe quan
tita
tive
am
ino ac
idrequ
iremen
t.
12
. Wha
tarethecauses o
fdie
taryessen
tia
lam
ino ac
id de
fic
iency?
13
. Wha
taresomeo
fthe p
rob
lemstha
toccurin de
term
ining essen
tia
l
am
ino ac
idrequ
iremen
ts?
14
. Exp
lain br
ief
ly howtheessen
tia
lam
ino ac
idrequ
iremen
tscan be me
t
inap
rac
tica
ldie
t fo
rmu
lat
ion
.
15
. Ineva
lua
ting pro
tein qua
lity
, whyisthe b
iolog
ica
l me
thodcons
ide
red
mo
re accu
ratethanthechem
ica
l me
thod?
 2
1
LIP
IDS ANDFATTYAC
IDS

In
troduc
tion
Lipids are a d iverse group o f organ ic compounds tha t areimpor tan t
componen tso fp lan t, an ima l
, and m icrob ial memb ranes . Theya reinso luble
in wa terbu tso lub lein nonpo lar,o rgan icso lventssuchase theranda lcoho l.
Dietaryl ipid hastwo ma infunc tions: as sou rce of me tabo lic energy and as
sou rce o fessen tia lfa tty ac idstha t have spec ificfunc t
ionsinthe bodysuch
asfo rce llula rst ruc ture and ma intenance o fthein tegrityo fb iomemb ranes .
Lipids a reimpor tan t componen ts o ff ish d iets becausethey supp ly a
concen tratedsou rce o fene rgytha tis we llu ti
lized byf ish . They a lsosupp ly
essen tialfa t ty ac ids wh ich canno t be syn thesized byf ish. As a sou rce o f
dietary ene rgy ,l ipids have been shownto spa re some p roteinfo rg row th.
Lipids a re a lsoimpo rtan t sou rces o fs te rols, phospho lip ids, andfa t
-so luble
vitam ins . Fa t
ty ac idsf rom d ietaryl ipids may a lso se rve as p recurso rs o f
steroid ho rmones and p ros tagland ins.
The ob jective o fth issec tionisto acqua in tthereade rabou tcommonfa t
ty
acids,the ir nomenc latu re andfo rmu las, and d if
fe rentia te betweensa tu rated
and unsa tura ted fat ty ac ids ; to know how env i ronmenta l fac tors
(tempe rature , salin i
ty,d iet)in f
luencethefa tty acid compos i
tion offish; the
mechan isms o ffa tty acid b iosyn thes is and ox ida t
ion , andfac torstha tfavo r
fa t
ty ac id b iosyn thes is and ox idation ;the e ffects o fl ipid pe roxidation and
thefunc t
ion o fan tioxidan ts ; andto unde rstandtheimpo r
tance o ffa t
ty ac id
pro fi
lesin f ish nu trit
ion , and d ifferences in the essen tialfa tty ac id
requ iremen ts o f wa rmwa te r and co ldwa terfishes .

Typeso
fLip
ids

□ T
rig
lyce
rides orfa
ts a
refo
rmed bythereac
tion o
fglyce
rolw
ithfa
tty
acid moleculesandhenceca lled
glycerides . Thus when a
triglycerideis hydro lyzed,3
moleculeso ffattyacid andone
moleculeo fglycero
la reformed.
Triglycer
ides do no t occur as
componen ts of biomemb ranes
bu t they accumu latein the
adiposeo rfa tt
issues. Theya re
the pr imary means by wh ich
an imals store energy.

□ Phospho
lip
ids are es
ters o
f
fattyacidsandphospho ricacid
g
lycero
l t
rig
lycer
ide
(H3 PO4) and a n itrogenous
base. Theresu lt
ingcompound
is cal
led a phospha tidic ac
id. Some o ftheimpo rtan
t phospho l
ipids a re
phospha tidy
l choline( leci
thin)
, phospha tidyle thanolamine(cepha l
in),
phospha tidyl serine, and phospha tidy
linos i tol
. They a rethe ma in
componen ts o
fb iological memb ranes
.
22 CHA
PTER2 NUTR
ITIONINTROP
ICALAQUACULTURE

Nameo
fX-OH Fo
rmu
lao
f-X
cho
line HO- CH2- CH2- N+
(CH3
)3

e
thano
lam
ine h o –c h 2 –c h 2 –n +
h3

- C- coo
-

□ Waxes are esters o

flong
-cha
in fa
tty ac
ids and o
fhigh mo
lecu
lar
we
igh t monohydric alcohols
. Liketriglyce
rides, they are sources o
f
energy s
toredin p lants and an
imals and serve as pro
tective coat
ing
.
They are so
lids at ambienttemperature.

a
. some areeste
rs o
f long
-cha
in a
lcoho
ls, R1
-CH2OH andlongcha
in
fa
tty ac
ids
, R2-COOH
O
I
I
fo
r examp
le. R2
-CO
-CH2R1

b
. some a
re e
the
rs, R2
-CH2
-O-CH2
-R1

□ S
tero
idsare usua
llypo
lycyc
liclong
-cha
ina
lcoho
ls. Theya
rep
recurso
r
of sex or other hormonesinf ish and shrimp and are biologically
impor tantin the reproductive processes. S tero
ids have the same
generalstructure consis
ting of afused-
ring system. Cho
lesterol is a
phys io
logically important sterol and is w
idespread in b iological
membranes , especia
llyin anima ls
.

s
tero
id nuc
leus cho
les
tero
l
 CHAP
TER2 L
ipid
s and Fa
ttyA
cid
s 23
□ Sph
ingomye
lins do no
t con
tain g
lyce
rol
, bu
tarefa
tty ac
id es
ters o
f
long-cha
inam inoalcoholsphingosine
. Theyarelip
idcomponen
tso
fthe
bra
in and ne
rvet issue ofplants and an
imals.

Gene
ralFunc
tiono
fLip
ids

The genera lfunc t

ions a re :
1. Sourceso f metabol
icene rgy,adenos inet riphospha te(ATP ). Theycon tain
approx ima telytwicetheene rgyo fproteins andca rbohyd rates.
2
. Sources o fessen tia
lfa t
ty ac ids(EFA ) which a reimpo r
tan tfo rg rowth
and su rvival. EFA canno t be syn thesized bythe an imali tselfo rare
synthes izedininsu fficien t amoun tsfor good grow th and hasto be
providedfo rinthe d iet . Examp les:lino lenic,linoleic,a rachidon ic ac
id
(ARA )
,e icosapen taeno icac id(EPA ),anddocosahexaeno icac id(DHA )a re
essentia lfa t
ty acidsinf ish andc rustaceans .
3. Essen tia l componen ts o f ce
llular and sub -cel
lular membranes .
Examp les : phospho l
ip ids and po lyunsa tura tedfa t
ty acids.
4
. Sources o fs tero
ids wh ich per formimpo rtan tbiologicalfunc t
ions such
as ma in tenance o f membrane sys tems ,lipidt ranspo r
t, and p recursors
ofstero id ho rmones .

Fa
ttyAc
ids

S
truc
tureand C
lass
ifica
tion
Fa
ttyac
idsareanimpor
tantcons
tituentofl
ipid
s. Ove
r40fattyacidsa
reknown
tooccu
rnatura
lly
.Theycanberepresentedbythegenera
lfo
rmu la:

CH3
(CH2
)nCOOH

whe
re: nva
riesf
rom0to24andisusua
llyanevennumbe
r

Most natura
lly occurringfa t
ty acids con tain a single ca rboxy
l(COOH )
group atoneend and as tra
igh t unb ranched hyd rocarbon(C )chain. Fatty
acids may be saturated (no doub le bond ), where all ca rbon atoms are
fi
lled with hydrogen, or unsa tura ted , where one or more carbon a tom
lacks a hydrogen a tom . Unsaturated fat ty ac ids may e i
ther be
monounsa turated (one doub le, bond ),o r polyunsatura ted (PUFA ) wi
th
two or more doub le bonds or h igh ly unsaturated fa tty acids (HUFA )
contain
ingfour doub le bonds o r mo re.
24 CHA
PTER2 NUTR
ITIONINTROP
ICALAQUACULTURE

Polyunsaturated fatty acids normal

ly have a methylene (-CH2-)
in
te rrup
ted sys
tem of double bonds
. The commonfat
ty ac
ids are shown
in Table 2
.4.

T
able2
.4 Thecommonfa
ttyac
ids
Common Name Chem
ica
l Name Sho
rthand
No
tat
ion*

Sa
tura
ted
Form
ic, Ace
tic
, Prop
ion
ic 1
:0,2
:0,3
:0
Bu
tyr
ic, Va
ler
ic Bu
tano
ic ac
id, Pen
tano
ic ac
id 4
:0,5
.0
Cap
roic Hexano
ic ac
id 6
:0
Capry
lic Oc
tano
ic ac
id 8
:0
Cap
ric Decano
ic ac
id 10
:0
Lau
ric Dodecano
ic ac
id 12
:0
My
ris
tic Te
tradecano
ic ac
id 14
:0
Pa
lmi
tic Hexadecano
ic ac
id 16
:0
S
tea
ric Oc
tadecano
ic ac
id 18
:0
Arach
idic 20:0
Behem
ic 22:0
L
ignoce
ric 24:0

Unsa
turated
Pa
lmi
tole
ic ac
id Hexadeceno
ic ac
id 16
:1 n
-7
O
leic ac
id Oc
tadeceno
ic ac
id 18
:1 n
-9

Po
lyunsa
turated
L
ino
leic ac
id Oc
tadecad
ieno
ic ac
id 18
:2 n
-6
L
ino
len
ic ac
id Oc
tadeca
trieno
ic ac
id 18
:3 n
-3

High
ly unsaturated
Arach
idon
ic ac
id E
icosa
tet
raeno
ic ac
id 20
:4 n
-6
E
icosapen
taeno
ic ac
id 20:5n
-3
Docosahexaeno
ic ac
id 22:6n
-3

Nomenc
latu
re
Fat
ty ac ids are g
iven a common name bes ides their chemica
lformu la
and sho rthand abbrev
iation. Infa t
ty ac
id nomenc la
ture, afatty ac
idis
ident
ified bythefo rmula:A :Bn-3 , A:B n-6, A :B n-9, sometimes n is
des
igna ted as ω(omega) whe re
, Aisthe number o f carbon atoms , B
isthe number o f double bonds , n-3
,n -6
,n -9isthe posit
ion ofthefirs
t
double bondf rom the me thyl end ofthefatty acid
. For examp le, the
 CHAPTER 2 L
ipids and Fa
tty Ac
ids 25
numerical designat
ionfor pa lmitole
ic or hexadecenoic ac
idis 16 :1 n-7.
This meanstha t pa
lmitoleic acid has 16ca rbons andcontains one double
bond wh ich appears onthe seven th carbon, numberingfromthe me thyl
end ofthefa t
ty acid chain.

Based upon the classif

icationin Tab le 2.4, unsa turatedfa t
ty acids
may be div
idedin to three ma jorfam i
lies: the o
leic o rn -9 ser
ies, the
lino
leic orn -6 se
ries, and thelinolenic or n-3 series.
Each fam i
ly name represents the shor test cha in member o fthe
group. Theshor thand notation andthe irrespectives tructuralformu las
are g
ivenin Tab le 2.5.

T
able2
.5 Unsa
tura
tedfa
ttyac
idsfam
il
ies
Sho
rthand*
C
lass Fam
ily S
truc
tura
lfo
rmu
la
no
tat
ion

n
-9 O
leic 18
:1 n-9 CH3- (CH2
)7- CH = CH- (CH2
)7- COOH
20:1 n
-9

n
-6 L
ino
leic 18
:2 n
-6 CH3- (CH2
)4- CH = CH - CH2- CH = CH- (CH2
)7- COOH
18:3 n
-6
20:3 n
-6
20:4 n
-6
22:4 n
-6

n
-3 L
ino
len
ic 18
:3 n
-3 CH3- CH2- CH = CH- CH2- CH = CH- CH2- CH = CH- (CH2
)7- COOH
20:5 n
-3
22:5 n
-3

*Nu
mbero
fcarbon(C)atoms:numbe
rofdoub
lebondsandpos
it
iono
fthef
irs
tdoub
lebond
,coun
tingf
romthe me
thy
l
(CH3
)end o
fthefa
tty ac
id

The degree o f unsa turat

ion o ffatty acids influences the phys ical
propertyo fthefa ts. Ingeneral, unsaturatedfatty ac
ids are morereactive
chem ically and have lower me l
ting po ints than the correspond ing
saturatedfa tty acids. Plantoils arel
iquid a troomtempe rature because
they have h ighe r propor
tions of unsaturatedfa t
ty acidsthan do an imal
fats which tendto be so l
ids.
Mostfish a re di
fferentfrom terrestria
l an imalsintha ttheirtissues
containfairly high amoun ts ofn-3 highly unsaturatedfatty ac
ids(HUFA ).
26 CHAPTER 2 NUTR
ITIONINTROP
ICALAQUACULTURE

Fa
ttyAc
id Compos
it
iono
fFish

The composi
tion offatty ac
ids in f
ish is a
ffected by a number o
f
env
ironmenta
lfactorsl
ike sal
ini
ty, tempe
rature, and d
iet
.

1
.Sa
lin
ity
Fishl ivein env ironmen ts o fvary ing sa l
ini
ty. The ma jorfa t
ty ac ids
inl ipids of marine andf reshwa te
rf ishes are shownin Tab le 2 .6
. The
fatty ac id compos ition o ffreshwa ter and mar inef ishes d ifferf rom
each o ther:
a
. ma rinespecies have h ighe rcon ten to flong-cha in(20and 22ca rbon )
monoeno ic acids wh ilef reshwa ter spec ies have h igherleve ls of
medium -chain(16and 18ca rbons ) monoeno ic ac
ids;
b
. ma rine species contain mo re h igh ly unsatura ted fa tty acidsthan
freshwa terspecies;
c
. thera t
io o fn-3fa tty ac idsto n -6fa tty acidsis g reaterfo r ma r
ine
thanf reshwa terspecies .
Di
fferencesinfa tty ac id compos i t
ionis re flected alsoinf ishtha t
migratef romf reshwa terto ma r
ine env ironmen t
. Asf ish mig ratef rom
freshwa te
rtoseawa ter,the irfattyac idcompos itionchanges . The n -3/
n-6ra t
iois h igherfo r m ig rato
ryf ishesl ike sme l
ts and sa lmon .

T
able2
.6 Ma
jorfa
ttyac
idsinl
ipidso
f ma
rineandf
reshwa
terf
ishes
*

Pe
rcen
t Fa
tty Ac
ids
Fa
tty Ac
id
Ma
rine Freshwa
ter
A B C D

14:0 3
.7 2
.2 2
.8 6
.7
14
:1 0
.1 0
.2 1
.0 0
.7
16
:0 12
.6 17
.0 16
.6 14
.6
16
:1 9
.3 4
.1 17
.7 14
.7
18
:0 2
.3 3
.2 3
.3 1
.5
18
:1 22
.7 21
.4 26
.1 18
.2
18
:2 n
-6 1
.5 2
.0 4
.3 3
.7
18
:2 n
-3 0
.6 1
.0 3
.6 3
.6
20
:1 7
.5 5
.4 2
.4 1
.6
20
:4 n
-6 1
.4 0
.9 2
.6 2
.4
20
:5 n
-3 12
.9 6
.7 2
.7 8
.2
22
:1 6
.2 9
.4 0
.3 0
.4
22
:4 n
-6 0
.1 0
.6 0
.4 0
.4
22
:5 n
-3 1
.7 2
.3 2
.0 1
.5
22
:6 n
-3 12
.7 16
.1 2
.0 6
.0

To
talsa
tura
ted 18
.6 22
.4 22
.7 22
.8
To
tal monoenes
med
ium 32
.2 25
.7 44
.8 33
.6
long
-cha
in 13
.7 14
.8 2
.7 2
.0
To
taln
-3 27
.9 26
.1 10
.3 19
.3
To
taln
-6 4
.1 3
.5 7
.3 6
.5
6
.8 7
.5 1
.4 3
.0
Ra
tio n
3-
n-6

*A=A t
lan
ticcod C=Sheepshe
rd
B=Chinooksalmon D=Alew
ife
Sou : A
rce c
kman 1976
 CHAPTER 2 L
ipids and Fa
tty Ac
ids 27
2
. Tempe
ratu
re
Tempera tureis a ma jorfac tortha t causes d if
ferencesinfa t
ty acid
compos it
ion.F ishtha tlivein wa rmwa ters con tain mo resa tu
ratedfa tty
acids than f ish tha tlivein co lder wa ters. F ish reared a th igher
tempe ra turesaccumu late mo resa tura tedfattyac idsthanthesamef ish
acclima ted atlowe rtempe ra tures
F
ishandc rus
taceansa reab letoad jus ttheirlevelsofpo lyunsaturated
fatty acidsto ma intain memb ranein teg r
ity andfunc t
ioninthe co ld.
Most PUFAsrema ininthel iqu id stateeven a tlowtempe ratures while
satura ted fatty acids congea l and so lid
ify a t coldertempera tures ,
therefore ,highleve ls o f unsa turatedfa t
ty ac idsin co ldwa terfishis
necessaryfor ma in tenance o f membranef luidity.

3
. D
iet
Dietisthela rgests ing lefac tortha ta ffectsthefa tty acid compos i
tion.
Unde r norma lcond itions ,thefa t
tyac idcompos itiono ffishcomesf rom
thesesou rces :
a. fatty acids de r
ivedf romthe d iet
;
b. fatty acids de rivedf rom non -l
ipid sou rces by biosyn thesis and
c. fatty ac ids derivedf roml ipid sou rces by b
iosyn thesis
The effecto fd iet onthefa tty ac id composition o ffish and sh r
imp
lipids has been demons t
ra ted. Whenthe sh rimpPenaeus se ti
ferus is
fed a dieth ighinl ino leic ac id(18 :2n-6 ) andlowin C20 and C22 PUFA
fo rone mon th,theshr imp ’scon ten tofn -3and n-6fa ttyac idsis mod i
fied
(increasein n -6and dec reasein n -3), hencethen-6/ n-3ra t
ioinc reases.
Afterthree mon ths,the n -6/n -3ra tioo fsh r
impcon tinuestore f
lecttha t
ofthe d ie taryl ipid (Tab le 2.7 )
. The seasona l var iation o fthefa tty
acid compos ition off ish may a lso bein f
luenced bythe changesinthe
compos ition o fthe ir na tura lfood .

le 2
Tab .7E
ffec
tofd
ietonfa
ttyac
idcompos
it
iono
fsh
rimpPenaeus setiferus
| Compos
ition o
f Penaeus se
tife
rus l
ipid
Fa
tty ac
id 0 mon
th 1 mon
th 3 mon
ths D
iet

14
:0 0
.6 0
.5 0
.5 1
.6
16
:0 14
.8 13
.4 15
.0 15
.5
16
:1 11
.2 8
.7 10
.0 7
.9
18
:0 5
.1 2
.3 2
.2 1
.7
18
:1 13
.1 22
.9 20
.0 28
.4
18
:2 n
-6 2
.3 18
.1 14
.1 32
.3
18
:3 n
-3 2
.8 2
.1 1
.3 4
.4
20
:4 n
-6 11
.6 9
.4 10
.3 0
.7
20
:5 n
-3 10
.4 8
.7 9
.7 2
.6
22
:6 n-3 11
.3 6
.3 6
.9 0
.3

To
tal sa
tura
ted 26
.6 22
.6 25
.6 25
.0
To
tal monoenes 18
.2 25
.2 22
.2 30
.1
To
taln
-6 13
.9 27
.5 24
.4 33
.0
To
taln
-3 24
.5 17
.1 17
.9 7
.3
n
-6 0
.57 1
.61 1
.36 4
.5
Ra
tio
n
-3

Source: Cas
tel
l19
81
28 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

B
iosyn
thes
iso
f Fa
ttyAc
ids

Fish can biosynthes izefa t

ty ac idsf rom a non -
lipid source(acetate)
. The
process beginsf rom ace tyl CoAand unde rgoescha ine longat
ion byadd ing
two carbon un i ts at at ime asi t progresses through the b iochemica l
pathway. The enzymes tha t catalyze the syn thesis offatty ac
ids a re:
acetyl CoA carboxy lase and FA syn the tase. Ace tate f
ragments are
added to afa t
ty ac id bythe process o f chain elonga t
ion. The reaction
occursin bo ththe m i
tochondr ia and m icrosomes . Becausefa t
ty acids
are synthes ized from two -carbon ace ty l units, they usua l
ly have an
even number o f carbon a toms . Pa lm itate is the ma jor fatty ac id
produced by de novo b iosyn thesis.

Denovo syn
thes
iso
fsa
tura
tedfa
ttyac
ids

Ace
tate → C4 → C6 → C8 → C10 → C12 → C14 → C16
:0 (cha
in e
longa
tion
)
pa
lmi
ticac
id

Inthe syn thesis o

f unsatu ratedfatty acids
, desatura t
ion o rremova lo fa
hydrogen mo lecu
leisanadd it
ionals teptha tisrequiredto make a doub le
bond( -C=C -
). Desa turat
ion o ffa
tty acids occursinthe m icrosomes . Of
the monounsa turatedfa t
ty ac ids howeve r
, only those o fthe n-5 ,n -7,
n-9 , n-11 and n-13 series a reformedf rom non-lipid sources
. De novo
biosynthes
is ofsatu ra
ted and monoeno ic fattyacidsisfur therillustrated
in Figure 2.4. The ma in s i
tesforde novo fa tty acid synthesis arethe
liver and the ad ipocytes. The process occurs when the ce l
ls have
abundance o fene rgy and have h igh ATP con tent
.

F
igu
re2
.4
Denovo syn
thes
iso
fsa
tura
ted monoeno
icfa
ttyac
ids

PUFAs are made by a se r

ies ofcha in elongation and desa turation steps.
The cha
in elonga t
ion and desa turation o fthe n-3 andn-6fa t
ty acids are
shownin F igure 2.5.
In common w ith other ver
teb rates,fish canno tdi rec
tly synthesize n-3
and n-6fat ty ac
ids de novo f rom non -l
ipid sources
. D ietary precursors
such as o
leic,linole
ic, andlinolenic acids should be suppliedinthe ird ie
t.
 CHAPTER2 L
ipid
s and Fa
tty Ac
ids 29
Biosynthes is o ffa tty ac ids
may be contro l
led by the
compe ti
tion be tweenthe d i
ffe rent
series offa tty acids fo r enzymes
tha t desa tura te andenzymestha t
cha in elonga te. H igh leve ls o f
n-3 inh ibit the e longa tion and
desa tura tion o fthe n -6 ser ies.
The potency o f inh ib it
ion is
ranked asfo llows:n -3> n -6> n -9.
Thus , aprope rba lanceo fn -3and
n-6essen tia lfa t
ty ac idsis needed
informu latingthe d iet
. A f
terthe
biosyn thes is o ffatty ac ids, two
majo rtypes o flipids a refo rmed :
the tr ig lycer ides and the
phospho lip ids . The t riglycer ides F
igure2.5
B
iosyn
thesiso
funsa
tura
tedfa
ttyac
ids
are u ltimate ly s tored in fa t
depos its wh ilethe phospho lipids
are incorporated in the
biomembranes .

Ox
ida
tiono
fFa
ttyAc
ids

Long cha infa tty ac ids , comb ined ast rig lycerides,p rov idethelongte rm
storagefo rm o fene rgyinthefa to r ad iposet issue o fthe an ima l. When
ene rgy demands a re g rea t
,fa tty ac ids a re broken downto y ield ene rgy.
The mos timpor tan t mechan ism by wh ich these s to ragefa tty ac ids a re
degradedin a s tep -w ise manne rto y ie
ld ene rgyis known as β -oxida t
ion .
Accord ing tothe β -oxida t
ion mechan ism , the β -o r 3r dcarbon a tom
fromthe ca rboxy l end o f thefa tty acid cha inisthe s ite a t which oxygen
isin troduced dur ingox ida tion . Ca tabol ism beg ins w i
ththeca rbony lgroup
by conve rtinglong -cha infa tty ac idsto ace tyl CoA , wh ichis subsequen tly
oxidizedinthe c itric ac id o r Krebs cyc le. Inth is p rocess , two -carbon
fragmen ts a re success ive ly removedf rom thefa tty ac idinthefo rm o f
acetyl CoA . Fa tty ac id ox ida tion occu rs ma inlyinthe m itochondr ia and
thisisfo llowed by comp lete ox ida tion o fthe ace ty l CoAinthe c itric acid
cycle. I tis one o fthe pa thways through wh ich ce lls de rive ene rgyfo r
syn thesis of ATP and u ltima tely produce ene rgy.
For examp le, the β -ox ida tion o f palm itic acid, C16 :0 , requ ires seven
(7) cycles o fβ -ox ida tions and g ives riseto e igh t(8) mo lecules o f acety l
CoA wh ich en tersthe c i tric ac id cyc le and p roduces ATP . The ATP y ield
forthe comp lete ox ida tion o f one mo lecu le of palm itic ac idis 106 ATP .
Thel iveristhe ch iefo rgan conce rned w ith β -oxidation o ffa t
ty ac ids.
The ox ida tion o ffa tty ac ids is the ch ief source o f energyin the
catabo l
ism o flip ids. The pa thways o fl ipid syn thes is(anabo lism ) and
catabo l
ism occurs s imu ltaneous ly bu tinthe d ifferen t pa rts ofthe ce ll.
30 CHA
PTER2 NUTR
ITIONINTROP
ICALAQUACULTURE

L
ipidPe
rox
ida
tion

One ofthe cha racteris ticreac t

ion o fl
ipids which are exposedto oxygenis
thefo rma t
ion o fperox ides. Th isreactionis ofgreatp racticalimportance as
itleadstothe de terio ra t
iono ftissuesanda lsocausesspo ilageoffoods. This
isduetothep roduct
iono ffreeradicalssuchas, ROO∙,RO∙ , OH ∙duringpe rox
ide
forma t
ion . Thesef reerad icalsreac t by hydrogenremova l and ava r
iety of
addit
ionreac t
ionstha t damageenzymes ,p rote
ins,vitam ins and otherlipids
making o ther nu t
rien ts unava ilabletof ish
. An tioxidan ts are substances
thatp rotectlipidsf rom pe roxidation. The use oflip
idsinf ish d
ietsrequ i
res
the use o fapp rop r
ia te an tioxidants(see Chap te
r5 )
.

Impo
rtanceo
fFa
ttyAc
idP
rof
ilesinF
ish Nu
tr
it
ion

Bothfishandvege tableo i
ls havebeenfound use fulinf ishfeeds . Thefa t
ty
acid compos it
ion ofcommon ly used o il sourcesinf ish d ietsis shownin
Table 2 .8. Incompa rison w i
th o the rvege tab
leo i
lso rfats,f isho ils conta
in
ag reate rvarie
tyo funsa turatedfa ttyac idsoflonge rca rboncha in(20o r22
carboncha inlength)
, mos to f wh ich be longtothe n -3fam ilyo ffattyacids.
Thelong -chain n-3fattyac idsgene ral
ly make upone -fourthtoone -thi
rdo f
al
lfa ttyac idsinfisho i
ls, whe reas ,longcha infattyac idsin mos tvegetable
oi
ls se ldom exceed 5% andisf requen tlylessthan 1% . The d ietarylipid
requ i
remen toff
ishcan be de rivedf romthe irfattyacid p ro f
iles.

Tab
le2
.8 Essen
tia
lfa
ttyac
idcompos
it
iono
fva
riousl
ipidsou
rces(g
/100gfa
ttyac
id)
L
ipid sou
rce 18
:2n6 18
:3n3 20:5n3 22:6n3

P
lan
tsou
rces
Com o
il 58 1 0 0
Coconu
toi
l 2 0 0 0
Co
ttonseed o
il 53 1 0 0
L
inseed o
il 17 56 0 0
Pa
lm o
il 10 1 0 0
Pa
lm ke
rne
loi
l 2 0 0 0
Rapeseed o
il 15 8 0 0
Peanu
toi
l 30 0 0 0
Soybean o
il 50 10 0 0
Sun
flowe
roi
l 70 1 0 0

Ma
rinean
ima
lsou
rces
Cape
lin o
il 5 0 7 5
Codl
ive
roi
l 5 1 16 14
Cu
ttle
fishl
ive
roi
l 1 2 12 18
He
rring o
il 1 1 8 5
Po
llackl
ive
roi
l 2 0 12 7
Sa
lmon o
il 3 0 10 10
Sa
rdine o
il 3 1 13 10
Sho
rt-neck c
lam o
il 1 1 19 14
Sk
ipjack o
il 5 3 7 12
Squ
idl
ive
roi
l 3 3 12 10

Sou
rce
: Tacon 1987
 CHAPTER 2 L
ipids and Fa
tty Ac
ids 31
Essen
tia
lFa
ttyAc
id Requ
iremen
tso
fFish

Fishrequ i
re n-3 and n-6fa tty acidsinthe i
rd iets
.L inolen
ic ac
id(18 :3n-3)
,
linoleic acid (18 :2n-6 )
,e icosapentaeno ic ac id (EPA , 20:5n-3) and
docosahexaeno ic ac id (DHA , 22 :6n-3)a re needed by f ish. Fa ilure to
providethesefa tty acidsinthe d iet can causeimpa irmen tofg rowth and
prolongedlack o fthesefa t
ty ac idsinthe d ietleadto dea th.
The essentialfatty acid(EFA ) requ i
remen ts have been establishedfo r
mostcu l
turedfishes. Asumma ryo ftheseva luesisp resentedinTab le 2
.9.
Fish species di
fferinthe ir EFA requ irements.

T
able2
.9 Essen
tia
lfa
ttyac
idrequ
iremen
tso
ffishandsh
rimp

F
ish spec
ies Requ
iremen
t Re
ference

As
ian sea bass 0
.5% n
-3 PUFA Bo
rlongan and Pa
razo 1991
0
.5% n
-6 PUFA

Ayu 1% 18
:3n-3 Kanazawa e
tal
.1982
1% 20
:5n-3

Ca
rp 1% 18
:2n
-6 Wa
tanabe e
tal
. 1975
1% 18
:3n
-3 Takeuch
i & Wa
tanabe 1977

Chum sa
lmon 1% 18
:2n-6 Takeuch
ieta
l. 1979
1% 18
:3n-3
0
.5% n-3 HUFA

Coho sa
lmon 1
-2.5% 18
:3n-3 Yu & S
innhube
r 1979

Ee
l 0
.5%18
:2n-6 Takeuch
ieta
l. 1980
0
.5%18
:3n-3

Grouper 1% n-3 HUFA M

illamena and Go
lez 1998

M
ilk
fish 1
-1.5% n-3 PUFA Bo
rlongan 1992

N
ilet
ilap
ia 0
.5% 18
:2n-6 o
r 20
:4n
-6 Takeuch
ieta
l. 1983

Ra
inbowt
rou
t 0
.8%n-3 HUFA Cas
tel
leta
l. 1972
0
.8%n-3 HUFA Wa
tanabe e
tal
. 1974
18
:3n-3 20% o
flip
id Takeuch
i & Wa
tanabe 1977
n-3 HUFA 10% o
flip
id

Red seab
ream 0.5 n-3 HUFA Yone e
tal
. 1978
0
.5% 20
:5n-3

Tu
rbo
t 0
.8%n-3 HUFA Ga
tesoupe e
tal
. 1977

Ye
llowta
il 2% n
-3 HUFA Desh
ima
ru e
tal
. 1984

T
igersh
rimp 2
.6% n
-3 PUFA Ca
tacu
tan 1991
<5% n
-6 PUFA
0
.5% n
-3 PUFA

Ku
ruma sh
rimp 0
.5-1% n-3 PUFA Kanazawa e
tal
. 1980

Rainbowt rout(Onco rhynchus myk iss)requires about 1% 18 :3

n-3inthe
d
iet. Comb in
ing 18 :3n-3and 18 :2n-6inva rious proport
ions did notimprove
growthra te o
re f
ficientfeed conve rsion. In contrast
, carp(Cyprinuscarpio),
one ofthe mos timpo rtantcu lturedf ishinJapanrequ ires bo
th 18:2n-6and
18:3n-3. The bes t weigh t gain andfeed conve rsion are obtainedinf ish
g
iven a dietwith a m ixtureo f 1% 18 :2n-6and 1% 18 :3n-3
. Theee l(Anguil
la
32 CHAPTER 2 Aquacu
lture Nu
tri
tion

japonica), another cu l
tured warmwaterf ish
, has a requ irementfor both
18
:2n-6 and 18:3n-3, but a
t alevelof0 .5%. At rop
ical herb
ivore
,N i
let
ilapia
(Tilapia ni
lotica
) requ ires n-6 rather than n-3fatty acids. The
ir d
ietary
requiremen tfor 18:2n-6 or 20:4n-6is about0.5%inthe d iet
.
H
igh ly unsa tu ra ted n-3fa ttyac ids(n -3 HUFA )a reessen t
ia linthe nu t ri
tion
of some ma r
ine f ish such as red sea b ream(Ch rysophy rs ma jor) , and
yellow ta il(Se riola qu inque rod iata) . The requ i
remen tforlong -cha in
polyunsa turatedfa tty acidsis duetothel imi ted ab il
ity o fsome ma rine
fishes to add more carbon a toms orto remove hydrogenf rom d ietary
precu rso rs. Thus , mos tco ldwa terand ma rinef ishrequ ires n -3 fa t
tyac ids .
Stud ies onthe essen tialfa tty ac idrequ iremen ts o f wa rmwa te rfish and
shrimp spec iesinthe Ph i
lipp ines showtha t some spec iesrequ ire bo th n -3
and n -6fa tty acids wh ile o the rs on ly n-3. Grow thinh ib i
tion a t ce rtain
levelso fn -3and n -6hasa lsobeenobse rvedin wa rmwa te rfishes. Mi
lk fish
(Chanoschanos )cu lturedinseawa te rrequi res n -3 PUFA . Goodg row th and
surv ival we re ob tained us inge ithe rlinolenic(18 :3n -3)o rn -3 HUFA asl ip id
sou rces . As ian sea bass(La tesca l
ca ri
fer)juven ilesrequ i re both n -3 and n -
6 PUFAa t0 .5%inthe d ieto ran n -3/n-6ra t
io o f1 .0.G roupe r(Epinephe lus
coioides)requ ires abou t 1% n -3 HUFA . Injuven ilet ige r sh r
imp( Penaeus
monodon) , abou t2 .6% d ieta ry PUFAenhancesg row th wh ileleve ls of 18 :2n-
6g rea terthan 5% have a nega tivee f
fect ong row th. Thus ,d i
fferentspec ies
requ ire d i
fferent EFAs andthe d i
fferences a re mo re ma rkedin wa rmwa ter
thanin co ldwa te rfishes.

Gu
ide Ques
tions

1
. Def
inetheterm“lip
id”
. Namethetwo ma jorfunc t
ions ofl
ipids
.
2
. Whataretheimpo r
tanttypes o
flipids? Br
ief
ly def
ineeachcomponen
t.
3
. Showthe gene
ralformulaforfatty ac
ids
. Exp la
in brie
fly
.
4
. Whata rethethree majorfamil
ies o
f po
lyunsa
tura
tedfa tty ac
ids
5
. Namethe envi
ronmen talfacto
rstha taf
fec
tthefatty ac
id compos i
tion
o
ffish
.
6
. Whatarethetwo ma
jortypes o
flip
idsfo
rmed a
fte
rbiosyn
thes
is o
ffa
tty
ac
id?
7
. Where a
rethe b
iosyn
thes
izedfa
tty ac
ids o
rtr
iglycer
ides u
ltima
tely
s
tored?
8
. Unde
r what condit
ions o
r nut
rit
ionals tate o
fthe animalisfat
ty ac
id
ox
ida
tionfavored? Under what cond
itionsisitslowed down?
9
. Whatis βeta
-ox
ida
tion o
ffat
ty ac
ids?
10
. Desc
ribethephenomenonofl
ipidperox
ida
tion
. Whyisi
tofg
rea
tprac
tica
l
importance?
11
. Why doesthetype o
flip
idrequ
iredfo
r wa
rm wa
terf
ish d
if
ferf
rom co
ld
wa
terfish?
12
. Wh
ich amongthef
ishspec
iess
tud
iedrequ
ires mo
re n
-6ra
the
rthan n
-3
fa
tty ac
ids?
13
. Di
ffe
rent
iate be
tween
: a) sa
turated vs
. unsaturatedfa
tty ac
ids b
)
po
lyunsa
turatedvs
.high
ly unsa
turatedfat
ty ac
id.
14
. Namesomeo
ilstha
tare usedinf
ishd
ietsandexp
lain whytheya
re u
sefu
l.
CARBOHYDRATES
In
troduc
tion
Carbohydra tes(CHO )a re ala rge g roup o fo rganic compounds common
in p lan ts wh ichinc lude s imp le suga rs,s tarches , cellu loses, gums , and
rela ted subs tances. They con ta in C, H , and O w ith ara tio o f hyd rogen
to oxygen o f2 :1 wh ichis s imila rto H2Ohencethe name“ca rbohyd rate”.
However the ra tio o f2:1 is no ta lways t rue . The genera lformu lais
Cn(H2O )n. Carbohydra tes a re genera lly composed o f ca rbon, hyd rogen ,
and oxygen and some times n itrogen and su lfur . Chem ically,
carbohydra tes on hydro lys
is y ield po lyhydroxy a ldehydes( the a ldoses)
and ke tones( the ketoses). Theyfo rmthela rgest pa r
to fthe an ima l’sfood
supp ly and make up 75 pe rcento fthe d ry weighto fp lan ts.
Carbohydra tes are a cheap source o fene rgy and can spa rethe mo re
expens ive pro te
in as an energy source . The pro tein spar ing e ffects o f
carbohydra tes andl ip
ids shou ld be max im izedin orderto reducefeed
cos ts.S ince ca rbohyd rateisthe mos t econom icald ieta ry ene rgy sou rce,
as much d iges t
ible carbohydra te asthef ish can u ti
lizeis usedinf ish
diets. Carbohydra tes such as s tarch,f lou r
,a lginates , aga r, carrageenan ,
andguargum a re a
lso used asfeed b inde rstoimp rovethe wa te rs tabili
ty
off ish and sh rimp d ie
ts.
Af ter s tudy ingth is section, the reade r shou ld be ab leto d if
feren tiate
amongthe var iousfo rms o f carbohydra te andthe i
rs ign if
icanceinf ish
nut rition;d istinguish be tween u til
ization o fca rbohyd rates by wa rmwa ter
andco ldwate rfishesandknowhowd ie
ta ryca rbohyd ratesa re madeava ilable
tof ish .

C
lass
if
ica
tiono
fCa
rbohyd
rates

Carbohydra tes are c lass ified as

T
able2
.10C
lass
if
ica
tiono
fca
rbohyd
rates
monosacchar ides ,d isacchar ides
Monosaccha
rides(s
ing
le g
lycose un
its
)
or oligosacchar ides , and po lysac ­
Pentoses (C5H1
0O5
) Hexoses(C6H1
2O6
)
char ides depend ing on whe ther Arabinose Glucose
the mo lecu leis made up o f one , Ribose Galactose
two or many s imp le sugar un its Xylose Fructose
Xylulose Mannose
(Tab le 2 .10 ). They are c lass ified
D
isaccha
rides(2 g
lycose un
its
) and o
ligosaccha
rides(2to 10 g
lycose un
its
)
accord ingto d igestibil
ityas d iges t­
ible, par t
ially d igestible, andin­ Sucrose Raff
inose
Mal
tose Stachyose
digestible. Suga rs,s ta rches , dex ­ Lactose Verbascose
trin, and g lycogen a re d iges tible
Po
lysaccha
rides(G
lycan
, >10 g
lycose un
its
)
carbohydra tes whe reas, ce l
lu lose ,
Sta
rch
dietaryf ibers, and hem icellulose Dex trin
are ind iges tible carbohydra tes . Glycogen
Galac togens , mannosans ,inu lin, Cellulose
Hem icellu
lose
and pentosans are cons idered Lign in
partially diges tible carbohyd ra tes . Chitin
Pectin
Gums and muc i
lages
Algina tes, aga
r, ca
rrageenan(ex
trac
tsf
rom seaweeds
)
34 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

A
. Monosaccha
rides
The monosaccha rideisthefundamen tal un itf rom wh ich a l
lca rbohy ­
drates a re de rived . They a re cha racterized bythe numbe ro fca rbon
atomsthey con ta in, C3 , C4 , C5 , e tc., and by the i rs truc tura l
configu ra tion e ithe ras a ldose o r ketose. The ca rbohyd ra teg lucoseis
an a ldose because o fthe p resence o fan a ldehyde g roup wh ilef ruc tose
is a ke tose because o fthe p resence o fa ke tone g roup . Mos t common
suga rs a re a ldoses ra the rthan ke toses . The irs truc tureinc ludes a
carbony lg roup and seve ral hyd roxylg roups .
Mos t monosaccha ridesa reob tainedbyhyd ro lys iso f mo recomp lexp lan t
subs tances . Hydro lysisis a chem ical reac tion whereby a comp lex
subs tanceis b rokenin to sma lle r un its bythe add ition o f wa te rinthe
presence o fa ca ta lyst. The monosaccha rides a re o ftenre fe rredto as
simp lesuga rs. Twose rieso fs imp lesuga rs a recomme rciallyimpo rtan t:
the pen toses o rf ive -carbon -atomsuga rs andthe hexoses o rs ix -ca rbon -
atom suga rs. The hexoses a re abundan tin na ture , bu tthe pen toses ,
ribose and deoxyr ibose , occurinthe s truc tures o f RNA and DNA ,
respec t
ive ly .
Pen toses The pen toses havethe gene ra lfo rmu la C5H1 0O5 . Two
pen toses a re o f comme rc ialimpo rtance and bo th a re a ldopen toses :
xylose and a rab inose . Xy loseisfo rmed
by the hydro lys is o f pentosans .
Cons iderab le amount o f xy lose are
fo rmedinthe pu lp ing o f woodth rough
hydro lysis o f hem icellu lose . Arab inose
is produced by the hydro lys is o f gum
arab ic o r whea tb ran .
Hexoses The hexoses have the
D
-arabinose D-xy lu
lose genera lformu la C6H1 2O6 . The hexose
(ana ldose ) (ake tose) sugars : ga lac tose , and g lucose are
aldoses , wh ilef ruc toseis a ke tose . The
mos t abundan t carbohydra tes in p lan ts and an ima ls are e i ther
hexoses o r comp lex mo lecu les wh ichfo rm hexoses on hyd rolys is .
Glucoseisthe mos t common o fthe a ldohexose sugars andis
comme rc ially producedf romthe hydro lys is o f co rns ta rch . I tisthe
basic mo lecu leforthe syn thes is o fs ta rch and ce llulose . G lucoseis
ofg rea timpo rtancein nu tr
ition , asi tisthe ma jo rend p roduc to fthe
digest ion o fca rbohydra tes by nonrum inan ts . Glucose , ox id ized to
ca rbon d iox ide and wa te r,is
the p rima ry ene rgy sou rce o f
humans .
Fruc tose is the on ly
impor tant ketohexose in
na ture andis the swee test
ofthe ca rbohyd ra tes. When
cane o r bee t suga r(suc rose )
is hydro lyzed , one mo lecu le
off ruc tose and one mo lecu le
ofg lucose a refo rmed .
D-glucose D-galac tose D-
fruc tose Galac tose does no t occur
(analdose ) (ana ldose ) (ake tose) in thef reeform in na ture .
 CHAPTER2 Ca
rbohyd
rates 35
The hydro lysisoflactoseorm ilksuga rp roducesga lactoseandg lucose.
Isome rism .G lucose,f ructose , and ga lactose have the same
molecu lar formu la but the ir struc tural formu las d i
f fer in the
arrangemen to ftheira toms w ith in a mo lecule. These sugars a re
optica lisomers because when p lacedin a po lar
iscope ,g lucose and
galac toserotatethe plane ofpola rizedlighttother ight
, wh ilefructose
rota tes thel ight to theleft
. Thus ,a lthough they have the same
molecu larformu lasthey d i
fferinthe irind ividualp roperties.

B
. D
isaccha
rides and O
ligosaccha
rides
Theo ligome rso fsuga rsare made upo ftwototen monosaccha ride un its
andf requen tlyoccurasd isaccharides. D isaccha ridesa reacomb ination
oftwo mo lecu les o f monosaccha ride . Thefo rmu la , C12H2 2O1 1
,shows
that one mo lecu le of water wasremoved astwo monosaccha rides a re
comb ined . Hyd ro lys
isresu ltsincleavageo fthe mo leculeandfo rma tion
ofthe hexoses . Theth reeimpo rtan td isaccha rides a resuc rose ,lac tose
and ma l
tose .
Suc rose,the commontab le suga r,is made up o facomb ina tion o f
one mo lecu leo fg lucoseandone mo lecu leo ffruc tose.I tisde rivedla rge ly
fromsuga rcaneandsuga rbeets wh icha resou rceso fcomme rcia lsuga r
.
When suc roseisconsumed by an ima ls,i tis hyd rolyzedtog lucose and
fructose , wh ich a re degraded by me tabo l
ic p rocessesto p rov ideene rgy.
Lac tose ,o r m ilk suga r
,isfound on lyinthe m ilk ofa l
l mamma ls.
Nearly ha lfo fthe m i
lkso lidsislac tose hence hyd rolysis oflac tose w ill
give a mo lecule o fglucose and a mo lecu le ofga lactose. Lac toseis o f
specialin te restin human nu tr
ition.
Ma ltose occu rs naturallyin seeds o fs tarch-p roducing p lan ts .I tis
formedbyhydro lyzings tarch wi
ththeenzymeα-amy lase. When ma l
tose
isfur the r hyd rolyzed by the enzyme α-g lucos idase,two mo lecu les o f
glucose a re p roduced .

Suc
rose Lac
tose(afo
rm) Ma
ltose(afo
rm)

C
. Po
lysaccha
rides
The polysaccharides areformed bythecomb inationofhexoseso rother
monosaccha r
ides. Theya reofhigh molecula
r weightand mostlyinsoluble
in wateranda recons ideredthe mos timportantnu t
rientsofplanto r
igin.
Upon hydro lysis by acids or enzymes , they are broken downin to
variousin termed ia
te produc ts andf inallyinto thei
rs imp le sugars
(Figure 2.6
). The po lysaccha
r ides havethegene ralformula(C6H1 0O5)n
.
36 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

F
igu
re2
.6
Summa
ry o
f hyd
rolys
is o
f ca
rbohyd
rates
.

Starchisthe p r
inc ipalstorageform o fcarbohyd ratesin plants andis
foundinthetube r,rh izomes, and seeds.Itisthecheapes tfoodstuffand
serves ma in
lyin human and an ima l nut
rition as a source o f energy
.
Starchis hydro lyzed by e i
ther acids or enzymesin to dextr
in, ma ltose
,
andf ina
lly glucose. Bo th plants and anima ls contain enzymes,a lpha-
and beta- amy lase, that hydrolyze s
tarch. Abou t 20to 26 mo lecules of
glucose are producedf rom each mo lecule ofstarch.

S
tarch(X=abou
t20to26
)

Glycogenis manu factured by mamma ls ando theran imalsf romg lucose

in the b lood andisfoundin the musc le t
issue and l iver. I tis the
sto ragefo rm o fca rbohyd ratein an ima ls, as starchisin p lan ts.
Celluloseisthe ma jors t
ruc tural componen to fp lant cel
l wa ll andis
the mos t abundan t subs tance in the p lantk ingdom . Ce l
lu lose is
essen t
iallyinso luble and ex treme ly resis
tan tto deg radation by enzyme
action . I t cons ists of severalg lucose un i
ts and can be hydro lyzed to
glucose by s trong ac ids. An ima lslack enzymes ca l
led ce llulase tha t
hyd rolyze ce lluloseto g lucose.
Hem icelluloses a recomposed o fa m ixture ofhexose and pen tose un its
.
Hyd rolysiso fhem icelluloseyieldsglucoseandapen tose, usuallyxy lose.It
isthe p r
inc ipal componen to fp lant cell wal
l. Un like cel
lulose,i tisless
resis tantto chem ical degrada t
ion and can be hyd rolyzed by are lat
ively
mild ac idt rea tmen t.
 CHAPTER2 Ca
rbohyd
rates 37
Lign inisfoundinthe cobs and hu lls andthef ib rous po r
tion o froo ts
,
stems ,andleaveso fplan ts.I thasacomp lexs t ructu recons istingo fca rbon
to ca rbon bonds and e the rl inkages wh ichisres istan tto ac id and a lka l
i.
Chitinisthe ma jor s t ruc tural componen to fthe r ig id exoske leton o f
inve rtebra tes such asinsec ts,C rus tacea , and a lso occu rsin ce ll wa lls of
algae ,fung i, andyeas ts.I tisapo lysaccha ride w ith n it rogen a tomsas we l
l
as C ,H ,and Oandis composedo fN -ace tylD -glucosam ine.L ikece l
lu lose,
chitinhasas tructu ralro leandafa iramoun to f mechan ica ls trengthbecause
theind ividua ls trands a re he ldtoge the r by hyd rogen bonds .
Pec tinisfound pr imar ily onthe spaces be tween p lan t cell wa l
ls and
may a lsoin f
iltrate the ce ll wa l
li tself. Pec tin canno t be hydro lyzed by
mamma lian enzyme pec tinase bu tis d iges ted by m icrob ial ac tion. I t
can be ex tracted w ith ho to r cold wa ter andfo rms a ge l.
Plan tgums a reformed a tthes i
te o fin ju ryo rby aninc isiononthe ba rk
.
They a re comp lex,h igh ly b ranchedres idues con tain ing D -glucu ronic and
D-ga lac turon ic acids a long w i
th o the rs imp le suga rs such as a rab inose
and shammose . They a rev iscousf luids wh ich become ha rd when d ry and
are used comme rcially asth icken ing agen ts o rs tab ilize rsfo r emu ls ions.
Gum a rabicis a we l
l-known comme rcia l gum .
Alg ina tes, aga r
, and ca rrageenan a re ex trac tsf rom seaweeds ,the g iant
kelp,G racilaria sp .andKappaphycus sp .,respec t
ively . Thesea reimpo rtant
binde rsinf ish andc rus tacean d iets.

U
ti
liza
tiono
fCa
rbohyd
rates

Aqua tican ima lsdono thaveaspec ificd ietaryrequ iremen tfo rca rbohyd ra tes
bu tthe irp resencein d ie ts may p rovide aninexpens ive sou rce o fene rgy.
The ab i
lity o ff ish to u ti
lize d ie tary carbohydra tes for energy var ies
cons iderab ly. Mos t carn ivorous spec ies have more l im ited ab ility
compared w ith omn ivorous and herb ivorous spec ies. Amy laseis an
enzymetha td iges ts sta rchesin to suga rsinthein tes t
ine o ff ishes . The
activity o fin testina l amy laseis h ighe rin omn ivo rousf ishes ,l ike m ilk f
ish ,
ti
lap ia , and ca rp,thanin ca rn ivo rousf ishes, such asra inbowt rou t and
yellowta il. F ib rous ca rbohyd ra tesinthefo rm o fce l
luloseis essen tia l
ly
ind igest ib le by some f ish spec ies and does no t make a pos itive
con tr
ibu t
ion to the ir nu tr
ition . Thus , theleve lof crudef iberinf ish
feedsistyp icallyres trictedtolessthan 7% o fthe d iettol im i tthe amoun t
of und iges ted ma te r
ialtha t en te rsthe cu lture sys tem .
Theava ilab i
lityo fva r
iousfo rmso fca rbohyd ratestof ishandthe irnu tritive
valuerema ins unc lear. The h igh ly d igest
ibleca rbohyd rates(ca rbohyd rate
withsma ll mo lecu lar weigh tandsho r
te rchainleng thsuchasg lucose ) have
high nu t r
itive va luefo r coldwa terf ishes bu ta re no teffec tive ly u t
ilized
by o therf ishes . Warmwa terf ishes such as red seabream and carp
utilize s tarch more e ffec t
ive ly than dex tr
in and g lucose . F igure 2 .7
showsthe g row th andfeed e fficiency o fca rpfed d iets con tain ing s ta rch,
dex trin o rg lucose a t 42%leve l. Thecha inleng th ofva riousca rbohyd ra tes
andf requency o ffeed ing a f
fec tsthe iru ti
lization byf inge rling ca rp. The
sta rch d ietresu ltsin bes t weigh t ga in andfeed e f
ficiency compa red w ith
othe rca rbohyd rates w i
thsho rte rcha inleng thsuchas dex tr
inandg lucose .
Thetype andleve ls ofca rbohyd rateinthe d iet have a lso been shown
to a ffectthe grow th o f penae id shr imps. T iger shr imp u tilizes s tarch
38 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

better than dextr in and

glucose . Whenthe d ietary
starch leve l is increased
from 20 to 30% , and the
dieta ry proteinis dec reased
from 40to 30% , weigh tga in
andfeed e fficiency a re no t
affected but prote in
eff
ic iencyra tioisinc reased
(Tab le 2.11). O thers tudies
showtha ttige r shr
imp can
utilize treha lose and
suc rosebe tterthang lucose .
Glucose is a lso poor ly
uti
lized by o the r species o f
penae ids such as kuruma
shr imp . Furthermore ,
dif
fe ren ttypes andleve ls of
carbohydrate in the d iet
F
igu
re2
.7 sign if
icant ly in fluences
G
rowth andfeede
ff
ic
ien
cy(inparen
thes
is)o
fcarpfed
d
ietsconta
in
ing42%ofα-s
tarch
,dext
rinorg
luco
se.
surv ivalo fjuven i
le t iger
Sou
rce: W
atanabee
tal
.19
88 shrimp .

Tab
le2
.11 Meansfo
r weigh
tgain
,feedef
fic
iencyrat
io(FER
),prote
ineff
iciencyrat
io(PER),
andsurv
ivalrateo
ftigershr
impfeddie
ts wi
thdif
feren
tcarbohydra
tesou rcesandleve
ls

Carbohyd
rate sou
rce We
igh
t ga
in (%) FER PER Su
rviva
l(%)

40% p
rote
in
20% g
lucose 207
.52d 0
.38d 0
.71c 55
.77c
20% dex
trin 370
.99a
b 0
.47b
c 1
.05b
c 65
.39b
20% s
tarch 408
.17a 0
.50a
b 1
.11b 64
.42b

35% pro
tein
25% g
lucose 232
.38d 0
.35d 0
.91c
d 47
.29°
25% dex
trin 328
.99b
c 0
.44b
c 1
.06b 75
.00a
b
25% s
tarch 388
.71a
b 0
.48a
b 1
.25b 80
.81a

30% p
rote
in
30% g
lucose 152
.44c 0
.26c 0
.76c
d 55
.94°
30% dex
trin 272
.68c
d 0
.40c
d 1
.20b 71
.55a
b
30% s
tarch 387
.36a
b 0
.54a 1
.51a 74
.36a
b

Co
lumn mean
sha
vingacommonsupe
rsc
rip
tareno
tsign
i
fican
t
lyd
i
ffe
ren
t(P>0
.05
).
Sou
rce
:Sh
iau 1
992
 CHAPTER2 Ca
rbohyd
rates 39
Gu
ideQues
tions

1
. Wha
t doesthete
rm “carbohydra
te” mean?
2
. Wha
tarethefunc
tions o
f ca
rbohydra
tes?
3
. D
ist
ingu
ish be
tweenl
ipid and carbohydra
tes
.
4
. Wha
tareca
rbohydra
teschem
ica
lly? Whyisg
lucoseca
lled ana
ldose
,
f
ruc
tose a ke
tose?
5
. G
ivethree c
lass
ifica
tions o
fca
rbohydra
tes and g
ive one examp
le o
f
each
.
6
. Wha
tisisome
rism? Why a
reg
lucose
,fruc
tose
, and ga
lac
tose op
tica
l
isome
rs?
7
. Wha
tisthe ma
jorend produc
tofca
rbohyd
rate d
iges
tion?
8
. Showby meanso
fchem
ica
lequa
tion howg
lucoseis madef
roms
tarch
.
9
. Wha
tca
rbohydra
tetypeisthe pr
inc
ipa
lcomponen
toftheexoske
leton
o
finsec
ts andc
rus
taceans?
10
. Name theimpor
tan
tbindersfor aquacu
lturefeeds ob
tainedf
rom
seaweeds
.
11
.Ifl
ipidg
ivestw
ice as muchene
rgyasca
rbohyd
rates
, whythenshou
ld
ca
rbohydra
tes beinc
ludedinthe d
iet?
12
. Name some good ca
rbohyd
rate sou
rcesinf
ish d
iets
.
ENERGY
In
troduc
tion
Ene rgyis de fined asthe ab ility o rcapac ityto do wo rk. Ene rgy mayex ist
in d ifferen tfo rms and do d ifferen tk inds of wo rk. Aqua tic an imalsrequ i
re
foodto supp lythe ene rgythey need . The ene rgyva lue offoods maycome
from ca rbohyd ra tes,fa ts, and p ro teins and can be measu red direct
ly by
means o fa bomb ca lor
ime ter. Ene rgyisrequ iredto do mechan ical wo rk
(musc le ac tivity ), chem ical work(chem ical processes wh ich take p lace
in the body ), e lectrica l work (nerve act ivity )
, and osmo tic work
(ma in tain ingthe bodyf luids a t equ i
librium w i th each o ther and w ith
the med ium , whe therf resh , brack ish o r sea wa terin wh ichthe an ima l
lives). F ree energyistha t wh ichisle ft ava ilab lefor b iolog ica
l ac t
iv i
ty
and g row th a fte rthe ene rgy requ iremen tis me t
.
The quan t
i ty and cos to f ene rgy wh ichis ava ilableforthe g rowth o f
the spec ies be ing cu l
turedis mos timpor tan tf rom the po into fview o f
the aquacu lturis t. Theene rgyrequ i remen tso fthean ima lvaryinquan ti
ty
accord ing to the spec ies, feed ing hab its , s ize, env ironment and
reproduc tive s ta te. The ene rgy supp l
ied byfoodis one o ftheimpo r
tan t
cons ide ra tionsin de term iningi ts nu tr
itional va lue . Ene rgyis exp ressed
in k iloca lo r
ies(kca l)o rk ilojou le(kJ ). Ak i
loca lo rieisthe amoun tof hea t
necessaryto ra ise thetempera ture o f one k ilogram o f wa ter 1 °C. The
Joule(J )isthe un itofene rgyinthe me tr
ic sys tem and
one kca lis equa lto 4 .186 kJ . Fo r examp le , 70 kca lis
equa lto 293 .02 kJ .
Afte rs tudy ingth is sec t
ion,thereade rshou ld be ab leto
dif
feren t
ia tethefo rms o fene rgy andthe ir measuremen t.
unde rs tand d ieta ryene rgy me tabo lism ,theene rgy ba lance
equa tion and fac tors tha t in fluence d ie tary energy
requ iremen to ff ish ; and unde rstand the s ign i
ficance o f
optima lp roteinto non -p rotein ene rgyinf ish d iets.

U
ti
liza
tiono
fEne
rgy

The gross ene rgy(GE ) value offeedistheto ta

l ene rgy
containedinthefeed . No ta llofthe GEis ava i
lab leto
the an ima l
. D igestible energy (DE )o f afeed is the
di
fference be tween the GE and the energy excre tedin
thefeces. The ene rgy ava i
lablefo rg row thistha t wh ich
rema ins a f
ter the requ irements for me tabo lism,
reproduc t
ion,e tc., have been supp lied(F igure 2.8).
The use o fdigestible energyva luesis a mo re sens ible
and prac tical way o f express ing the energy va lue o f
feedstuf
fs. Theo ret
ically, me tabol
izab le energyvalues a re
a more exac t measure o fthe d ietary energy used for
metabo l
ism by the t issues . However , d irect F
igure 2
.8
measuremen to f metabo l
izable ene rgyinf ishis dif
f icu lt Ut
i
liza
tionofene
rgy
.Theene rgyavai
lab
lefo
rgrowthi
stha
t
wh
ichrema in
saf
tertherequ
irementsfo
r me
tabo
l
ism,
andinvo lvescon finingthef ishin sma l
lvo lumes of wa te r reproduc
tion
,et
c.havebeensuppl
ied.
in me tabolism chambe rs.
42 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

The me tabolic rate or energy expend i

ture of smal
l aqua tic an
ima lsis
greater than tha to flarge an imals. Sma l
l animals growfas ter than
large onesinte rms of percentageincreasein weight per day
. Thus , the
feed requiremen ts o
f sma ll animals a
re higherthanthose o flarge ones,
when expressed as pe rcentage oftheir weights
.

Ene
rgy Me
tabo
lism

Energy is obta ined a fter ma jor nutr ients

carbohydra tes , fa ts, and pro teins undergo
several chem ica l processes such as ca tabo lism
and ox ida tion w i
th in the an ima l body . The
energyl ibera tedis usedforthe ma in tenance
oflife processes such as ce llular me tabo lism ,
grow th, rep roduc t
ion , and phys ica l activ ity.
Thef reeene rgyl ibe ratedf romtheca tabo lism
and ox ida tion o f ma jorfood nu t rientsis no t
uti
lized d irec tly bythe an ima l
, bu tist rapped
inthefo rm o fthe ene rgy -r
ich phospho rus bond
of adenos inet riphospha te or ATP(F igu re 2 .9)
.
Itis ATP wh ichisthe p rincipald rivingfo rcein
the ene rgy requ iring b iochem ical processes o f
li
fesuch as anabo lism o rsyn thesis , mechan ical
F
igure 2
.9
energy ,e lec tr ica l energy , osmo tic work , and
Chem
icals
tru
ctu
reo f ATP
. Adenos
inetr
ipho
spha
tecon
tains
aden
ine(a
),r
ibose(b)
,andt r
iphospha
teuni
t(c
). energy for trans ferr ing subs tances aga ins t
concen tra tion grad ients . The ro le of ATP in
cel
lu lar ene rge ticsis shownin F igure 2 .10 .

F
igu
re2
.10
Thero
leofATPince
l
lula
rene
rge
t
ics
.ATPi
sthep
rin
cipa
ldr
iv
ingfo
rceintheene
rgy
-requ
ir
ingb
iochem
ica
l
pro
cesseso
fli
fe
.

ATPis an ene
rgy-r
ich molecule becauseitstr
iphospha te un
it contains
two phosphoanhydr
ide bonds. Ala rge amounto ffree ene
rgyisl ibe
ra ted
when ATP is hydro lyzed to adenos ine diphosphate (ADP ) and
orthophosphate (P i
) or when ATP is hydro lyzed to adenos ine
monophosphate (AMP) and pyrophospha te(PP
i)
, as shown below:
ATP + H2O — ADP + P i + H+
ATP + H2O — AMP + PP i + H+
 CHAP
TER2 Ene
rgy 43
Undertypical ce
llular cond
itionsthe ΔGforthesereac t
ionsis-12 kcal
per mol
. ATP , ADP, and AMP a reinte
rconver
tible. The numbe rof mo
les
of ATP tha tis liberated in the comp le
te ox idat
ion o f 1 mole o
fa
carbohydratelike g
lucose and 1 mole ofafat
ty acidlike pa
lmita
teto CO2
and H2Oa re 30 ATP and 106 ATP ,respect
ively
.

Ene
rgy Ba
lanceand D
ieta
ry Requ
iremen
t

Since a
llb
iologica
l sys
tems obeythelaws ofthermodynam
ics
,the ene
rgy
balanceequat
ioncan berepresented asfo
llows
:

C=P+R+U+F

whe
re:

C(consump tion) - thegrossene rgycontentofthefoodingested

P(produc tion) - energy u
til
izeding rowth mate
rials
R(respiration) - netlossofene rgyas hea
t
U(urina ryloss) - energylostin nit
rogenousexcretory p
roduct
F(feca
lloss ) - energylostinthefeces

A genera l
ized energy budge tfor young carnivorous and herbivorous
f
ishesfed on na tura lfood has been developed by Brett & Groves. An
ene rgy budgetisthe amoun tofene rgyin percentofingestedfoodtha tis
uti
lizedfor each ma jor process such asfor grow th and reproduction,
digestion, resp
ira
tion, urinary, andfecalp roduction
.

Carn
ivores : 100 C =29P + 44R + 7U +20F
Herb
ivores : 100 C = 20P + 37R +2U +41F

*B
ret
tand G
roves1979

whe rethef igures are expressed as pe rcentage o finges tedfood ene rgy o r
percen tageo fenergyde rivedfromea tenfood . Me tabolizab leenergyva lues
forf ishin k iloca
lories pe rgram o f substance a re:4 .5,3 .3
, and 8 .0for
prote in, carbohydra te and fa t, respec tive ly. The energy needs for
main tenance and vo lun tary swimm ing ac tivity mus tfi rst be sat
is f
ied
before ene rgy can be made ava ilablefo rg row th.
Fish ,likeotheranima ls,eatp rima r
ilytosa tisfythe i
rene rgyrequiremen ts.
Whenthefeed haslowene rgydens i
ty,fisha reab letocompensa tebyea t
ing
moreo fthefeedbu tw ithince r
tainl imits. The refore,itisessen t
ialthatfish
have easy accesstofeed o ra re given a pa la table ration w ith su
fficient
ene rgyto mee ttheir ene rgy needs.

D
ieta
ryEne
rgy Requ
iremen
t

Fish d
ietsshould have anop t
imal proport
ion ofene rgyfrom dietarylipids
,
carbohydrates, and prote
ins . An excess or a de ficiency o
fl ipids and
carbohydrates
, which are nonprote
in ene rgysources, mayresu ltinlowe r
growth. Ifthe d ietis deficien
tin nonpro tein energy, protein whose
functionisforgrow th andrepa i
ro ftissues wi
ll be used asene rgy source
44 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

ratherthanfo rg row th. Simi

la r
ly,i
fthe d ie
t contains an excess ofl
ipids
and carbohyd rates,f ish appe
titeis met even before enough p ro
tein(and
possibly o
ther nu tr
ients)is eaten to mee tthe demand for max imum
protein synthes is and growth . Many fac tors in fluence the energy
requiremen ts off ish:

1. wa ter tempera ture - me tabolic ra te and consequen t

ly energy
requ iremen tsfo r maintenanceinc rease w i
thtempe rature.
2
. an ima lsize- me tabolicrate and consequen tly ma intenance ene rgy
requ iremen ts, dec rease wi
thinc reasing an ima lsize.
3
. phys io logicals tatus - energy requ irements increase dur ing
produc tion o fgonads andreproduc tion ac tivity such as m igration
dur ing spawn ing.
4
. wa terf low ra te - energy requ i
remen tsfor ma in taining a certain
positionin wa terinc rease with aninc reasein wa terf low rate.
5
.l igh texposu re-ene rgyrequ iremen tsfo rvo lun taryac tiv
ity decrease
duringres t periods atn ight
.
6
. wa ter qua lity and s tress - po l
lu tan ts, increased sa l
inity
, low
disso lved oxygen , crowd ing or overs tock ing increase energy
requ iremen tsfor ma intenance.

Gu
ide Ques
tions

1
. Wha
tis ak
iloca
lor
ie? G
ivetheequ
iva
len
tofk
iloca
lor
iein k
ilo
jou
les
.
2
. Wha
tisthe d
iffe
rencebe
tweeng
rossene
rgyinfeedandene
rgyexc
reted
inthefeces?
3
. G
iveth
reeexamp
les o
fene
rgy
-requ
iring b
iolog
ica
lprocesses
.
4
. D
iffe
ren
tia
te be
tween d
iges
tib
leene
rgy and me
tabo
lizab
leene
rgy
. Why
is me
tabo
lizab
leene
rgy mo
re d
iff
icu
ltto measu
re?
5
. Wha
tis ATP? Wha
tisi
tsfunc
tion?
6
. Wr
ite andexp
lain br
ief
ly theene
rgy ba
lanceequa
tion
.
7
. Wha
tarethefac
torstha
tin
fluencetheene
rgyrequ
iremen
ts o
faqua
tic
an
ima
ls?
8
. Whyisi
timpo
rtan
tto p
rov
ide an op
tima
lpropo
rtion o
fpro
teinto non
pro
tein ene
rgyinf
ish d
iets?
V
ITAM
INS
In
troduc
tion
Vitam ins a re organ ic compounds tha ta re requ iredinthe d ietin sma ll
amoun ts and un like p roteins,l ipids, and ca rbohyd rates, have unre lated
chem ica ls truc ture . Th is d istingu ishes v i tam insf rom o ther organ ic
mac ronu trien ts(p roteins, ca rbohyd ra tes,l ipids ). The chem ica ls tructu re
ofv itam insisremarkab ly diverse andthey a re usua lly ofsma ll mo lecula r
size (mo lecu lar we igh t usua lly <1000 )
. They have spec i
fic regu latory
func t
ions and a re necessaryfo r no rma lg row th , ma intenance o f hea lth,
and reproduc tion . Insu ff
icient amoun ts in f ish d iets may resu ltin
nu tr
i t
iona l de ficiency d iseases , poo rg row th, andinc reased suscep tibil
ity
to d iseases andin fections. They do no tg ive ene rgy bu ta re neededin
enzyme sys tems to has ten or ca talyze reac tionsin energy u ti
lization .
Vitam ins a re essen t ialfo rthe regu lation o f me tabo lisminthe ce lls and
forthet rans forma tion o f ene rgy. W ithou tv itam ins , resp iration,g row th,
musc le con trac t
ion , and o ther phys ica l activ ities canno t occu r
. Many
are usedtoform coenzymes or as par to fan enzyme sys tem . Abou t 15
vitam ins are knownto be needed byf ish.
This sec tion descr ibes the var iousl ipid-so lub le and wa ter-solub le
vitam ins , the ird i
fferences , phys iolog icalfunc t
ions , and the symp toms
ofv itam in de ficienc iesinf ish. I ta lso shows a summa ry of nu tr
itiona l
deficiencys igns andtherequ i
remen ts o fva riousf ishspec iesfo rvitam ins .

C
lass
if
ica
tiono
fVi
tam
ins

Vitamins are c lassif

ied accord ingtothe ir solub il
ity: wa ter-soluble and
lipid-soluble. The wa ter-solub le vitam ins, wh ich are the B comp lex,
and asco rbic acid orv i
tam in C and some co factors,d issolvein wa ter bu
t
no tin oi
ls andfa ts. Onthe con t rary,thel ipid-solub lev itam ins,A, D,E
and K d issolvein o i
ls andfa ts
.
Manyofthe wa ter-solublevitam insfunc tione itherd irec t
lyo rina mod i
fied
fo rm as acoenzyme . Coenzymes a reorganic mo lecu lestha t he
lpenzymes
become b iologically active
. In ce r
tainins tances , th isfunc t
ional role o
f
vitamins has been used as a meansto assessthe nu trit
iona lstatus ofan
an imalw i
threspec ttotha tvitamin . Wa ter-solublev itam insa reexcretedin
the ur ine, wh i
lefa t-soluble vitam ins are morel ike lyto appearinthe
bi
le andthus exc retedinthefeces .

□ Wa
ter
-so
lub
lev
itam
ins
Amongthe group o f water -soluble v
itamins areincludedthe B complex
v
itamins, th
iam inor B1,riboflavinor B2,pantothen
icac ido
r B5,py
ridox
ine
or B6
,andcyanocoba lam ino r B12,b
iotin
,n iac
in,choline
,inosi
tol
,fol
icacid
,
and asco
rbic ac
id o
rv i
tam in C.

Th
iam
in (V
itam
in B1
)
. Th
iam
inis a comp
lex subs
tancetha
t con
tains
su
lfur, wh
ich g
ivesi
t a charac
teris
tic pungen
t odor
.I tplays an ac
tivero
le
in carbohydra
te metabolism asthe coenzyme , thiam
in py rophosphate
.
Of part
icularimportance is the ro
le ofthiam inin the conversion of
46 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

pyruvate to ace tyl CoA . Th iamin is

destroyed by hea t unde rsligh tly a
lkaline
and mo ist cond itions . However , the
vi
tam in is re latively s table to dry hea t,
andis re tainedinfeed pe l
le ts duringthe
pellet
ing process and s torage . Wet or
moist f ish d iets shou ld be used
immed iatelyto preven tloss o fth iamin by
hydrolysis.
Th iamin is essent ial for growth ,
reproduc tion and norma ld iges t
ion. Lack
F
igure 2
.11
of thiamin in f ish resu l ts in loss o f
Thiam in(vitamin B1)def
ic
iencyinhumpbackgroupe
r.
Me chanicalin
juriesw i
thhemo r
rhagesonthebodysurfa
ce, appe t
ite, poor grow th, musc le at
rophy ,
pecto ra
lfinsandabdomen ,ande r
rat
icswimm
ingbeha vio
r instab ility and sens itivi ty to shock ,
ares ignso fvi
tam in B1
-def
ic
ientd
iet
Sou rce: Ko esharyan
ie ta
l.2001 hemorrhages and errat ic sw im ming
behavior(F igure 2.11 )
.

Riboflavin(V i
tam in B2 )
.R iboflavinis an orange pigmen ted mo lecule.I t
isinvo lvedin energy me tabolismintheform o fthe coenzymesf lavin
mononuc leo
tide (FMN ) andf lav in adenine d inucleotide (FAD ). These
coenzymesfac i
litatethe breakdown o fenergy-yie
lding nut r
ien ts such as
am ino acids
,fa tty ac
ids, and py ruvic ac
id. R ibof
lavinisre latively mo re
stable to heatthan th iam in bu tis destroyed byl ight andin s t
rong ly
alkal
ine o r ac
idic solutions. Ing redients and rations shou ld be s tored
in dark bags or non -
transparen ttight containers and pro tectedf rom
 CHAPTER 2 V
itam
ins 47
l
ightto preventloss ofribo
flavin ac
tiv
ityinthefeed.
R
iboflav
in defic
iencyinf ish resultsinformat
ion o flens catarac
ts,
photophob
ia, hemorrhagiceyes
, anemia,losso
fappeti
te, and poo
rg row
th.

Pan to thenic Ac id (V i tam in B5 ). Panto thenic acidis an organ ic acid

tha t con ta
ins n itrogen.I tp lays an essen tia
l rolein p rotein,lipid, and
carbohydra te me tabo l
ism as a componen to fthe coenzymes , ace ty
l
coenzyme A , and acy l car rierp rotein. These coenzymes a reinvolvedin
the syn thesis offa t
ty ac ids, cholesterol,s te
roid ho rmones, phospho lip
ids,
and hemog lobin. The sod ium or ca lcium sa lto f panto thenic acidis
rela tively stable and can beincorpora tedin to either dry or mo istfish
diets. I tis s tablein neu t
ral solution bu tis des troyed by hea ta teither
alka line or ac id pH . In d iet prepara t
ion , somelossisincur red dur ing
autoc laving and excess ive hea t.
Pan tothen ic acid de f
ic iency resu ltsin c lubbed g i
lls
,loss o f appet
ite,
nec ros is
, cellular atrophy ,slugg ishness , and poo rg rowth.

N
iac
in (N
ico
tin
ic ac
id)
.Niac
in o
rnico
tin
ic ac
id p
lays an essen
tia
lro
le
inthe me tabolism o fca rbohyd rates,lipids, and prote
ins, as acomponen t
oftwo h igh ene rgy mo lecules, n icot
inam ide adenine d
inucleot
ide(NAD )
and nico t
inam ide aden ine d inuc leot
ide phospha te(NADP ). NADH and
NADPH a reimpo rtantin a numbe ro f oxidat
ion and reduc t
ion reactions
that occur w ithin cells. These coenzymes a re essent
ialforthere lease o
f
energyf rom food nu trients and are a lso
involved in the syn thes is o ffa tty acids
and cho lesterol, respec tively.
Niacin de f
ic iency resu lts in loss o f
appetite
, poor g row th,les ions and edema
of the co lon,jerky mo tion , weakness ,
muscle spasms , andf in erosion .

Pyr
idox
ine (V
itam
in B6
)
. Pyr
idox
ine is
essent
ial in pro
tein me
tabo
lism as the
48 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

coenzyme , py r
idoxa l phospha te, whichisinvo lvedinthe non -oxidative
degrada t
ion o f am ino acids,inc ludingtransam ination, deamina t
ion and
decarboxy lation . The ava ilabili
ty o fv itamin B6 in foods can be
subs tantial
lyreduced by p rocessing. Itis susceptibleto destruction by
lightin neu tral and alkaline solutions hence die
ting redients shou ld be
protectedf rom exposureto sun l
igh t
.
Pyridox ine de f
iciency resu ltsin nervous d isorders, hyperirri
tab i
lity,
edema o fthe per itoneal cavity
, anem ia, rapid and gasp ing brea thing,
andloss o f appetite.

B
iot
in.B
iot
in p
laysanimpo
rtan
tro
leinthe me
tabo
lism o
fca
rbohyd
rates
,
lipids and proteins, as a coenzymeforreac t
ionsinvolvingtransfer and
remova lofca rbon dioxidefrom one compoundto ano ther. As such,itis
essentialforthe syn thes
is offatty ac
ids and catabo
lism ofcer
tain am ino
acids. Itis alsoimpo rtantin cellimmun ity
, as an act
ivatorof bacte
ria-
destroying enzyme ,lysozyme .
 CHAPTER2 V
itam
ins

Bio
tin de f
iciencyleads to reduced growth and ac
tiv
ity
,loss
ofappe t
ite, sk
in diso
rders
, musc le a
trophy
,lesionsinthecolon,
spastic convu ls
ions, skin lesions, and abnormal swimm ing
behavior.

Coba lam in (V itam in B1 2

). Coba lam in or B1 2is essentialfo r
norma l ma tura tion and deve lopmen t
, intheforma tion o fred
blood cells and ma intenance ofthe ne rvetissue asthecoenzyme
cobam ide. The v i
tam inis des troyed by hea tingin a h ighly
alkaline med ium andinthe p resence of ascorbic acid.S tabi
lity
ofv i
tam in B1 2is highinfeeds s tored at mode ratetempe ratures.
In salmon ids ,v itamin B1 2deficiency resu lts in high var iabil
ity of
fragmen ted erythrocytes andin hemog lobinva lues,resemb ling monocy t
ic,
hypochrom ic anem ia. Poor appe t
ite and poor grow th have a lso been
observed in coba lam in def
ic iencyin f ish. The requ iremen tfor and
metabo l
ism o fv itamin B1 2infish has no t been adequa tely studied
.
50 CHAPTER2 NUTR
ITIONINTROP
ICALAQUACULTURE

Folic Ac id (Fo lacin,P teroylglutam ic acid). Fo

lic acidisimpo rtantin
protein me tabolism as acomponen tofthe coenzymete trahydrofol
ic acid
.
This coenzyme is needed for the syn thesis of hemog lob
in,g lyc
ine,
methion ine , cho l
ine, and pur ines . H igh tempera ture or pro longed
heating as we ll as ac
id pH w ill des
troyfo lateinfeeds .
Macrocy tic anem ia occursin the red b lood cell
-produc ing tissue of
the an terior kidneyinf ish. Other signs tha t have been observed a re
poor g row th,le thargy
,b lindness, me lanism and dark sk in colorat
ion,
andf ragilef ins.

fo
licac
id te
trahyd
rofo
licac
id

Asco
rbic Ac
id(V
itam
in C
). Ascorb
ic ac
idis a wh
ite c
rys
tal
line powde
r
tha tis essen t
ia lin ma in tain ingthein tegr
ity o fconnec tivet issues ,b lood
vesse ls , bone t issue , wound t issue and as a co fac torfor numerous
hydroxy la t
ion reac tions . The v itam in also ac ts as a s t rong b iologica l
reduc ingagen tandisrequ iredfo rtheconve rsion o ffolictote trahyd rofolic
acid ,t ryp tophanto se ro tonin , andinthe syn thes is o fs te roid ho rmones .
Asco rbic ac id is eas ily des troyed by hea t and pro longed exposureto a ir
and a lka line med ium . Ox ida tion o fvitam in C or des truc tionin the
process o ffeed prepara t
ionfo rf ish can be reduced by us ing coa ted o r
pro tec ted forms o f L-ascorb ic ac id . S tab le,b io log ica lly-equ iva lent
der iva t ives are ascorba te-2-monophospha te (AMP ) and ascorba te-2-
polyphospha te(APP ). Asco rba te -2 -sulfate(C2 )is a s tab le s toragefo rm o f
Asco
rbicac
id vitam in Cfoundintheth ick de rma llaye ro ff isht issue .
Scurvyisaspec i
ficsynd rome
caused by v itam in C de ficiency
in humans and o the r an imals.
Itis charac ter ized byimpa ired
collagen format ion and
widespread cap il
lary
hemo rrhag ing . Inf ish ,vitam in
C de ficiency causeslordos is,
sco liosis(F igu re 2 .12 )impa i
red
collagenfo rma tion, changesin
the ca rt
ilage , exop tha lmic eye ,
eye opac ity, hemo r
rhag ic sk in,
Figure 2
.12
Ascorbic ac
id(v i
tamin C
) def
iciency
. Deformed spinalcord liv er, ki d ne y, int estine , a nd
of humpback g rouperas aresul
to fvi
tamin C def
iciency musc le.
Sou
rce
: Koesha
ryan
ieta
l.2001
 CHAPTER2 V
itam
ins 5
1
Inos
ito
l. Myo
inos
ito
lis an essen
tia
l componen
tofthe
inosi tol conta ining phospho l
ipids and is animpor tan t
structura l componen to fskeletal, hear t
, and bra int issues .
It ma intainsthein tegri
ty ofce l
l memb ranes,p reven tsthe
accumu lation o f cholesterolinfa t
tyl iver d
isease , andis
involved , with cho l
ine ,in norma ll ipid me tabolism .I ta lso
plays animpo rtantro lein grow th o fliver ce
lls, cho lesterol
transpor t, andin the syn thesis ofr ibonuc leic ac id . The myo
inos
ito
l
compoundiss table and w ithstands no rmalfeed p rocess ing
and s torage cond itions.
Deficiencysymp tomsa repoorg row th,d is
tendeds tomach ,
skinles ions , andinc reased gastric emp ty
ingt ime .

Choline. Cholineisimpo rtant as componen tofacety

lcho l
ine
andthe phospho lipids, leci
th in, and sph ingomyelin. As
such ,i tp lays a v ital role in the ma intenance o f cell
structure andt ransm ission o f nerveimpu lses. Itis a lso
impor tan tinl ip id transportw ithin the body . Cho l
ineis
widely distr
ibu ted infoods .Itis usua llyincorpora tedin
fish diets as cho line hydroch loride . cho
line
De f
iciency s igns inc lude impa ired fat me tabo l
ism,
fat ce l
l necros is syndro me , poor growth , and
hemorrhag ic k idneys and intest ine .

□L
ipid
-so
lub
lev
itam
ins

Re
tino
l(V
itam
in A
).V
itam
in Ais a gener
ic term o
fal
l
compounds o therthan ca rotenoidstha t hasre tinol activity.
Itis an essent ial componen to fv isua lp igmen ts and is
requiredfo rthe ma in tenance o fep ithe l
ial cells. V itam in A
is also requ iredforthe re lease of pro teolyt
ic enzymesf rom
lysosomes . De f
iciency o fv i
tam in Ainf ishleadstosymp toms
such as exoph thalm ia, eyeles ion, anorex ia, asc ites,lens
deforma tion and opercu lum de forma tion (F igure 2 .13)in
fish. Hyperv itaminos is or excess v itam in A has been
describedinf ish andinvo lves enlargemen to fthel iver and
spleen , abnorma l growth , sk in les ions , ep ithe l
ia l
keratiniza t
ion , abnorma l boneforma tion, and hyperp las ia
of head ca rt
ilage.
Codl ivero i
l and o the rlive roi
lscon tainv itamin A( ret
ino lo r F
igure 2
.13
Ope
rcu
lumde
form
ityinhumpbackgroupe
risgene
ral
ly
ret
inyl es ters
). Syn thetic vitam in A( re t
inyl pa lmitate)is o ften cau
sedbyadef
ic
iencyinVi
tamin A
.
usedto supp lemen tra tionslowinf ish mea ls o r carotenes,a Sou
rce
: Koesha
ryan
ieta
l.2001
precurso rofv itam inA. Ca rotenoidsa refoundinphy toplank ton
and a re changedtov itam in Ainthel ivero ffish.
52 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

re
tino
l,v
itam
in A β
-ca
rotene

Cho
leca
lci
fero
l(V
itam
in D
). There aretwo ac
tivefo
rms o
fvi
tam
in D
,
ergocalci
fero l(vitam in D2 ) and cho lecalci
fe rolo rv i
tam in. D.3V itam in D
plays animpo rtantro leinthe p roper use o fca lcium and phospho rusto
form bones and tee th. I t promotes norma l bone format ion or
mineral
ization . Cho leca lci
fe rolcan besyn thesizedinthesk in bysun l
igh t
irradiat
ion o f7 -dehyd rocho les
terol whichisfound unde rthe sk in. I tis
some times ca lledthe sunsh ine v
itam in
. S incethe v itam inisfa t soluble
and accumu la
tesinl ipids tores,fishliveroilis ar ich sou rce ofv i
tam in D.
The deficiency symp toms o frickets and abno rmal bonefo rma t
ion has
been desc ribedinf ishfed alow v itamin D d ietin wa ter tha t con tains
low amoun ts ofcalcium . Poo rgrowth, andte tany o f whiteskeletal musc le
have also beenrepo rted.F ish w i
th hype rvitam inosis D exh ibi
timpa ired
growth , lethargy , and dark
co loration . H igh doses o f the
vitam in mob ilizes calcium and
phosphate and may lead to
frag i
le bones .

α-tocophero
l (V
itam
in E
).
Tocophero lis animpor tantfa t
-
solub le an t
iox idant w ithin the
anima l body , pro tecting react
ive
compounds such as h ighly
unsaturated fat ty ac ids, and
vi
tam ins A and Cf rom ox idat
ive
damage byt rapp ingf reerad ica
ls.
cho
leca
lci
fero
l,v
itam
in D
Thees ter, α-tocophe ro lacetateo r
phospha te
,iscommon ly used as
ad ie
t supp lemen t becausei tis
more s tablethanthef reeform ,
wh ich is rap id ly lost by a ir
oxida tion orin the presence o f
uns tab le react ive me taboli
tesin
fish o ils. Vitamin E has
phys io logic an tioxidan t activ
ity
in grow ing an ima lsincluding
fish . An interact ion ex ist
between v itamin E and
selen ium, a metabolic
ant iox idant ; hence v itamin E
requ irements are greater in
α
-tocophe
rol
,the mos
tac
tivefo
rm
o
fvi
taminE selen ium-dep letedf ish.
 CHAPTER 2 V
itam
ins 53
Fishfed vitamin E -def
icien
td iet containing
rancidfa t exhib
itlow surv ival
, poor g rowth,
fragile erythrocytes and fragmentat ion,
emac ia
tionand darken ingo fbodyco lor(Figure
2.14)
. An excess(hype rv
itaminosis)ofvitam in
E can cause poorg row th,tox
icl iverreaction,
and eventua l
ly, death.

Menad
ione (V
itam
in K
). Vitamin K is
essentialforthe ma in tenance o f norma lb lood F
igure 2
.14
P
ictureofhumpba ckgroupe
r
coagulation byfac ilitatingthe produc tion o f
showingemacia
tion,darken
ingof
various plasma pro teins. I t may a lso p lay bodycolo
r,pe
techiaattheba se
animpor tant partin e lectron transpor t and o
fope rcu
luma sare sul
tofranc
id
d
ietaryfa
tandvi
tam in E
oxidative phosphory lation . Synthe tic def
ic
iency.
menadioneis agood supp lemen tfor adequa te Sou
rce
: Koesha
ryan
ieta
l.2001

vi
tamin K intake . V i
tamin Kis lab ile to
oxidation and exposure to u l
trav iolet
radiat
ion . Diets con tain ing vitam in K mus t
be kep tdry, protectedf rom a ir oxida tion o r
oxidat
ion by u ltraviolet rad ia
tion.
Vitamin K de ficiency s igns inc lude
prolonged b lood c lot t
ing , anem ia, and
hemor rhagic g
il
ls, eyes , andvascu lart issues .
menad
ione
,vi
tam
in K
V
itam
in Requ
iremen
tso
fFish

Vitam in requ iremen ts off ish have been s tud ied by us ing vitam in tes t
diets. The qua li
ta tive requ iremen tis de te rm ined byfeed ing one g roup o f
fish a d iet comp le tein v itam ins, and ano the rg roup ,ad ietf rom wh ich
thev itam in be ings tudied has been de le ted. Grow th andtyp ical de fic
iency
signs a re no ted. I fthe v itam inisfoundto be essen t
ial,the quan titative
requ iremen tfo rtha tv itam inisthen de term ined. The common app roach
istofeed g roups o ffish,tes td ietscon ta iningg radedleve ls o
fonev itam in
wh i
le allo ther v i
tam ins and componen ts o fthe d ie tare kep t cons tan t
.
The “brokenl ine” ana lysis o fthe grow th da tais usedto de term inethe
min imum d ietary requ iremen to ffish . Qua litative and quan t
i tat
ive
expe rimen tsonv itam inrequ iremen ts haveshowntha tthefourfa t-solub le
and 11 wa ter-so lub le vitam ins prev ious ly men tioned are requ ired by
fish.
Therequ iremen tfor av itam in may be a ffected byva riousfac tors such
as spec ies, s ize o r age,d ie tary nu trien tleve ls,d ietcompos i
tion, phys ical
prope rt
ies o fd iet, and cu lturecond i
tions . Fo rins tance ,therequ iremen t
fo rvitam in E mayinc rease asthe po lyunsa tura tedfa tty acidleve linthe
dietinc reases . Inlow -dens ityex tens ivecu l turein ponds o rlakes , natu ral
food a ref requen tly abundan t enoughto prov ide essen tialvitam ins . In
high dens ityin tens ive cu lturein ponds , cages , and raceways , na tura l
foodisl im ited, thus v itam ins mus t be supp liedin the d ietto ach ieve
norma lg row th o ff ish.
54 CHAPTER2 NUTR
ITIONINTROP
ICALAQUACULTURE

The v itam in requ iremen ts by d ifferent spec ies vary grea tly accord ing
tothe i r usua lfeed ing hab it and capac ityto syn thesizethem . A summa ry
ofthev itam in de ficiency symp tomsis g ivenin Tab le 2.12. Some v i
tam ins
canno t be syn thes ized by mos t an ima ls hence there is an abso lute
requ iremen tforthem to preven t the occur rence o f specific def
iciency
diseases .
Ce rtain vitam ins may be des t
royed du ringfeed manu facture by hea t,
moisture,a l
te rationsin pH ,the p resence o fsome me tals
,lipid oxidation,
etc. Des truct ion o fvitam in C due to ox idationis a prob lem infeed
manu factu re. Somev itam ins area lsolos t durings to rage,thusfeedshou ld
be used soon a fter pelleting. In crus taceanfeed ,a l
lowance shou ld be
madefo rtheleach ing o fvitam insf romfeed pe llets. On ly afew stud ies
have been done onthe v itam in requ iremen to f aqua t
ic an imals
. The
supp lemen ta tionleve ls o f each v i
tam ininf ishfeeds a re o f
ten higher
thanthe requ iredleve lsto p rov
ide a sa fety ma rgin.

Tab .12 V
le2 itam
in de
fic
iencysymp
tomsinf
ishes
V
itam
ins De
fic
iency symp
toms

V
itam
in B1 Equ
il
ibr
iumloss
,ir
ritab
il
ity
,le
tha
rgy
, ne
rve d
iso
rde
r, ano
rex
ia, musc
le a
trophy
,
ca
tarac
t, convu
lsions
, sk
in d
isco
lora
tion
, edema

V
itam
in B2 F
in e
ros
ion
,frag
ile sk
in, eyeles
ion
, pho
tophob
ia,i
ris p
igmen
tat
ion
, co
rnea
vascu
lar
iza
tion
, anem
ia, ano
rex
ia,a
tax
ia, ca
tarac
t,c
loudylens
, da
rk sk
in
co
lora
tion
, poo
rfeed e
ffic
iency

V
itam
in B5 Exuda
ted g
il
ls,f
rag
ile sk
in, le
tha
rgy
,liver nec
ros
is, pros
tra
tion
, er
rat
ic
sw
imm
ing
,atax
ia,g
il
lat
rophy
,clubbed g
il
ls, da
rk sk
in co
lora
tion
, de
rma
tit
is

V
itam
in B6 Equ
il
ibr
iumloss
, exoph
tha
lmus
,ir
ritab
il
ity
, ne
rve d
iso
rde
r,rap
idr
igo
r mo
rtis
,
b
lue s
lime
,er
rat
ic sw
imm
ing
, anem
ia,a
tax
ia, convu
lsions
, edema

V
itam
in B1
2 Anem
ia, ano
rex
ia

V
itam
in C Exoph
tha
lmus
,frag
ile g
il
l,k
idney
,live
randsk
in, eyeles
ion
,le
tha
rgy
,lo
rdos
is,
p
ros
tra
tion
, sco
lios
is, anem
ia, ano
rex
ia, asc
ites
, car
ti
lage abno
rma
li
ty,low
d
iseaseres
istance

Inos
ito
l Fa
tty l
ive
r, sk
in les
ion
, anem
ia, ano
rex
ia,d
istended s
tomach
, poo
rfeed
e
ffic
iency

B
iot
in Fa
ttyl
ive
r,f
rag
ile e
ryth
rocy
tes
, co
lon and sk
inles
ions
,blue s
lime
, ano
rex
ia,
musc
le a
trophy
,da
rksk
inco
lora
tion
,convu
lsions
,gi
lldegene
rat
ion
, poo
rfeed
e
ffic
iency

Cho
line Fa
ttyl
ive
r,f
rag
ile k
idney
, poo
rfeed e
ffic
iency

N
ico
tin
ic ac
id Anem
ia, ano
rex
ia, edema
,frag
ile sk
in, co
lonles
ion
,le
tha
rgy
, pho
tophob
ia,
musc
le spasm
, wh
ite musc
lete
tany
, poo
rfeed e
ffic
iency

Fo
lic ac
id Poo
rfeed e
ffic
iency
,frag
ilef
ins
,le
tha
rgy
,anem
ia,ano
rex
ia, da
rksk
inco
lora
tion

V
itam
in A Eros
ion o
ffins
, exoph
tha
lmus
,frag
ile k
idney and sk
in, eyeles
ion
, ano
rex
ia,
asc
ites
,lens de
forma
tion
, edema

V
itam
in D Sco
lios
is, wh
ite musc
lete
tany

V
itam
in E Exoph
tha
lmus
,fa
ttyl
ive
r,f
rag
ile e
ryth
rocy
tes
, anem
ia, asc
ites
, ce
roidl
ive
r,
muscu
lar dys
trophy
, edema
, ep
ica
rdi
tis

V
itam
in K Anem
ia,s
low c
lot
ting b
lood

Sou
rce: Ha
lve
r1989
 CHAPTER2 V
itam
ins 55
The speci
fic v
itamin requi
remen
ts o
fsomef
ish spec
iestha
t have been
s
tud ied a
rein Tab le 2
.13.

Tab
le 2
.13 Summa
ryo
fthev
itam
inrequ
iremen
tso
fva
riousspec
ieso
ffishandsh
rimp
At
lant
ic Channe
l Common Paci
fic Ra
inbow Ye
llow
Ayu Ee
l Sea bass T
ilap
ia Sh
rimp
salmon ca
tf
ish ca
rp sa
lmon trou
t tai
l
Th
iam
in 1
2 1 R R R 10-15 40 11
.2 60
R
ibo
flav
in R 40 9 7
-14 R R 20
-25 9 11
.0 25
Py
ridox
ine 5 1
2 3 5
-6 R 5 15
-20 9 11
.7
Pan
tothen
ic ac
id R 50 1
5 30
-50 R R 40
-50 40 35
.9 75
Coba
lam
in R R 0
.15-0
.02 0
.21 NR 0
.2
N
icot
inicac
id 100 1
4 28 R 150
-200 300 12
.0 40
B
iot
in 0
.3 R 1 R 1
-1.5 0
.4 0
.67
Inos
ito
l 400 NR 440 R R 300
-400 510 423 400
Cho
l
ine 350 400 4000 R 600
-800 11100 2920 600
Fo
l
icac
id 3 R R 6
-10 2
1 1
.2 1
0
Asco
rbic ac
id 50 300 60 R R 700 100
-150 400 R 122 200
V
itam
inA 10000IU 1000
-2000IU 10000IU 2000
-2500IU 7000IU 5
.68 5000 IU
V
itam
in D 2000IU 250
-500IU 3000IU NR 0
.1
V
itam
in E 35 100IU 50IU 1000IU 200 R 30IU 200 50
-100 119
.0 100
V
itam
inK R 10IU R 50 NR 5

Va
luesare mg
/kgdie
tunlesss
tatedothe
rwise
.
R
,requ
ired
, NR
,notrequ
ired
,IU
,inte
rnat
ionalun
it
s.
Sou
rce
: Ha
lve
r1989
;Wi
lson1991
;D’Ab
ramoe
tal
.1997
.

Thetab le showstha t not only dothe vitam ins required vary butthere
is also a var ia t
ioninthe d ietaryrequ i
remen tlevelI tistherefore very
di
fficu l
tto de term ine arecommendedleve lo fvi
tam in supplemen ta
tion
tha t w i
ll be sat isfac
tory for a l
lf ish spec ies
. The essent ial
ity and
requ iremen tsforthese v i
tam ins generally have been determined based
on we ight ga in, surv ival rate,tissue s torage , and o ther specif
ic
deficiency s igns .
56 CHAPTER2 NUTR
ITIONINTROP
ICALAQUACULTURE

Gu
ideQues
tions

1
. De
finethete
rm v
itam
in. Wha
tisthe gene
ralfunc
tion o
fvi
tam
ins?
2
. Why mus
tvi
tam
ins beinc
ludedinthe d
iet?
3
. D
ist
ingu
ish be
tweenl
ipid
-so
lub
le and wa
ter
-so
lub
lev
itam
ins? Name
the v
itam
ins tha
t are l
ipid
-so
lub
le; name the v
itam
ins tha
tare
wa
ter
-so
lub
le.
4
. G
ive names andsymp
toms o
f4v
itam
in de
fic
iency d
iseases
. Tabu
late
asfo
llows
:
Name o
fthe De
fic
iency d
isease Symp
toms
de
fic
ien
tvi
tam
in
a
.
b
.
c
.
d
.

5
. Wha
tisthe s
tab
le s
toragefo
rm o
fvi
tam
in Cinf
isht
issues?
6
. Wha
tisthe resu
lto
fvi
tam
in A de
fic
iencyinf
ish?
7
. Wha
tvi
tam
ins a
reimpor
tan
tbio
log
ica
l an
tiox
idan
ts?
8
. Wha
taretheconsequenceso
fhype
rvi
tam
inos
iso
rexcessv
itam
in Ain
f
ish?
9
. Wha
tvi
tam
inis essen
tia
lfo
r ma
intenance o
f no
rma
lblood c
lot
ting?
fo
r bonefo
rma
tion?
10
. Wha
tvi
tam
in p
lays animpor
tan
t ro
lein ca
lcium and phosphorus
me
tabo
lism?
11
. Wha
tarethefac
torstha
taf
fec
tthe requ
iremen
tfo
r av
itam
in?
12
. Whyisthe
re hype
rvi
tam
inos
isforl
ipid
-so
lub
le v
itam
ins?
M
INERALS
In
troduc
tion
Minera ls are inorgan ic substances that have many impor tant
func t
ionsin the an ima l body . They are requ ired to ma intain many
metabolic processes andto prov ide ma terialfor ma jor st
ruc tures(e .g
.
bones, tee th, exoske leton o f crus taceans )of aqua tic anima ls. They
are also requ ired for ma intenance o f osmo tic pressure , acid-base
balance(e .g.,the regu lation o fb lood pH , hemo lymph , urine and o ther
bodyf luids) andthe properfunc tion ing of musc les and ne rves. Un like
carbohydra tes,fats, and pro teins , they do not prov ide energy bu tthey
may serve as componen ts o fenzymes ,vitamins , and ho rmones . Fish,
especially mar inef ish spec ies,l ivein an env ironmen t tha t contains
many o fthe m inerals tha tthey needfor grow th and surv ival
.
This sec tion discusses the mac ro, micro, and t race m inerals; their
physiologicfunc tions; and de fic
iency s igns and symp toms. I talso gives
a summary o fthe m ineralfunc t
ions and m inera lrequ iremen ts off
ishes
and sh rimp .

C
lass
if
ica
tiono
fMine
rals

Abou t 20inorgan ic elemen ts are required to meetthe st ructural and

metabolicfunc tions o fliv
ing o rganisms. They are groupedin to mac ro,
micro
, andt race m inerals.
The macrom inerals ca lc
ium , phosphorus , magnesium , po tassium ,
sodium andch lorine a rerequiredinthe d ietinrela
tive
lylarge rquan tities
than m icrominera ls
.
The microminera ls are chrom ium , copper, cobalt,i ron, iod ine ,
manganese , molybdenum , selenium , and zinc. Sofa r
,t race m ine rals
such as a luminum , arsenic, cadm ium, lead, mercury,n ickel,s i
licon ,
t
in, and vanad ium have no knownfunc tioninfish.

Gene
ralfunc
tionso
fmine
rals

The biochem icalfunc t

ions ofm ineralsin aqua t
ic an imals are s
im ilarto
thoseinland an imals
,w i
ththe excep tion o fosmoregu la
tion. M inerals
may serve as components o f hard- tissue matr ices, softt issues ,
metal
loproteins, and as co factors o r activators of enzymes . The mo re
soluble minera ls (calcium, phosphorus , sod ium , po tassium, and
chlor
ine)func t
ionin osmoregu lation andin ma intenance o f acid-base
balance and membrane po ten t
ials.
Unlike vertebrates , there are severa l phys iological requirements
unique to crus taceans, par ticularly in the mo l
ting cyc le wh ich is
necessary in crustacean growth . A lthough some minera ls are
temporar ily stored in the t issues , such as the hepa topancreas , a
signi
ficant amoun tislos t dur ing mo lting or ecdys is. A summary o f
functions of m ineralsis shownin Tab le 2.14.
58 CHAPTER2 NUTR
ITIONINTROP
ICALAQUACULTURE

Tab .14 Summa

le2 ry o
fmine
ralfunc
tions

|M
ine
ral Func
tion and Essen
tia
li
ty

Ca
lcium S
truc
tura
lcomponen
tofha
rdt
issue
; Co
-fac
torfo
renzyma
tic p
rocesses
; Musc
le
func
tion and p
rope
r ne
rveimpu
lset
ransm
iss
ion
; Osmo
regu
lat
ion

Phospho
rus Componen
tof ha
rdt
issues and o
rgan
ic phospha
tes(e
.g ATP
, phospho
lip
ids
,
coenzymes
, DNA and RNA
); Bu
ffe
rfo
rthe ma
intenance o
f no
rma
l pH o
fin
tra
-
and ex
tra
-ce
llu
larf
luids

Po
tass
ium Ca
rbohyd
rate me
tabo
lism and p
rote
in syn
thes
is; Osmo
regu
lat
ion
; Ac
id-base
ba
lance
; Phospho
ryla
tionreac
tions

Magnes
ium Me
tabo
lism o
ffa
ts,ca
rbohyd
rates and p
rote
ins
; Ce
llu
larresp
ira
tion
;In
tra
- and
ex
trace
llu
lar homeos
tas
is; Phospha
te t
rans
fer andth
iam
ine py
rophospha
te
reac
tions
; Osmo
regu
lat
ion

Coppe
r Func
tionsinhema
topo
ies
is andinnume
rouscopper
-dependen
tenzymes(e
.g.
cy
toch
rome c ox
idase
,fe
rrox
idase
); Componen
tof hemocyan
in

Coba
lt Sou
rcefo
rmic
rob
ialsyn
thes
is o
f B1
2inin
tes
tine

I
ron Heme con
tain
ing enzymes (e
.g. cy
tochromes
, ox
idases
, perox
idase and
ca
talases
); Co
fac
torfo
r enzymes
; Hemog
lob
in

Iod
ine Neuromuscu
larfunc
tions
; In
termed
iary me
tabo
lism; Syn
thes
is o
fthy
roid
ho
rmones

Manganese Co
fac
toro
fva
rious enzyme syn
thes
is

Se
len
ium Componen
tofg
luta
thione pe
rox
idase

Z
inc Co
fac
torin severa
l enzyme sys
tems
; Componen
tof a large number o
f
me
tal
loenzymes

Sou
rce
: Ha
lve
r198
9

M
ine
ralAva
ilab
il
ity

In gene ral,b ioavai

lab i
lity o
f m inerals has beenfound to be pos it
ively
correla
ted w ith their solubi
lityin wa te
r. H igh ly soluble sa
lts appearto
be bene f
icial
, bu ttheirleach ing ratesf rom d iets haveto be measured .
In add ition toinorgan ic sources, organ ic che lates and comp lexes of
minerale lemen ts a
re use ful means o f del
ivering m inerals
. Cons idering
thesel im i
tations, water solub leinorgan ic salts orb ioavai
lable organic
salts are preferablefo r usein m ineralp remixes .

Mac
rom
ine
rals

Ca
lcium
Calciumis mos t
lyfoundinthe ske leton and sca les o f bonyf ish andin
the exoske leton of crustaceans . F ish sca les are animpor tants ite of
calcium me tabolism and depos i
tion . As idef romi ts struc turalfunc tion,
calciumisimpor tantin phys iologica l processesinc luding me tabolism ,
nerve and musc le contrac
tion, ne rveimpu lset ransm ission , maintenance
F
igu
re 2
.15
of cellin teg r
ity, osmoregu la
tion , and ac t
iva tion o fimpor tant enzymes .
Ca
lciumandpho sphorus Calcium and phospho rus def
ic iencies can cause so ft-shellingin sh rimps
def
icienc
ies. Sof
t-she
l
ledshrimp (Figure 2 .15).
ma
inlyduetoca lc
iumand
phosphorus def
icienc
ies ha
s Fish mayto tal
ly or part
ially mee tthe ir calcium requ iremen tthrough
causedlo s
sesinthesh r
imp abso rptiono fcalciumf romthe wa terv iathe irg i
lls
,f insand o ralepithe l
ia.
industry
.
Sou
rce
: Lav
il
la2
001 The g i
lls a rethe mos timpor tan ts ite of ca lcium regu la tion. Gene rally,
 CHAPTER2 M
ine
rals 59
calc
ium f rom feed ingredien
ts for examp
le,f
ish mea
l, may meet the
requirementsof mostfish
, however
,itisacommon p
ract
icetosupp
lemen t
thefeed w i
th calc
ium .

Phospho
rus
Phospho rusis ano the r ma jorm ineralrequ ired byf ish andisanimpo rtan t
cons ti
tuen to f nuc leic acids and ce l
l memb ranes . Itis d irectlyinvo lved
intheene rgy-produc ingce l
lularreac tions andis acomponen tofessen tial
molecu lessuch as adenos inet r
iphospha te(ATP ). Feedisthe ma insou rce
ofphospho rusforf ish becausetheconcen t
ra tiono fphospho rusin na tu ral
watersislow . Consequen tly,the needfo ra d ietary source o fphospho rus
is more cr iticalthanfor ca lcium becausef ish mus te ffectively abso rb,
store, mob i
lize, and conse rve phospha tein bo thf reshwa ter and seawa ter
env i
ronmen ts .
Mos tf ish spec iesrequ ire a dietaryinc lusion o f0 .5%to 1 .0% ava i
lab le
phosphorus . Gene rally
, phosphorus requ iremen ts are no ta f
fected by
dietary ca lciumleve ls. In con tro
lled exper imen ts, the grow th o f both
common ca rp andt rou t have been shownto be pos it
ive ly correlated w ith
dietary phosphorusleve ls bu t no tw i th ca lciumleve ls. The apparen t
phospho rus ava ilabilityva luesfo r seve ral phospho rus sou rces have been
estima ted. They a re : ca lcium phospha te monobas ic, 46% ; calcium
phospha te d ibas ic, 14% ; calcium phospha te tribasic, 5 .7% ; sod ium
phospha te monobas ic , 70% ; and po tass ium phospha te monobas ic, 68% .
Dietary ca lcium and phospho rus areimpo rtan tinthe p reven t
ion o fso ft-
she l
lingin shr imps .

Sod
ium
, po
tass
ium
, andch
lor
ine
Sodium, po tassium , andch lorinea rethe mos tcommonino rgan icelemen ts
foundinf ish . Theya reabundan te lectro
ly tesinthebodyanda reessen tia
l
for a numbe ro f physiologica lprocesses . Appropr iatelevels o ftheseions
arerequiredfo rprope rfunc tioningo fcellsandfo r maintainingne rvefunc tion.
Fishread i
lyabso rbthesee lemen tsf romtheaqua tic med ium , especiallyin
marine wa ters, andthet issueleve ls oftheseions a re ma inta ined as a
resultofosmo regu la
tion. Theya re abundan tinsea wa terandincommon
feedstuf
fs usedinprepa rationo ff ish diets. Thus,d ietarysupp lemen tation
ofthese elemen tsis norma lly no trequ ired.

Magnes
ium
Alarge propor tion o f magnes ium inf ishis con tainedin the ske letal
tissue. Magnes iumis an essen tia
l cofactorin many me tabolic react
ions.
These enzymesinc ludethe phosphok inases, thiokinases, phospha tases,
pyrophospha tases, and am ino acy l synthetases. I tis also neededin
ske le
taltissue me tabol
ism , osmo regulation
, andin ma intaining musc le
tone .
Generalsymp tomso f magnes ium de f
iciencyincludereduced we ightgain
and poo rfeed conve rs
ion . Inra inbowt rout, magnes ium de f
iciencyleads
torena lcalcinosis andf lexibil
ity ofthe musc le, part
ly dueto aninc rease
in ex tracel
lularf luid volume . Mos tfeeding redients, especiallythose o f
plant or igin, are abundant in magnes ium thus , magnes ium
supp lemen tationin prac ticald ietsis generally notrequ ired.
60 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

M
icrom
ine
rals

Coppe
r
Coppe risanimpo rtantcomponen to fanumbe ro f metalloenzymestha ta re
involvedin a w ide variety o
f me tabo l
ic processes. I tis assoc iated w ith
cytochrome c ox idase ofthe e lectront ransport cha inin ce l
ls.I tisthe
prima ry oxygenca r r
ierin hemocyan in ofcrustaceans and mo l
luscs. F ish
appears toto lerate copperin the d ietthan d issolved copperf rom the
water
. Concen trations of0.8-1.0 mg coppe r perliterin wa ter aretox ic
to manyf ish species. Feeding red ien
ts such asf ish so lubles
,k r
ill meal,
and yeas t con tain relat
ively highleve ls of coppe r
. Shr impin genera l
canno t mee tthe ir physiolog
ica l requiremen tfor copperf rom seawa ter.
In wh i
te shrimpPenaeus vanname i, symp toms ofde ficiencyinc lude poo r
growth , reduced copperleve lsin the carapace , hepa topancreas and
hemo lymph , and en largemen to fthe hea rt
.

I
ron
Die
taryi ronis essen tialinf ishfo rb loodfo rmation, maintaining no rma l
hemog lobin con tent, hema tocri
tva lue, and cellsize. A min imum d ietary
concen trat
iono f 150µg pe rgram d ietisrequ i
redto p reven
ti ron deficiency
resultingin hypochrom ic, m icrocy t
ic anem iain red sea bream and
common ca rp
. D ietaryi ron de ficiency has no t been observedin sh rimp.
In con trast,d ietary supp lemen ta t
ion o f shrimpfeed w ithi ron grea ter
than 150µg per gram has been found to g ive poor growth . S ince
crustacean d iets gene ral
ly con ta in po lyunsaturatedfa t
ty acids, excessive
supplemen tat
iono fferrousi roninthed iet maya f
fectdietstabil
itythrough
increasedl ip
id ox idation .

Manganese
Manganeseisimpo rtante i
ther as a co
factorthat activates metal-enzyme
comp lexes or as an integra l par to f metalloenzymes in prote in,
carbohydra te
, andl ipid metabo l
ism . The up take o f manganesef rom
water byf ish has been demons trated butitis mo re eff
icient
ly abso rbed
fromfeed . Howeve r, duetothe po tent
ialinhibitory ef
fects of phyt
ic acid
on the b ioava ilab i
lity of manganese , feed supp lemen ta
tion may be
desirable. D
ie tary defic
ienciesinf ish have resultedin poor grow th
,
skeletal abno rma li
ties, embryo mo rta
lit
ies, and poo r hatching rates.

Se
len
ium
Seleniumis anin teg ra
l componen to fthe enzyme g lutathione pe roxidase
tha tp ro
tects cells and memb ranesf rom de leter
ious effects of peroxides.
In con junction w ith vitamin E , this enzymefunc t
ions as a b iological
ant ioxidant wh ich protects b io logical membranes aga inst l ipid
peroxidation. Se len iumimpa rts a p rotec
tive ef
fect aga instthetox icity
of heavy me tals such as cadm ium and mercury . The se len ium
requ iremento ffishva r
ies withthe po lyunsa turatedfattyac id andv itam in
E con tento fthe d iet. Bo th selen ium and v i
tam in E are requ ired to
preven t muscu lar dys trophyin A tlan t
ic salmon . In gene ra
l, prac tical
diets containingf ish mea l contains adequa te amoun to f selenium and
does no trequ ire supp lemen tation.
 CHAPTER 2 M
ine
rals 6
1
Z
inc
Zincis anin tegra l componen to f me talloenzymes
includ ing dehydrogenases ,a ldolases , pep tidases,
and phospha tases . Approx ima tely 20 d i
fferent
enzymes has beenfound to con tain z inc. Many
metabolicfunc tions a re af
fec ted by z inc de f
iciency.
Fish can accumu latez incf rom bo th wa ter andfeed
but d ietary z inc is more e fficient ly absorbed .
Practica ld iets con ta in feeds tu f
fs tha t are good
sources o fzinc ;for examp le,fish mea l. Howeve r
,
zinc b ioava ilab i
lity is genera l
ly very low , thus
supp lemen tat
ionis essen t
ia l. Feeds tuffstha tare F
igure 2.16
Z
incde f
iciencyca taract
:
relat
ive ly highin phy tate mayfur therreduce z inc non
-a va
i
labil
ityo fzincduetoh ighle
velsof
bioavailabili
ty. Ca taractis a common symp tom of ca
lc
iumandpho sphorusinf
ishfeedhas
zinc de fic
iencyinf ish(F igure 2 .16). causedlosse sinthefishfarm
ingindus
try
.

T
race m
ine
rals
Info
rma t
ion onthe d ie
ta ryrequ i
rements ofothermicromine
ralsisl imited
.
A numbe roftrace elemen ts a
rerequiredfor adequatef
ish nu tr
ition but
theirdietary def
icienc ies have not been reported. Coba lt acts as a
componen tofvitamin B1 2
. Sul
furisrequiredforthesynthesis o
fcys te
ine.
Chromiumisimpo r
tan tin no rmalg
lucose andl ip
id metabol
ism .

M
ine
ralSupp
lemen
tat
iono
fPrac
tica
lFishD
iets

Although the d ietary m inera l requ iremen ts o ffish have no t been we ll

estab l
ished , prac t
ica ld iets are usua lly supp lemen ted w ith a minera l
prem ix. Prac ticald iets may norma lly con ta
in subs tan t
ial amoun ts o f
endogenous m inera ls, thus a comp lete m ineral prem ix may no t be
necessary . Excess ive m inera l supp lemen tationincreases the cos to f
feed and may reducethe b ioavailab ility ofo ther minerals andinc rease
phosphorus po llution . Many comme rcialfeed b inders may a lso con tain
high leve ls of ca lcium and magnes ium tha t may reach undes irab le
levels o f these m ine ralsinthefeed .
Most stud ies on the minera l requ irements o ff ish have been
conduc ted us ing semipur if
ied d ie ts. Dietary requ irements are
determ ined byfeed ing gradedleve ls o fthe e lemen t being studied and
measur ing the phys iolog ical response o fthe tes t anima l. M inera ls
requ ired by f ish are ca lcium , magnes ium , phosphorus , and m icro
elemen ts such as coppe r,iod ine,i ron , manganese , selenium and z inc.
Ingenera l,the d ie taryrequ iremen tsfor m inerals are poorly unders tood
because o fthe d i
fficu ltyin dev ising m inera l-def
icien tdiets and the
need to dep lete tissue m inera ls tores .

M
ine
ral Requ
iremen
tso
fFish

A
lthoughinorgan ic elemen ts a
re requ
i redfo rthe no rma llife processes
offish, know ledge ofthe ir mineral nu tr
itionis s t
ill one oftheleas t
knowna reasoffish nutr
ition. Unl
ikeo the r nu
trien
ts,s igni
ficantamoun ts
of minerals can be absorbedf romthe ir external environmen tthusi tis
62 CHAPTER2 NUTR
ITIONINTROP
ICALAQUACULTURE

dif
ficu l
t to con trol the die tary intake o fthe m ineral being s tudied .
Calcium ,sod ium, potassium ,iron,zinc ,coppe randse leniuma regene rally
derived byf ishf rom the rear ing wa ter. The exchange o fionsf rom the
surround ing wa ter across the g il
ls and sk in offish comp l
ica tes the
measuremen ts of m ineral requiremen ts. In terac
tion be tween m ine rals
furthe rcomp l
icatesthe assessmen to fdietaryrequ iremen ts
. Sofa r,the re
is veryl imitedin forma t
ion onthe d ietary requiremen tsfo r mine ra
ls by
fish.
A summa ry ofthe known d ietary m ineralrequ iremen ts ofvariousf ish
spec iesis presen tedin Tab le 2.15 and the ir defic
iency symp toms a re
summa r
izedin Tab le 2.16.

T
able2
.15 Summa
ryo
fthe m
ine
ralrequ
iremen
tso
fva
riousf
ishandsh
rimpspec
ies
Channe
l Common Japanese N
ile Ra
inbow Ku
ruma T
ige
r
ca
tfish ca
rp ee
l t
ilap
ia t
rou
t sh
rimp Sh
rimp

Ca 0
.27% 1
.0-2
.0% 1
.0%

P 0
.33
-0.45% 0
.6-0
.7 % 0
.58 % 0
.8-1
.0% 0
.7-0
.8 % 1
.0-2
.0% 1
.0%

Mg 0
.04 % 0
.04
-0.05% 0
.04 % 0
.05
-0.07 % 0
.30%

Cu 3µg
/g 3 -4µg
/g 3µg
/g

Fe 150µg
/g 170µg
/g

Mn 13µg
/g 13µg
/g 12µg
/g

Zn 20µg
/g 15
-30µg
/g 10µg
/g 15
-30µg
/g

Sou
rce
: Wa
tanabee
tal
.1988
; Bau
tis
taand Ba
ticados1
988

T
able2
.16M
ine
ralde
fic
iencysymp
tomsinf
ishandsh
rimp
M
ine
rals | De
fic
iency symp
toms |

Ca Poo
rgrow
thinchanne
lca
tfish
; So
ft-she
ll synd
romein c
rus
taceans

P Poo
rgrow
th & ske
leta
l abnorma
lityin common ca
rp & ra
inbowt
rou
t; Lowfeed e
ffic
iency and h
ighl
ipid
con
ten
tincommon ca
rp; Low ashin who
le body andve
rtebra
teincommon ca
rp,ra
inbowt
rou
tand channe
l
ca
tfish

Mg Poo
rgrow
th and h
igh mo
rta
lityincommon ca
rp andra
inbowt
rou
t;S
lugg
ishness andconvu
lsionincommon
ca
rp, ra
inbowt
rou
t and channe
l ca
tfish
; H
igh Ca con
ten
tin bone o
f ca
rp and ra
inbowt
rou
t; Ano
rex
iain
channe
l ca
tfish
; Ske
leta
l abno
rma
lit
ies andrena
lca
lcinos
isinra
inbowt
rou
t

Cu Poo
rgrow
thincommon ca
rp andra
inbowt
rou
t; Dwa
rfisminJapanese ee
l

Co Poo
rgrow
thincommon ca
rp

Fe Anem
iain common ca
rp

I Dwa
rfisminJapanese ee
l

A
l Dwa
rfisminJapanese ee
l

Zn Poo
rgrow
th,h
igh mo
rta
li
ty,e
ros
ion o
ffins and sk
in,low Zn and Mn con
ten
tin bone o
fcommon ca
rp and
ra
inbowt
rou
t; Dwa
rfisminra
inbowt
rou
tandJapanese ee
l; Ca
tarac
tinra
inbowt
rou
t;Low Ca
, Mg and Pand
h
igh mor
tal
ityin common ca
rp; Low Zn and Mnincommon ca
rp andra
inbowt
rou
t

Sou
rce
: Wa
tanabe e
tal
.198
8
 CHAPTER2 M
ine
rals 63
Gu
ideQues
tions

1
. Wha
tare m
inera
ls?
2
. Wha
tarethe 3 g
roups o
fmine
rals? G
ive examp
les o
feach g
roup
.
3
. Wha
tarethe gene
ralfunc
tions o
f m
ine
rals?
4
. Wha
t m
inera
lis neededfor norma
lforma
tion o
f bones
, sca
les and
tee
thinf
ish?
5
. What e
lement is a component o
f thyro
id hormones? o
f
me
tal
loenzymes?
6
. Wha
t m
ine
ralis a componen
tofv
itam
in B1
2?
7
. Wha
t m
ine
ral pro
tec
ts b
iolog
ica
l memb
ranes aga
ins
tlip
id ox
ida
tion?
8
. Wha
t m
ine
rals a
reimpor
tan
tto preven
t so
ft-she
llingin shr
imps?
9
. Whyisthe needfor a d
ietary source o
f phosphorus mo
re cr
itica
l
thanfor ca
lcium?
10
. Wha
t m
ine
ralis a componen
tofg
luta
thione pe
rox
idase?
11
. Wha
t m
ine
rals a
re neededfo
rbloodfo
rma
tioninf
ish? hemocyan
in
forma
tionin mo
llusks and c
rus
taceans?
12
. Wha
tisthe consequence o
firon de
fic
iencyinf
ish?
13
. Whyis m
ine
ral nu
tri
tion o
ffish one o
ftheleas
t known a
reas o
ffish
nu
tri
tion?

Summa
ry

P
rote
insandam
inoac
ids

Am ino ac ids a rethe bas ic un ito fp rote ins and have a gene rals truc tu re
with an am ino g roup( -NH2 ) and aca rboxy lg roup( -COOH ) bondedtothe
alpha -ca rbona tom . Thena tu reo fthes idecha in,re ferredtoasthe Rg roups ,
are the bas ic d ifferences among am ino ac ids . The am ino acids a re
dividedin totwo g roups: the d ispensab le or non -essen tial amino ac ids
and theind ispensab le or essen tia
l am ino ac ids . O fthe 20 na tura lly
occur ring am ino ac ids, ten a re essen tial because they canno t be made
bythef ish o r canno t be madein amoun tstha t sa t
isfythe requ i
remen t.
These a re: arg inine,h is t
id ine , isoleuc ine ,leuc ine, lysine , me thion ine ,
pheny lalan ine , threon ine,t ryptophan , and va line . The non -essen tial
am ino ac ids can be made by the an ima lf rom carbon and n itrogen
precursor compounds . Cys teine(wh ich can rep lace pa r
to f methion ine )
andty ros ine(wh ich canrep lace pa rto f pheny la
lan ine),g lycine,glu tam ic
acid,g lutam ine , aspartic ac id , aspa rag ine ,a lanine ,p roline, and se r
ine
are non -essen tial am ino ac ids .
Proteins are po lymers o f am ino ac idsjo ined by pep tide bonds . The
struc ture o f pro teinsis comp lex, with no obv ious regu lars tructu re. To
fac i
litate s truc ture determ ina tion,i tis cus toma ryto de f
inefourleve ls o f
organ iza t
ion . Pr imary s truc tureisthe o rde rin wh ichthe am ino ac ids
are cova len tlyl inked toge ther . Secondary s t
ruc tureis the hydrogen
bonded a rrangemen to fthe po lypep tide cha in. Tertiary s t
ruc tureinc ludes
the three d imens ional arrangemen to fa ll the a tomsin the pro tein.
64 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

Qua ternary s t ruc tureisthe ar rangemen to f subun itsin mu lt

i-subun i
t
proteins . P rote inis d iges ted and b roken downtoi ts componen t am ino
acids, wh ich a re used bythe o rgans andt issuesto make new p rote
ins
(grow th) or rep lace ex is t
ing pro teins (ma in tenance )
. Excess can be
conve rtedto ene rgytofue lthe ac t ivity o
fthef ish.
S
ince p rotein qua lityis de te rminedla rge
ly byi ts am ino acid compos it
ion
and d iges tibi
lity,in fo rma tion onthe p rote
inrequ iremen to ffishisl imi
ted
withou t da ta onthe essen tial am ino ac idrequ iremen t
. The nu trit
iveva lue
ofa d ietary p ro teinis dependen t ontheex tentto wh ichi tsessen t
ial amino
acid compos it
ionfu lfi
llstherequ iremen tofthe o rgan ism . A de ficiencyin
any essen tial am ino ac id w i l
lresu l tin poo rg row th . Thel imiting am ino
acido fap roteinistheessen tialam inoac idtha tisp resen tinlowes tamoun t
relat
ivetotherequ iremen t.P rote
insf roman ima lsou rcesa re mo re digest
ible
than p roteinf rom p lantsou rces
. P roteinsf rom p lan t sources a re defic
ient
in some essen tial am ino ac ids; such as , methion inein soybean mea l, and
lysinein com mea l.P rope rcomb ina tion ofp rote insf rom bo th an ima l and
plant sou rces p rov ide d iets w ith ba lanced am ino ac id p rof
iles.

L
ipidsandfa
ttyac
ids

Lipids a recompoundstha ta reinso lub lein wa ter andso lub lein nonpo la r
organ ic so lven t. L ipids a re componen ts o fce l
lu lar memb ranes ,p rovide
essen t ialfa tty ac ids and a re good sou rces o f ene rgy. Tr iglycer ides and
waxesa rethes toragefo rmso ffattyac ids , wh i
lephospho lipidsa reimpo r
tan t
componen ts ofb io log ica l memb ranes such assph ingo lipids . S teroids a re
precurso rs o fsex ho rmonesinf ish and a reimpo rtan tinthereproduc tive
processes .
Fa ttyac idsa rea lipha ticca rboxy licac idsw i
tha — COOHg roup . Theimpo rtan t
fatty ac ids have be tween 2 and 24 ca rbon a toms and may be comp letely
satu ratedo rhaveone(mono -unsa turated) ,twoo r more(po lyunsa turated,PUFA )
or mo re(h igh lyunsa tu ra ted, HUFA )doub lebondsintheca rboncha in
. Essen tial
fattyac idscanno tbesyn thes izedinthebodyand mus tbep rov idedinthed ie t
.
The essen tialfa tty ac idsforf ish a re e ithe rthe po lyunsa tura ted o rthe
highly unsa turatedfa t ty acids .
The compos i
tion o ffa tty ac id in f ish is a f
fec ted by a number o f
env ironmen talfac to rs: sa l
in i
ty,tempe ra tu re
, and d iet. D ietisthela rges t
sing le fac tortha ta f fec ts thefa tty ac id compos ition . Thefa tty acid
requ iremen t var ies amongf ish spec ies . Mar inef ishes need more n -3
PUFAs . L ino len ic , l ino leic , arach idon ic, e icosapentaeno ic and
docosahexaeno ic ac ids a re essen tialfa t
ty ac idsfo rf ish. L ipids such as
cho les tero l andlec ith in a re a lso needed by sh rimps . F ishl ipids have
more n -3fa tty ac ids wh ile p lan tl ipids con tain more n -6fa tty acids .
Somef ish requ ire n -3fa tty ac ids; some need bo th n -3 and n -6fa tty
acids , wh i
le o the rsrequ ire n -6fa tty ac ids. The d ieta ryl ip idrequ iremen t
offish can be de rivedf romthe irfa t
ty ac id p rofi
les .

Ca
rbohyd
rates

Carbohydrates a
ree i
thera ldehydes or ke
tones withtwo or mo re hydroxyl
groups. A carbohydratelike glucoseis ca l
led an aldose because o fthe
presence o
fan a ldehyde group, whi
lef ructoseis ca
lled a ketose because
 CHAPTER2 M
ine
rals 65
ofi ts ke tone group . The s imp lest form o f carbohydrates are
monosacchar ides , compoundstha t con ta in a single ca rbony lg roup and
two or more hydroxy l groups . Monosacchar ides can undergo var ious
reactions wh ich g iver iseto o l
igosaccha rides and po lysaccha rides. Three
impor tanto ligosacchar ides a rethe d isacchar ides suc rose ,lactose and
maltose
. Suc roseis a commontab le suga r
,lac tose occu rsin m ilk, and
maltose is obta ined f rom the hydro lys is ofs tarch , an impor tant
polysaccharide . Po lysaccha ridesa refo rmedbythecomb ina tiono fhexoses
oro ther monosaccha rides. S tarch wh ichisfoundin p lan ts whileg lycogen
whichoccursinan ima ls areene rgys torage po lyme rso fg lucose. Ce l
lulose
and ch i
tin are po lysaccha r
ides based on s ingle monome r un i
ts ofg lucose
and N -acetylg lucosam ine, respec t ively.
Carbohydra tes a re a cheap sou rce o fene rgy and,l ikel ipids, can spa re
protein as an energy source . The ab i
lity off ish to ut il
ize d ietary
carbohydra tesfo r ene rgyva ries. Ca rnivo rous spec ies have mo rel im ited
abili
tythan omn ivorous and he rb ivorous spec ies. D ietsfor ca rn ivorous
fishes containlowe r ca rbohydra televe lsthanthose o f herbivores .S ince
carbohydra teistheleas t expens ive ene rgy source ,itis advan tageousto
use as much ca rbohydra te asthef ish can u t
il
ize.

Ene
rgy

Ene rgyisde fined asthecapac ityto do wo rk.I t mayex istin d i

fferen tfo rms
anddod ifferen tkindsofwo rk. Ene rgyisob tainedf rom ma jorfoodnu tr
ien ts:
carbohyd ra tes ,lipids,andp roteinsand unde rgoseve ralchem ica
lp rocesses
such as ca tabo lism and ox idation w i
th in the an ima l body . Adenos ine
triphospha te(ATP )isthe pr incipa l drivingfo rceinthe energy -requ iring
biochem ica lp rocessesofl i
fe. Anene rgybudge tistheamoun tofene rgyin
inges tedfoodtha tis ut
ilizedfo reach ma jor processessuchasg row th and
reproduc tion . A gene ralized ene rgy budge tfo r carnivo res and he rb
ivo res
has been deve loped by Bre ttand Groves(1979 ).
Fish like other an ima ls, ea t pr imar ily to sat isfy the ir energy
requ iremen ts . Whenfedlow ene rgyfeed ,fish are ab leto compensa te by
eating mo re o fthefeed w ithin ce rtainl imits. Itis essen tia
ltha tfish a re
given ara tion w ith suf
ficien t energyto mee tthe i
r ene rgy needs.
Fromthepo into fv
iewo fanaquacu lturist,thequan t
ityandcos tofene rgy
whichis ava ilablefor g row th of cu lture spec iesis mos timpo rtan t.

V
itam
ins

Vi
tam ins a re comp lex compounds whose p resenceinthe d ietis necessa ry
for no rmalg rowth and ma intenance o f heal th and whoseinsu ffic
iencyin
thed ie tresu l
tsinthedeve lopmen tofspec if
icpa thologicconditions. Vitamins
are essen tialfortheregu lat
ion o fcel
l me tabo lism andfo rt ransforma tion o
f
energy . They may ac t as co-enzymes o r pa rto fan enzyme sys tem . They
occu rint race amoun tsin mos t naturalfood and canno t be synthesized by
mostf ishes hence , mus t be providedinthe d iet
.
Vitam ins are c lass
ifiedin to wa ter-solub le andl ipid-soluble. Wa ter-
solub le vitamins are hydroph il
ic wh ile l ipid-solub le vitamins are
hydrophob ic
. The wa ter-so luble vitam ins are th iam in (vitam in B1),
riboflav in (B2 )
, pyridoxine(B5 )
, pan tothen ic acid (B6)
, coba lam in (B1 2
)
,
66 CHAPTER2 NUTR
ITIONINTROP
ICAL AQUACULTURE

asco rbic ac id(vitam in C )

,n iacin,fo lic acid,inos ito
l, cho l
ine , and biot
in .
Lip id-solub le vitamins are ret ino l (vitam in A ), cho lecalciferol or
ergoca lciferol(D )
, α- tocophero l(E ) and menad ione (K ). Wa ter-solub le
vitam ins may be excre ted in the ur ine and rare ly bu ild up to tox ic
levels. In con trast
,l ipid-so
lub le vitam ins may be exc re
tedinthefeces o r
accumu lateinthet issues. A de r
ivative ofv i
tam in A p lays ac rucialrolein
vision. V itam in D con t
rols calcium and phospho rus me tabo l
ism ,a f
fect
ing
the s truc tura lintegrity of bone . V itamin Eis an an tiox idant, and the
presence o fvitam in Kisrequ i redinthe b lood clott
ing p rocess. Tox icity
mayoccu rf romanove rdoseofv itam insAand D . Mos tv itam inrequ iremen ts
are knownfo rjuveniles ofcoldwa ter, bu t notfor wa rm wa terfishes.

M
ine
rals

Minera ls are requ iredin theforma t

ion o f bone , sca les, and tee th, in
osmo regu lation,intheregu lationo fne rvefunc t
ionandin ma intenanceo fb lood
pH. Some m ine rals a recons tituen ts o fso ftt issues, enzymes ,v i
tam ins, and
hormones . The mac ro m ine rals: calc ium , phospho rus, magnes ium , sodium ,
potassiumandch lo rine,a rerequ iredinthed ietinla rgeamoun ts. The m icro
minerals: ch rom ium , cobalt, coppe r
,iod ine,i ron , manganese , mo lybdemum ,
selenium and z inc ,a re requ iredin much sma ller amoun ts. The t race
minerals have no knownfunc tioninf ishesand a re mos tlytoxic. Ingene ral
,
theb ioava ilabil
ityo fm inerals hasbeenfoundtoco rrelate withthe irsolub i
lity
in wa te r
. Wa ter -solub leino rgan ic sa lts o rb ioava ilab le organ ic salts a re
preferredfo r usein m ineralp rem ixes .
Mine ral nu tri
tiono ffishisoneo ftheleas tknowna reaso ffish nu tri
tion.
Unlikeo the rnu trien ts,fishcande rives ignifican tamoun tso fm ine ralsfrom
theirex terna lenv ironmen t
,thusi tis d if
ficu lttocon trolthe irin take o fthe
minera l beings tud ied. In terac tion be tween m ineralsfur thercomp licates
the stud ies on d ieta ry m ine ralrequ iremen ts o ffish. Howeve r,a lthough
fishes can de r
ive many m ine ralsf romthe wa ter, m ine ra
l supp lemen tsin
the die timp roveg row th, surv ivaland hea lth .

Sugges
ted Read
ings

P
rote
insandam
inoac
ids

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A
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.
 CHAPTER 2 The Essen
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l Nu
trien
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Ha
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fguppy
Poec
il
iaret
icu
lata
. Nippon Suisan Gakkaish
i 55
:1947-1953.

Wa
tanabe T
, Satoh S, Takeuch
iT. 1988
. Ava
ilab
ili
ty o
fmine
ralsinf
ish
mea
ltofish. As
ian F
ish.J.1
:175-195
.
 CHAPTER

3
Feed
ing Hab
itsand D
ige
stive
Physio
logyo
fFishes
ILDA G
. BORLONGAN
, REL
ICARDO M. COLOSO
,
and NELSONV
. GOLEZ

In
troduc
tion
Thischap terprov idesbas icin forma t
iononthefeed inghab itsandbehav ior,
andphys iologyo ffishesandc rustaceans .The mechan ismstha tcon trolthe
movemen tand d iges tiono ffood , me thodso fassess ing diges tib
ilityo ffeed ,
facto rs affecting d igestion and abso rpt
ion o ffood nu t
rien ts, andfeed ing
processesinf ish a re discussed . An unde rs tand ing ofthefeed ing hab its,
feed ing mechan isms , andthe d igestion and abso rption processescan he lp
fishfarme rs and nu tri
tion is ts maxim izethe useo ffeed. Thera te at wh ich
fish d igestthe i
rfoodiso fpr ima ryimpo rtancein de termin ingfeed ingra tes,
frequency , andra tion si ze. Know ledge ofthe d iges t
ive phys iologyo ffishis
also necessaryfor an e ffect ivefeedfo rmula tion andin choos ing a p rope r
feed ingreg ime.
Thischap tera imstoteachthereade r: thefeed ing hab itsand behav ior
offishes andc rus taceans ;thes tructuraladap tationinthe ana tomyo fthe
diges t
ivet rac
t;the var ious o rgans o fthe d igestive systems o ff ishes and
crus taceansandthe irfunc tions ; nutr
ientd iges t
ionandabso rptionbyf ishes
andthefa teo fdiges tedand und igestedfood ;thefac torstha taffec tthera te
ofd iges t
ion and abso rption ; andthefeed ing p rocessinf ish.

Feed
ing Hab
itsand Behav
ior

Thefeeding habi
ts and behavioroffishesre
fertothe processofthesearch
forandinges t
ionoffood. Thisalsoincludesthe manne randthes t
imul
ifor
feed
ing.
F
ishescan bec lass
ified accord
ingtothe i
rfood and die
t, wh
ichrefe
rto
the mater
ialsthey habi
tuallyea tas:

He
rbivo
res - thosethatfeedexc
lusive
lyon plan
t mater
ials
Ca
rnivores - thosetha
tfeedexc
lusive
lyonan imal ma
tter
Omnivo
res - thosethat der
ivethe
ir nut
rien
tsfrom both p
lan
ts
andan
ima
ls
P
lank
tivo
res- thosetha
tfeedon p
lank
ton
,the m
icroscop
ic p
lan
t
and an
imall
ifein wa
terinc
ludingbac
ter
ia
De
tri
tivo
res- thosetha
tfeedon decay
ing mat
ter
78 CHAPTER3 NUTR
ITIONINTROP
ICALAQUACULTURE

Food availab
ili
tyis a keyfactorin determ in
ing wha tthef ish w
ill ea
t.
Mostfishesare h
igh lyadaptab
leintheirfeed inghabitsand u t
il
izethe most
readilyavai
lablefoods. Table 3
.1summa rizesthe naturalfood andfeeding
habitsofcommon lyculturedspecieso
ffishandsh rimp .

T
abl
e3.1
. Feed
inghab
itsandna
tura
lfoodo
fsomejuven
ilef
ishandsh
rimp
F
ish Feed
ing Hab
it Na
tura
l Food

M
ilk
fish m
icrophagous M
icrop
lank
ton(
lab
-lab
), ben
thic a
lgae
p
lank
tivo
re (
fi
lamen
tous g
reen a
lgae
)

Seabass ca
rnivo
re F
ish o
fsame spec
ies o
rothe
rfish

Groupe
r ca
rnivo
re F
ish o
fsame spec
ies o
rothe
rfish

Snappe
r omn
ivo
re F
ish
,crabs
,stoma
topods
, mo
llusks
,crus
taceans
and o
the
r bo
ttom dwe
lle
rs
Ra
inbowt
rou
t ca
rnivo
re F
ish

Channe
l ca
tfish omn
ivo
re Insec
ts, sna
ils
, wo
rms
,plan
ts and gene
ralo
rgan
ic
deb
risin muddy bo
ttom

Common ca
rp omn
ivo
re P
lan
ts and o
the
rorgan
ic deb
risin muddy bo
ttoms

S
igan
id he
rbivo
re Mac
roa
lgae

Mu
lle
t omn
ivo
re Sma
lla
lga
l ce
lls and o
the
rorgan
ic deb
ris

Sh
rimp omn
ivo
re Sma
llc
rabs
, sh
rimps
, mo
llusks
,fish
, po
lychae
tes
,
a
lga
l ma
tte
r, and deb
ris

Ano the rc lass ifica

tiono ffeed ingbehav io ro ffishesacco rdingtothe manne r
o
ffeedinga re :
1
.P reda to rs aref ishestha tfeed on mac roscop ic an ima ls. They may
ei
the r becons tan tlyonthe move , hun t
ing and pursu ingthe irp rey,
orl ie-on -wa itto ca tch p reytha ts trayin tothe i rter r
ito ry. Some
preda to rsfeed uponsma llfisheso rinsec tsfounda tornea rthe wa ter
su rface .P redators ma inlyusev is iontohun tfo rp rey,a l
thoughsha rks ,
eels, and o therp reda to ryf ishestha tfeed a tn igh t may a lsore ly on
sme l
l,tas te, andla terall inesenseo rganstoloca tethe i
rp rey.
2. G raze rs feed onbo ttomo rgan ismso rp lank tonstha ta rese lec tive ly
consumed . Someg raze rsfeedona lgaeo rn ibbleonco ralree fstoea t
polyps . The ac tua l tak ing o ffood is by b ites wh ile brows ing
con tinuous ly.
3.S tra ine rs arethose wh ichf ilter o rgan isms , ma inly d iatoms and
crus taceansf rom wa ter. Thesef ishes sw imthrough r ich p lank ton
beds ,filterthe wa ter,andswa llowthesoup -
likeconcen t
ra te.S t raine rs
norma lly have nume rous ,f ine, ande longa tedg illrake rs.
4. Sucke rsarethose wh ichsuckin mudo rfood -con taining ma terialto
ob tainthe i rfood . Some times ,foodi tems a re separa tedf rom the
sed imen ts be fore be ingswa l
lowed ,a l thoughinsomeca t
fishes ,food
isinges tedtoge the rw i
thf loccu len tbo ttom depos its.
5. Pa rasi tes such aslampreys and hag fishes ,are ve ry d if
fe ren tfrom
othe rf in f
ishesinthe ir behav ior . They ob ta in nu trients by suck ing
bodyf luids o fhos tf ish.
 CHAPTER3 Feed
ing Hab
itsand D
iges
tivePhys
iologyo
fFishes 79
Ana
tomyand Phys
iologyo
fthe D
iges
tiveSys
tem

Inna tu re,the reisaw ideva r

ietyo ffoodava ilableon wh ichfishandc rustaceans
depend .F ishadap ttothei rfoodd if
fe rencesbyana tom icas wellasbehav ioral
means . Thus ,the re are many d i
ffe rencesinthe ana tomy and phys iologyo f
digestioninf ish.The reisas t
rongco r relat
ionbe tweentheana tom icalstructure
ofthe d iges t
ivet rac tandthefeed ing hab itso fthef ish. He rb
ivo rousfishtha t
depend onf ibrousfoods such as phy top lankton and mac rophy tes di
ffer
ana tom ically and behav ioral
lyf romca rn
ivo rousfishtha tconsume mea tand
othe r mo re diges t
iblefeeds . Ca rnivo rousf ishes have are lativelysimp le and
sho rt gut, w ithth ick mucosafo r abso rption. He rbivo rousf ishes have an
accesso ry mas ticatory appa ratus o ro therphys iologica l adaptationto helpin
breakingdownp lan tcell wallsbe forethed igest
ionp rocesss tarts, andalong ,
thingu ttoinc reasegu treten t
iont imeandenhanced iges t
ionandabso rpt
ion.

A
.Fishes
The digestive sys tem o ffishinc ludes the mou th, esophagus ,s tomach ,
pylorus,intestine,liver, and panc reas
. Ani llustrationo fthe digest
ivet ract
offourcommon lycu lturedfishestha tdif
ferinthei rfoodp referencesisshown
inF igure3 .1
.Thed igest
ivet ractistubularins truc ture
. The who led igestive
tractis oftenre ferredto asthe gu t andinf ish,the gu t usually hasfou r
div
isions:thesea retheheadgu t
,fo regut
, m idgut
, andh indgu t.Theheadgu t
,
whichisthe mos t anterior partincludesthe mou th(oralo r buccalcav ity)

Figure 3
.1
Diagramma ticrep
resen ta
tion ofthe d
igest
ive systems o
ffourfishes
,arrangedin o
rdero
finc
reasing gutleng
th.a)
Rainbowt rout(ca
rnivore)
;b ) Catf
ish(omnivore, ea
ting mo
re of an
imal sou
rce
);c) Ca
rp(omnivo
re, eat
ing more o
f
p
lantsou rce);d)Milk
fish(omn ivore,mic
rophagus plankt
ivo
re)
.
Sou
rce
: Sm
i
th1989
80 CHAPTER3 NUTR
ITIONINTROP
ICALAQUACULTURE

andg i
lls(b ranchialo rpharyngea lcav
ity). Theforegutbeg insa tthepos te r
ior
edge ofthe g i
lls andinc ludesthe esophagus and s tomach . The midgut
consists ofthein testines and pylor
ic caeca ,ifp resent. The m idgutisthe
longes t port
ion o fthe gu t and may be co iledin to comp licatedloops. The
hindgut inc ludestheen larged port
ion o fthein testines andtherec tum o r
anus. Each po rt
iono fthegu t hasaveryva riables tructurefo r adap
ta tion.
Theliverandpanc reasa reo rgansinvolvedind igestionbu ta refoundou tside
thetubu lars t
ruc ture.

1
. Headgu
t
Mou
thandva
riousinges
tion mechan
isms
Thef irst phase ofd iges tionistheinges tion o ffoodin tothe mou th .
The mou th has a var iety o f adap tationsfor cap tu r
ing , hand ling, and
so rting o ffood be fore en tryin tothe s tomach .F igu re 3.2A showsthe
diffe ren t shapes o f mou thinresponsetothe irfood adap tations .F ish
havetee ththa tvaryintype , numbe r, and a rrangemen t
. They se rveto
catchandho ldthep rey. Thea rrangemen tands tructureo fthetee tha re
rela tedtothe k ind offoodtha tthef ish no rma l
lyea t. Thereis as trong
co rre lation amongk indo ftee th,feed ing hab its, andfoodea ten .
Gene ra l
ly,the mo reac tivefeede rs haves trongjaws w ithsha rptee th
tob iteandsh redthefood . Some ma jo rk indso fjaw -teetha rethefo l
low ing:
ca rd iform ,v i
lli
fo rm,can ine ,inc isorand mo lariform(F igure3 .2B ).Those
feed ing on mo l
lusks and c rustaceans have sho rt heavytee th,s trong
enough to crush the mo llusk she ll. Zoop lank tonfeeders and mos t
plank t
ivo res have p ractica l
ly notee th. The shredd ing offoodis mos t
oftendoneinthethroa to rpha rynx . He re,ano the rse to fspecializedtee th
may befound . Again,the s t ructure,s i ze and shape o fthe pha ryngea l
tee th a re alsova r
iable.P lank tonfeede rs havef inerows o f pha ryngea l
tee th , wh ile mo lluskea ters havela rge bu tfla tc rownedtee th , wh ichis
bet te radap tedtoc rushingthe irfood .

F
igure 3
.2A F
igure 3
.2B
Va
ria
tionso
fthe mou
ths
truc
tureinf
ishes
. Some ma
jork
indso
fjawtee
th.
 CHAPTER3 Feed
ing Hab
itsand D
iges
tive Phys
iologyo
fFishes 81
2
. Fo
regu
t
Esophagus
Mos tfish havesho rt,wideesophagustha tse rves asat ransitionalarea
betweenthes tr
iated musc les ofthe mou th andthesmoo th muscles o f
thegu t. Mucusp roducingce llsa represen tintheesophagus . Ingene ral,
theesophagus se rves only as a passage way , however, enzyme ac tivity
has been de tectedinthe esophagus o fsomef ishesind icating a mo re
activero leo ftheesophagusinthe d igestion processinthesespec ies.
Othe rfishes withlong ,s l
im bodyshapesl iketheseawa te r
-adapted
eels have along esophagus . Osmoregu la
tion maytake p laceinthe
esophagusi f mucusis p resent. The mucusis muchth icker anter
iorly
thanpos ter
iorlyandise lectr
icallycha rged. Somerepo rts havesugges ted
tha t both pass ive and ac t
ivet ranspo rto fionsin tothe blood may a lso
takep laceintheesophagus w ithou tadditiono fwatersuchasthed i
lution
ofinges tedseawa terinf reshwa te reels.

S
tomach
Thestomachso ffishesva ryg reat
lyinthe irana tomicals
tructure dueto
adapta
tionstospec ificfoods. The re arefourgene ralconf
igurat
ions or
shapeso ffishstomachs . Theseinc lude:
a
. as tra
ights tomach w ith anenla rged po
rtion
b
. aU -o rJ-shapeds tomach
c
. as tomach shapedl ike a Y oni ts s
ide wherethe stemfacesthe
cauda lportion
d
. s tomachlessf ish, suchasinca rpsando the
rcyp r
inids

The s tomach has a con f

igu ra tion o r shape wh ichis conven ientfo r
conta iningfoodintheshapein wh ichi tisinges ted. Foodistempo rarily
storedinthes tomach wh i
letheres tisg radua llybeingp rocessedth rough
theo the rportiono fthed igestivet rac t
. Thes izeo rcapac ityofthes tomach
inre la t
iontothebody we igh tva riesbe tweenspec iesand
is usua l
lyrela tedtothein terva lbe tweenfeed ingsandto
thes i zeoffoodpa rt
icles. Gene rally ,f ishtha tea trelatively
sma ll,so f
tpa rticleshavesma lls tomach whe reasfishtha t
eatla rgefoodpa rt
icles,e .g. wholef ish ,o reata tin f
requen t
inte rvals havela rgers tomach .
The cecum o fthe Y -shaped s tomachis adap tedto
stretch pos te riorlytoaccommoda tela rgefoodpa rt
icles
orp rey . Incon trast,theabsenceo fas tomach hasbeen
sugges ted to bene f
itf ish adap ted tof reshwa ter( low
chlo ride concen tration) whe re s tomach ac idsimpose
added osmoregu latory pressure . Th is is to avo id
acid ifying large amoun ts o fa lka line food , as in
omn ivo rousf ishtha tea tp lant sou rces ,co rals,she lls,
and o thers.
In m i
lkf
ish ,thes tomachcanbed ividedin toca rd iac
and py loric po rt
ion . Theca rd iac po rtioniso ften mo re
Figure 3
.3
enla rged wh ilethe py lorics tomachis h ighly muscu lar Regions ofthe d
iges
tivetrac
to fmilk
fish Chanos chanos
. EP -
(Figu re3 .3
).Thepy lo r
ics tomachin tenselyg rindthefood epibranchial organ; E-esophagus; CS-cardiac stomach; PS-
pylor
ic stomach
; PC-pylo
ric caeca
;I-intes
tine
;L-l
iver
; GB -ga
l
l
particlesresu ltinginchyme(apas tel ike mass ). bladder
;P -panc
reas.
82 CHAPTER3 NUTR
ITIONINTROP
ICALAQUACULTURE

Thecapacityorvolumeofthestomachinrelat
iontothebody we
igh t
var
iesbe tweenspec
iesandref
lectsthesizeo
fthe mea
lthatcanbetaken
vo
lun ta
rily
.Itcanvaryfromassma llas 10%ofbody we
igh
tandasla rge
as 50%o fbody we
ightinasinglefeed
ing.

3
. M
idgu
t
The d igest
ion process ac tively con tinuesin to thein tes t
ines a fter
preliminary diges t
ioninthe s tomach . A l
lf ishes havein testines. The
leng thofthein testinesvariesf rom aslowas 1/5toas h ighas20t imes
the bodyleng th .Insomef ish ,thein tes
tines maybesho rtands traight
whileitcanbelong ,folded
,andloopedino thers. Ingene ral, herbivores
havelonge rin testinesthanca rn ivores. Althoughthe re aresomecases
ofove r
laps
,somegene rals
ta temen tscanbe madeongu tleng thinre lat
ion
tofeed ing hab i
ts o ffish(Table 3 .2
). W ithinthe samef ish spec ies,the
relativegutleng thcanchangeasfeed inghab itofthefishchanges . Gu t
leng this d
irec t
ly mo rere
latedtotheamoun tofindigestible ma terialin

le 3
Tab .2 Gene
ralobse
rvationsonfeed
inghab
itsandre
lat
ivegu
tleng
ths(
rat
ioo
fin
tes
tine
to bodyleng
th)infish
Feed
ing hab
its Re
lat
ive gu
tleng
th

Ca
rnivo
res 0
.2-2
.5

Omn
ivo
res 0
.6-8
.0

He
rbivo
res 0.8-20
.0

Feed
inghab
itandre
lat
ivegu
tleng
thinsomef
ishes
Spec
ies Food o
rfeed
ing hab
it

Gob
io gob
io Inve
rteb
rates 0
.80

Che
leth
iops e
longa
tus Zoop
lank
tons 0
.75

E
lop
ich
thys bambusa Ca
rnivo
res 0
.63

Bo
ril
ius meo
rei Ca
rnivo
res 0
.65
-0.80

Ca
tla ca
tla P
lan
ts,insec
tla
rvae 4
.7

Ga
rra dembens
is A
lgae
,inve
rteb
rates 4
.5

C
irrh
ina mr
igha
la A
lgae
, de
tri
tus 8
.0

Gadus mo
rhua Ca
rnivo
res 1
.05- 1
.50

Labeoca
lbasu He
rbivo
res 3
.75- 10
.0

Labeo ho
rie De
tri
tivo
res 15.0-21
.0

L
.ni
lot
icus A
lgae
, de
tri
tus 16
.9

L
.va
riega
tus A
lgae
, de
tri
tus 16
.9

L
.linea
tus A
lgae
, de
tri
tus 16
.1

C
tenopha
ryngodonide
lla P
lan
ts 2
.5

Do
ras g
rypus P
lan
ts 2
.8

Hypoph
tha
lmich
thys mo
lot
rix He
rbivo
res 4.6-7
.1

M
icrop
terussa
lmo
ides Ca
rnivo
res 0
.7-0
.9

Sa
lmosa
lar Ca
rnivo
res 0
.73-0
.80
 CHAPTER3 Feed
ing Hab
itsand D
iges
tive Phys
iologyo
fFishes 83
thefoodra the rthan whe the rthefood sou rceis o fp lan to r
an ima lo r
igin.F ishtha tinges tla rgeamoun tso fde tr
itus have
gu tleng thss im ilartothoseo fhe rbivo res.
Somef ishes possess py loric caeca . The re a re con f
lic ting
repor ts on the func tions o fthe py lor ic caeca in f ish .
Histo log ically,thepy lo r
iccaecaresemb lesthein tes tines. Mos t
stud iesind ica tetha ti tse rvesasanex tens iono fthein tes tines
thusinc reasingthe e ffec tive sur face a reafo rd igestion and
abso rp t
ion . O the rs tud ies showtha ti t ac ts as an accesso ry
food rese rvoir,fortemporary s to rage, poss ibly a dev isefo r
sav ing space . I tis c lea r, howeve r
,tha tra inbowt rou t caeca
takes upam inoac idsandsuga rsac rosstheap ica l memb rane
ofthe ep ithelial ce lls. E lec tron m icroscopy has showntha t
bothin tes t
ina landcaeca lce llsa reinvo lvedinl ipidabso rp tion ,
withcaeca lce llsbe ing mo re ac tive .
Thes truc tureo ftheabso rptivece llo fthein tes tinesre flects
its spec ializedfunc tionin d iges tion(F igure 3 .4 ). The ce l
l
con ta ins many m itochondr ia , wh ich prov ide energyfor
Figure 3.4
metabo lic p rocesses ; endop lasm icre t
icu lum , whe re pro teins
Schema ticrepresenta
tionofapor
tionofanabso rp
tive
(inc lud ingd iges tiveenzymes )a reassemb led ,andgo lgibod ies ce l
lfromthein test
ine, Mv-mic
rov
i
ll
i,M-mitochondr
ion,
whe reca rbohyd rates idecha insa rea ttachedtop ro teins.T igh t Ser- smoo th endoplasm icret
iculum, Re r
-rough
endop lasmicreticu
lum .
junc tions and desmosomes b indthe abso rptive ce llsin to a
sing le shee t onthe sur face o fthe mucosa . The nuc leusl ies
deepinthece l
l. The mos ts trikingfea tu reo fthece llisthep resenceo fa
brush bo rder, aprom inen ts truc tureonthesu rfacefac ingthelumeno f
thein tes t
ine. Theb rushbo rde riscomposedo fm inu tep rojec t
ionsca lled
microv illi. Onthe memb raneo fthe m icrov illusa refoundseve ralkinds
ofd iges tiveenzymes andt ranspo rtp ro teins .

4
. H
indgu
t
The hindgutisanex tensiono fthe midgu
t. D igest
ion hasbeenshownto
cont
inueinthe h indgu talthough w i
thag radua l
lydim in
ishingdiges
tive
orabso rpt
ivefunction, aninc reasedsecret
iono f mucus and apH nea r
neutral
. H isto
log
ical sections show a sudden changef rom co
lumna r
sec
re toryandabsorptivetoasquamousep i
theliumthatp roduces mucus.

5
. L
iver
Thel iverisanimpo rtant me tabo lico rgan .Itaidsind igestionbysec re ting
bi
le ,ag reen ishfluid wi
ths trongemu lsi
fyingp ropert
ies. Theb ileiss to red
inthega llb ladde randiscomposedo fa m ixtureofbilesalts,tau rocho late,
glycocho late . Bileacidsa rede rivedf rom me tabolismo fcho lesterol,and
deg rada t
ionp roduc tsofhemog lob in,b i
lirubinandb i
liverdin. B ilese rves
toemu lsi
fyl ipidsinthegu tand maycon tainothe r wastep roduc ts. The
bile duc t opensin tothe an terio rin testines orintothe py loric caecai f
presen t. F ishcanreabso rb b ileinthe h indgu teventhough mos tl ipid
up takeoccursinthean terio rin testine.Thel iverisa lsoas to rageo rgan
fo rlipids andg lycogen ors to reds ta rch. Insomef ishes,la rgeamoun ts
oflipidiss toredinthel iverto he lp ma inta
inbuoyancy . Ino the rfishes ,
glycogenisthe ma jorstored nu t r
ien t.
84 CHAPTER3 NUTR
ITIONINTROP
ICALAQUACULTURE

6
. Panc
reas
The panc reasisinvo lvedin manyimpor tantfunc t
ionsin d iges t
ion.
Panc reatic mo rpho logyisva r
iablein manybonyf ishes . In mos tf ishes,
unlikeinland an imals ,the reis no d iscrete panc reas . The panc reasis
dif
fused , sca t
te red, andembeddedinthe mesen te ries,inthel iver, and
clustereda roundtheb i
leduc t,orincomb ina tionso fsites. Inad if fused
panc reas, seve ral sma l
l duc ts openin tothein tes tine andthe py lor
ic
caeca . In othe rcases whe rethe panc reasisfoundins idethel ive r,the
panc reasde liversitssec ret
ionsd irectlyintothega llbladde r
. Thepanc reas
producesinsu lin and d igestive secretions, princ ipally pro teases and
bicarbona tes.Insu l
ins t
imu latesup takeo fam inoac idsf romthein tes t
ine
and mays t
imu lategrow th.Intheno rthernp ike,insu l
indec reasesb lood
am ino ac ids w ith up takein to skeletal musc les, wh ilein cod ,insu l
in
decreasesb loodg lucose .

B
.Crus
taceans
Thec rus taceand iges t
ivet rac tis mos tlys t raigh tandcons istso fthefo regu t,
the m idgu t
, andthe h indgu t(Figu re 3 .5 )
. Thefo regu tandthe h indgu ta re
lined w ithch itin . The m idgu ta risesf romtheendode rmandtheinne rmos t
laye rad jacen ttothelumenisthe mucosal ined w ithep i
the lialce lls wh ichis
composed o fs imp le co lumna r cel
ls. These a re suppo rted by a basemen t
memb rane , alaye ro fc i
rcu lar musc les andthen by alaye ro flong i
tud ina l
musc les . These musc lelaye rsa resu rroundedbyalaye ro ffib rousconnec tive
tissueo rse rosaandcon taina ne two rko fa r
te rial hemo lymphvesse ls.
Inthe an te rio r po rtion o fthe m idgu t ep ithelium a re s imp le co lumna r
cells w ith med ial nuc lei. The su r
face ad jacen ttothelumen a lsocon ta ins
mic rovill
i. Many m itochond ria, sec reto ry g ranu les and go lgi bod ies a re
presen tinep ithe lialce l
ls . Incon tras t,the pos terior po rt
ion o fthe m idgu t
epi the l
ium con tains manysquamousce lls wh ich sec rete mucus . The ce ll
sur facefac ingthelumena lsocon ta ins m ic rovilli
.
Ina lmos ta llc rus taceans ,the m idgu tis made upo foneo r mo repa irso f
glandu la r appendages ca l
ledthe hepatopancreas . The hepa topanc reas
con tains a w iderange o fd iges tiveenzymes
which hyd rolyzefood nu t
rien ts and a idin
break ing down bythe gas tric m i
ll. The
hepa topanc reasiscomposedo fs imp leb l
ind -
end ingtubu leso rdiverticu la wh ichopenin to
seconda rysec retion duc ts. These duc ts,in
turn , openin to the pr imary secre tion o r
collec tingduc t,through wh ichthesec re tion
is poured in to the m idgu t
, beh ind the
stomach . Thelumeno fthehepa topanc rea tic
tubu lecon tainsg ranu lar ma te rialandce l
ls
lining the mucosa are covered w ith a
microv illus b rush bo rde r
. Thetubu les a re
lined w ith anep itheliumin wh ich d ifferen t
celltypesa rep resent.Theapexo fthetubu le
Figure 3.5 con tainund if
fe rentiatedemb ryon ico rE -ce lls
D
iag ram o fthe d iges
tivesys
tem ofshr
imp with g
il
ls and muscula
tureremovedto Farthe r awayf romthe apex , ce lls beg into
show ma joro rgansystems.FA-fo
regu
t(ante
rio
rchambe r)
;AM -an
te r
iormidgutcecum;
FP-Fo regu t(posterio
rchamber);H-hea
rt
;O -ovary
; HP-hepatopancreas;M -midgu
t; di
fferen t
ia tein todeve lopingabso rptive,s to rage
PM -poster
io rm idgutcecum;A-anus orR -ce lls. Inthe p rox ima l reg ion o fthe
 CHAPTER3 Feed
ing Hab
itsand D
iges
tive Phys
iologyo
fFishes 85
hepatopanc rea t
ictubu le,inadd i
tiontothe Eand Rce l
ltypes ,a refoundla rge
dist
inct
ivesec retoryo rB-ce lls (F igu re3 .6).
Thecon ten tso fthegu to fc rustaceans a re movedfo rwa rd by pe r
istaltic
orcon tractile movemen tso fthelong itud inal musc les. Pe rista
lsisoccu rsin
theesophagus ,m idgu t,andh indgu t. Abso rpt
ionoccu rsinthean ter
io rpa r
t
ofthe m idgu tand d igestioninthe pos teriorpa r
t. The refore,itis necessa ry
thats trong an t
i-pe ristaltic movemen ts a lsotake p laceinthe m idgu tfrom
the pos teriorendtothean teriorpo r t
ion . In decapods ,if mand iblesa rmed
with sha rptee thfor chew ingfood a re absen t
,the reis agas tric m i
llw ith
movabletee thf ixedtop la telikeoss iclestha tserveassubs t
itutefo rg r
ind ing
thefood . Thegas tric m il
lisanadap ta t
iontoaseden taryex istence .Ita l
lows
crustaceansto swa llowthe i
rfoodf irst,then chew a tle isure wh ile hidden
from othe ran ima ls.
Thed igestiveju iceisproducedch ieflybythece l
lso fthehepa topanc reas
andt ranspo rtedtothes tomach . Insomespec ies, thepHisabou t5du ring
hunge randr isesto pH 6 .6onfeed ing.Ino thersspec ies, the pHis neu tral

F
igure 3
.6
Schemeo fdi
ffe
rentia
tion andfunct
ionofthedigest
iveglandtubule
. The E
-ce
llsdi
ffe
ren
tia
teinto F-ce
lls(
fibr
il
lar
)orR-cel
ls(s
torage
).
F-ce
llssec
rete enzymes andd i
ffe
rent
iatein
to B-ce
lls
. B-ce
lls absorbsma
llpar
tic
lesbypinocytos
is. R-cel
lsabsorb nu
tr
ien
tsands to
re
l
ipids,g
lycogen,coppe r
,phospho rusandothersubstances
.
Sou
rce
: Da
ll
, 1992
86 CHAPTER3 NUTR
ITIONINTROP
ICALAQUACULTURE

inthe hung ryanima lsandincreasestopH 7 .5af

terfeeding
. The d
igestive
juice contains various d igest
ive enzymes and b i
le sal
ts that actin
emu ls
ifyingfats
.
Absorpt
ion occurs ma inlyinthe hepa topancreas andinthe anter
ior
partofthe midgut. The R-cellsofthe hepa
topancreas havethecapacityto
absorb nutr
ients
.

D
iges
tionand Abso
rpt
ion

Digestionis a p rocess whe rebyinges tedfood nu trien ts such as p roteins ,

lipidsandcomp lexca rbohyd ratesa reb rokendownin to un itstha ta resma l
l
enoughto be abso rbed ac rossthe gu t wa ll. The p rocessis accomp lished
throughthe ac tion o fd iges tive enzymes . The ab il
i ty offishto d iges tfeed
dependsonthesec retiono fadequa tequan tit
ieso ftheapp rop r
ia tetypeso f
enzymes . Manyo ftheenzymesa res toredinaninac tiveo rproenzymefo rm .
Oncesec re tedin toafavo rab leenv i ronmen tfo rd igestion , usua l
lyin fluenced
by pH ,theseinac tive enzymes a re conve rtedtothe ac t
ivefo rm readyto
performthe irspec i
ficd iges tivefunc tion. Whenva r
iousnu trients havebeen
adequa telyd iges ted ,theya rethenabso rbed p r
ima rilyinthe m idgu t.
There a re seve ral mechan isms o f nu trien t abso rptioninf ishes -simp le
diffusion , activet ranspor t, and p inocytos is. Ins imp led if
fus ion ,so lutes
passthroughthe memb ranef rom an env ironmen to fh ightolow so lu te
concen tration w ithou t us ing ene rgy. Ac tivet ranspo rtd iffersf rom s imp le
dif
fusionasi trequ iresacon tinuoussupp lyo fene rgyandt ranspo rtsso lu tes
onlyinoned irec tionf romlowtoh ighso lu teconcen tration. Aca rrie rsys tem
tha tu t
ilizes Na+and ATPase ac t
ivityis neededinthe ac tivet ranspo rto f
glucoseandsomeam inoac ids . Pinocy tosis(ce lld rink ing)istheprocessby
which ma ter
ia ls are takenin to the ce llthrough aninvag ina tion and
subsequen tdisso lu t
iono fapa rto fthece ll memb rane .Th isp rocessenab les
thece lltoabso rbsome p ro teinsandl ip idsinin tac tfo rm . Abso rptivece lls
ofthe m idgu ta recapab leo funde rgoing p inocy tosis.

A
.Diges
tionandabso
rpt
iono
fpro
tein
s

Prote
insarehydrolyzedintoaminoac
idso
rpolypept
idecha
inso fafewamino
acids w
iththehelpo fenzymesknownasproteases
. Theprocesscanoccur
inac id
icorbasic pH.

1
. Peps
in
Peps inandhyd roch
lo r
icac idp layimpo rtantro
lesinpro te
ind igest
ionin
fishs tomachs . The pHop timumfo rpepsinisabou t pH 2.0. Pepsinis
syn thes
izedinthegas tricg landintheinac tiveformca l
ledpeps inogen .
Hyd rochloricacidconve rtstheinac tivepepsinogentotheac t
ive pepsin.
Peps inis anendopep t
idase and c leaves orcuts most peptide bondsin
thein teriorregion particularly whe rel
inkages areformed by a roma t
ic
am inoac ids(phenylalanine,ty ros
ine ,andtryptophan)
,andac idicam ino
acids(aspa r
ticandglu tam ic acid
).
 CHAPTER 3 Feed
ing Hab
itsand D
iges
tive Phys
iologyo
fFishes 87
2
.Tryp
sinandChymo
tryp
sin
Tryps in and chymo tryps in areinvo lvedinthe a lka line d igestion o f
proteins . These enzymes a re syn thes ized ,s to
red and sec retedin an
inac tiveform bythe panc reas andt ranspo r
tedtothe m idgu t andthe
pyloriccaeca . Tryps inogen isac tiva tedinthein testinebyen terok inase,
an enzyme secre tedf rom thein tes tina l mucosa . When ac tiva ted,
tryps inogen becomes tryps in. In turn , tryps in , act ivates
chymotryps inogen tochymotryps in . Bo tht ryps in andchymo tryps in
are endopep t
idases bu tcleave d i
ffe ren tl inkagesin a p rotein. Tryps in
cleavespep tidel inkages wh icha refo rmedbybas icam inoac ids,a rginine,
lysine ,andh istidine. Chymo trypsinc leavesl inkages w itha roma t
icam ino
acids, pheny lalanine,ty rosine , andt ryp tophan .
In m ilkfish, bo tht rypt
ic and chymo tryp t
ic ac t
ivities a re higherinthe
posteriorin tes t
inesthaninthean teriorpa r t
.Thepy loriccaecaa lsocon tains
highac t
ivitieso fbothenzymes .M ilkfishin testinalp ro teaseac t
ivityappea r
to havetwo pH op t
ima , one a t pH 7 .0to 7 .6 and ano the ra t pH 9 .5to
10.0.I ta lso hasatempe ra tureop timumo fabou t 50 °to60 °C.

PEPS
INOGEN TRYPS
INOGEN CHYMOTRYPS
INOGEN
↓ HC
l ↓ en
terok
inase ↓t
ryps
in
PEPS
IN TRYPS
IN CHYMOTRYPS
IN

Conve
rsiono
f peps
in,t
ryps
in,andchymo
tryps
inf
romthe
irinac
tivefo
rms

3
. Ca
rboxypep
tida
ses
Carboxypeptidasesa rea
lsosecre tedfromthepanc reasintheinactiveform.
Theseareexopep tidaseswhichc leavethe C-terminalam inoac ido
fpep t
ides
or proteins . There are two types : carboxypep tidase A and B .
Carboxypep tidase Aisactivetowa rds pro
teins w
i th aroma t
ic C
-terminal
am ino ac ids (pheny lalanine , tyrosine, and tryp tophan) wh i
le
carboxypep tidase Bactspreferen t
iallyonthosepep tides wi
thbas icamino
ac
ids(lysineanda rg
inine
).

4
. Am
inopep
tida
ses
Am inopep t
idases a re exopep tidasestha t act on N -
te rm inal peptide o f
proteins .
In summa ry, because o fthe comp lexity ofproteins ,the ircomp lete
digestion hasto p roceedin a numbe ro fs teps(Figu re 3.7). The protein
molecu leisf irst hydrolyzedin tore lat
ivelyla rge polypep t
idef ragmen ts
by endopep tidases. Thesef ragmen ts arethen hyd ro lyzed by enzymes
actingontheam inoandca rboxylbondsandf inal
lyd ipep t
idasesreduce
thepro teintoi tscons tituentam inoac ids.
Proteinscanbe abso rbed as who le proteins, peptides ,o rfreeam ino
acids.P rotein mac romo lecules may be abso rbedinthegu tepithelium
via pinocy tosis
. Ra inbowt routhaveg ranu lecellsinthelam inap ropriao f
thein test
inejus t unde rthe mucosa wh ich cou ld be pa rto fthis uptake
system .
88 CHAPTER3 NUTR
ITIONINTROP
ICALAQUACULTURE

For comp letely d igested

proteins, each am ino ac id hasi ts
own up take charac teristics. In
rainbowt rou t
,L -
leuc ine up takeis
shown to be spec ific for the L -
isomer , ac tive and sod ium-
dependen t andcou ld beinh ibited
byo the rneu tralam inoac idssuch
as L-va l
ine and L-me thion ine.
Lysine up takeis s lowed down by
the presence o fg lucose and
has tened by ATP . Some am ino
acids may betaken up by s imple
diffus ion; o thers may be
transpor ted by carr iers in a
sodium -
independen tsys tem wh ich
F
igure 3
.7
Sequenceofp ro
teindigest
ion.Theprote
in mo
lecu
leisf
irs
thyd
rolyzedin
topo lypep
tides usua lly occurin the baso lateral
byendopeptidases. Po
lypept
idesarefur
the
rhydro
lyzedtosma
llerdipep
tidesandfinal
ly memb rane.
in
to const
ituentam inoacids
.

B
.Diges
tionandabso
rpt
iono
fca
rbohyd
rates

Carbohydrases d iges
t comp lex carbohydra tes and a refoundinf ish
intestines. They a reve ryimpo rtan tin he rbivo rousf ish because o fthe
highlevelso fca rbohydra teinp lan ts. Theenzymeamy laseca talysesthe
diges
tiono fs tarch.The rea retwofo rms :α -amy lasewh ichac tsrandom ly
cleav
ingthecha infrom w i
thin and β -amy lase wh ichcu tsthecha in at
every two g lucose un its.A t the branched po int, ano ther enzyme ,
dextrinasedoesthe wo rk .Theamy laseanddex t
rinasep roduce ma ltose.
Ma ltase hyd rolyses ma ltosetog iveg lucose ,
thef ina lp roduc to fsta rchd iges tion(F igu re
3.8). Mos tfish haveamy lase;in he rb ivo rous
fish , suchast ilapia,i t maybep resen tina l
l
par ts o fthe d iges t
ive t rac t, whereasin
ca rn ivo rousf ishi t maybefoundon lyinthe
panc reas , pylo riccaeca andin testines . In
milkf ish (an omn ivo re), ex trac tsf romthe
in tes t
ines , panc reas , py loriccecaandl iver
showed h ighleve ls ofamy lase ac t
ivi ty. In
add ition , maltose ,t reha lose ,dex t
rin,s tarch ,
andg lycogen a rerap idly hyd rolyzedinthe
presence o f crude extracts from the
in tes tinesand py loriccaecao fm i
lk fish .
Othe r comp lex ca rbohyd ratestha ta re
po ten tia l sources o f energy bu ta re no t
read ilyd igesteda rece llulose ,acons tituen t
ofp lan tce l
l wa ll, andch it
in , acomponen t
of crus tacean exoske leton . The comp le
te
d
iges tion o f cellu loseis med ia ted bytwo
Figure 3
.8
e nz ym e s: ce llu la se a nd c ellob iase.
Represen ta
tiono
fdiges
tionandabso
rpt
ionofca
rbohyd
rates. C ell ul a se h yd rol ys es c e
l lulo s e to
 CHAPTER 3 Feed
ing Hab
itsand D
iges
tive Phys
iologyo
fFishes 89
disaccha r
idecellobiose, whichisthenac teduponbyce llob
iasep roducing
thef ina
lb reakdownp roduc t,glucose. Veryfewfishhavece l
lulaseactiv
ity
,
mos tlikely thein tes t
inal m icroflora active
ly prov ide the cellu
lase.
Cellulyt
ic bacteriaa re widely dis
tributedin natu re.
Carbohydrate abso rption has beentes ted by measur ingthe uptake
ofglucose,theusua lfinalendp roductofca rbohyd ratedigest
ion. Glucose
transpo rtappea rstobelowes tinca rnivores(e.g
.ca t
fish)and highestin
herbivores(e.g.ca rp).

C
.Diges
tionandabso
rpt
iono
flip
ids

Lipases hydro lyze es terl inkagesint riglycerides . The end p roduc ts o f

lipase ac t ivi ty a re g lyce rol andfa tty ac ids.L ipases a re de tec tedinthe
panc reas , py lo r
icceca ,in tes tine , andl iver.In m ilk f
ish ,lipase has a lso
beende tec tedintheesophagus .M ilk fishin tes
tina landpanc rea t
iclipases
appeartohavetwopHop tima -pH 6 .8to
8.0;andpH6 .4to8 .6,respec t ively . The
detec t
iono ftwo we l
l -defined pHop t
ima ,
one a ts ligh tly ac idic andthe o the ra t
alkaline pHfo r bo ththein tes tina l and
pancreat ic l ipases sugges ts a
phys iolog ica lve rsa tilityfo rlip idd iges tion
in mi lk f ish . Phosphol ipases are
enzymestha t hydro lyze phospho lipids .
Lip ids a retaken up bythein tes tina l
and ceca l ep ithe lium , par tly asfa t
ty
acids , mos tly as mono -g lycer ides and
partly as d rop lets, andt rans fer redin to
bloodandlymphvesse ls(F igu re3 .9). In
thein tes tina l ce ll,fa tty ac ids are re­
ester ified to tr ig lycer ides and are
transpor ted as verylow dens i tyl ipid
part
ic lesin tolymphvesse ls .Intheb lood ,
triglycer ides are transpor ted as
chylom icra . Thel ipidabso rp tionp rocess
inf ish ,a lthough much s lowe rthanin Figure 3
.9
D
iag ramma t
icrep
resenta
tiono
fdigest
ionandabso
rpt
iono
flip
ids
.
mamma ls,doesno td i
fferfundamen tally.

Measu
remen
tsand Ana
lys
isusedin D
iges
tionS
tud
ies

A
. Measu
remen
tofS
tomach Con
ten
ts

Analysisand measuremen tofstomachcon ten

tscang iveinformationonthe
feedingprefe
renceso fapart
icularspeciesas we
llasitsfrequencyo ffeeding
.
Several me
thodsare usedto measurestomachcon ten
ts, however
,a l
lo fthese
methods havethei
rlim i
tat
ions.

1
. The occurrence o
fa par t
iculartype offoodinthefish s
tomach can be
determined qual
itat
ively asin web s
tud ies bu
tthe quant
ityo
fthefood
presentcanno tbeob ta
ined bythis method.
90 CHAPTER3 NUTR
ITIONINTROP
ICALAQUACULTURE

2
. Eachk indo find iv idua lfoodo rgan isminthes tomachcanbecoun tedto
showthecho iceo fthef ishfo rce rtainfoodtypesf romagene ralpopu lation
ofpo ten tialfoodo rganisms . The me thodcanno tbe usedincases whe re
thefoodischewedo rwhenthefoodisno tind istinctun i
tsas w ithde tritus.
3
. The useo fs tomachf lush ing me thods measu resthed isplacemen tvolume
ofafoodtypeo ro ftheto talfoodvo lumeandis mos tapp licab leifthefood
isinl iqu idfo rm .
4
. Measuremen to fthe we t weigh to rthed ry weigh to ffoodasape rcen tage
ofthebodywe igh tisano therp rocedu refo rde term inings tomachcon ten ts.
This me thod may g iveinaccura te resu lts when heavyi tems such as
molluskshe lls, sando r muda re presen t
.
5
. Es tima te o ffu llness - ha lf
, th ird, quar ters ,e tc.-o fthe s tomach by
appea ranceisaninexac t me thod bu t useful unde rp racticalcond itions .
Gas tr icemp tyingra teandfood passagera te maybe de term inedina
numbe ro f ways . Th is can be done by k il
ling o r no tk i
llingthef ishto
ob tains tomachcon tents . In manycases ,ace rtain numbe roff ishisfed
a known amoun to ffood . Samp les a re ob tained pe riodica l
ly by se rial
slaugh te rtode term inegas tricemp tyingra te
. Onel im i
tationo fth is me thod
istha ti t assumestha ta l
lthef ish a te exac tlythe same amoun t. An
alterna t ive p rocedureistofeed anind ividua lfish a known amoun to f
food ,andthena fterace rta
inpe riods tomachcon ten tsa reremovedf rom
thef ish unde r anes thes ia. The s tomach con ten ts may bef lushed ou t
with wa te r us ing avar iety o fgadge ts.O ther me thods o f measur ingthe
amoun to ffoodinthes tomacho rin tes t
ines w ithou tremov ingfoodinvo lve
feed ingf ishad ie tw i
tha ma rke rlikeba riumsu lfa teo riron powde rand
usinganx -raytov iewthelabe lledfooda tva rioust imes . Th isisuse fu lin
fishtha ta retoosma l
lfo ro the r me thods .

B
. Measu
remen
tofd
iges
tib
ili
ty

Thef
irststepinevalua
tingthepo tent
ialofafeedstu
ffforinc
lusioninad iet
isto de
termine o
r measureits d
igestib
il
ity
. Digest
ibi
litycan be measured
ei
therbyinv ivoorinvit
ro methods.

1
. In vivo me
thods
The mos tcommon me thodo f measur ing diges t
ibilityisto addtothe d ieta
marke r, such asch rom ic ox ideo ri ron powde rtha tis ne i
the rdigested no r
abso rbed. The d ietisfedtothef ishandtheconcen trat
iono fthe marke ris
followedthroughthe d iges tivet ract astheind iges t
ible componen ts a re
exc re
ted . Fecesf romthef ishisco llected byoneo fthefo llowing me thods :
netting,feca lsettling,ands t r
ippingo rd issec t
iono ftheh indmos tparto fthe
digestivet ract(F igu re 3.10 )
.
Sometechn ica l prob lems a risein each o fthese me thods.V igorous
stripping mayremovepa r tso fthed igestivet ract. Feca lco l
lectionisdifficult
ifthe exc retais so ft and canno t be co llected as a so lid. The amoun ttha t
leachesou tfromthefecesastheys tayinthetankbo ttomo rinaco llecting
deviceisqu ite difficultto de termine . The refore, ca reshou ld betakentha t
thebes tfeca lcollec tion me thodfo rapa rticularspec iesis used .
 CHAPTER3 Feed
ing Hab
itsand D
iges
tive Phys
iologyo
fFishes 9
1
Endogenous marke rs such as cellu
lose, hydrolys
is res
istan
t
o
rganic ma
tter
, and ac
id-insoluble ash mayalsobe used.
Theapparentpro
teindiges
tibi
li
ty(%APD )ofafeedisdef
inedas:

(%Fp
) (%D
i)
APD(%
) = 1- X x 100
(%Dp
) (%F
)

where:
Fp =proteininfecesorin tes
tinalcontent
Fi = indicato
rinfeceso rintest
inalcontent
Feca
lse
tt
ling
Dp= proteinin die
t
D
i = ind icatorinthe d
iet

True prote
in diges
tibi
lity(%TPD )o f afeedis obtained by
cor
rectingtheamoun tofprote
ininintestina
lcontentsorfeces w
ith
pro
teino fendogenousorig
in withthe useoftheformula:

(%Fp) (%D
.) (%Di
) (%Fc
)
p S
tr
ipp
ingo
rdissec
tiono
ftheh
indgu
t
TPD(%
) = 1- X + X x 100
) (%F
(% Dp ) ) (%Dp
(%Fc
i ) F
igure 3
.10
Exper
imenta
l se
t-upfo
r in
-vivo d
iges
tib
il
ity
measu
rement
.
whe
re:
Fcp= pro
teininfecesorinintes
tinalcontentof
contro
lfish(
fishfeda non-prote
in d
iet
)
Fci= indica
torinfecesorin
testinalcontentofcon
tro
lfish

2
. In vitro assays

Inestima t
ingp roteindigest
ibil
itybyinv i
troassays
,tes tra
tionsa reincubated
withintest
inalex tracts offish a
t an optimumtempe raturefo r aspeci
fied
lengtho ft
ime(e .g. 24h).Thep roteo
lyticactiv
ityintheintest
ina lextrac
tw il
l
diges
tthep rote
incomponen tofthediet
. The inv it
rodiges
tibi
lityiscompu ted
usingthefo rmu la:

(N
r- N
i)
Inv
itro prote
in d
igest
ibi
lity(%
)= x 100
N
r
whe
re:
N
r=p rote
ininra t
ion
N
i = ind
igest
ible p
rote
in

Thecondit
ions unde
r wh ich invi
trotechniquesarecarriedouta rehighly
unphysio
log
ic,thuscau tion mus tbeexe rc
isedinex t
rapolatingtheresults
ofsuchexperimen
tstotheinv ivocondit
ion.A tbest
,theva lueofdigest
ibi
lity
obta
ined us
ingthistechn iqueisonlyanes tima
teo ftrue d
igest
ibi
lity
.
92 CHAPTER 3 NUTR
ITIONINTROP
ICALAQUACULTURE
Fac
torsA
ffec
ting D
iges
tionandAbso
rpt
ion

Inf ish , asin a l

l an ima ls, bo th ex trinsic andin trins icfac tors can a lter
thee fficiency o fdiges t
ion and absorp t
ion . These a rethoseinvo lv ingthe
feedi tse lf,feed ing prac t
ices , as we ll asthe preva il
ing cond ition o fthe
diges tivet rac t.
Feed compos i
tion ,d iges t
ib i
lity, and prepara tionin fluence d iges tion
andabsorp tion .D ie taryf iberis h igh lyind iges tiblein non -herb ivores due
tothe absence o fthe enzymes necessaryto breakdownthe comp lexce l
l
wallsfoundin thefeed . Roughage a re h ighinf ibe r
, thus the ir useis
lim itedin prac t
icalfeedfor non -herb ivores .P lan t prote ins are knownto
vary in the ir d igest ib il
i ty in f ish because o fd iffer ing am ino ac id
compos ition andsecondary as we l
laster tiarys truc tures o fthe pro teins .
Inadd ition ,i facer ta intypeo ffeedis knownto pass qu ick lythroughthe
diges tive t rac to f an an ima l, thefeed w ill no t be adequa tely d iges ted
because o finadequa te exposuretothe d iges tiveenzymes .
Thep resenceo fan t
inu tri
tiona lfac torsinfeedsuchast ryps ininh ibito rs
and tann ins w illalso decrease the e ff
ic iency o f pro tein d iges tion and
absorp tion . Process ingthefeed such ascook ingsoybeanto des troythe
tryps ininh ib itoris necessary be fore thefeed can be u til
ized byf ish .
Dehu ll
ing orremova lo fthe seed coa tofsomelegumes w il
lincreasethe
diges tibilityo fthefeedbecause unbrokenseedcoa tsa re no teas ilyd iges ted
andinsomecasescon tain highleve ls oftann in. Fur thermore , excess ive
amoun ts o ffeed g iventothef ish decrease d iges tibi
lity .
The e fficiency o fd igestion and absorp tionis a lso a ffected by the
cond ition o fthe d iges t
ive t rac t, pancreas , and thel iverinf ishes or
hepa topancreas in crus taceans . Nutr itiona ld iseases and bac ter ial
infec tionso fthed iges tivesys temcand im in ishthed iges tiveandabso rp t
ive
func tionsinf ishes andc rus taceans .

Feed
ing P
rocessinF
ish

Thecyc
leo
ffeed
ing p
rocessinf
ishisshownin F
igu
re 3
.11
.

Appe
titeandSa
tia
tion
Appetiteisthes tatetha tinitia
tesa rousalandfeed ingbehavior. Know ledge
onthefac torstha tt riggerappe t
ite andsa tiat
ion areveryimpo rtan ttofish
farme rs who wan tto max im izefeedconsump tion,g rowth, andconve rs
ion
eff
iciencieso fthe irfishs tockbyad justingfeedingschemes . Appe t
ite, which
iscon trolled bythe hypo tha lamus ,iss timulated bythegu tfu l
lnessando r
othe r metabo licchanges wh icha ffectfoodconsump tion.Ingene ral
,s tomach
expans iona fte rfeed inginh ib i
tsappe ti
te andgas t
ricevacua tions t
imu la
tes
appe ti
te. The op tima lin terva l between mea ls has been es tima ted to
correspondtogas tricevacua tion.
Severalfac tors bo th biotic and ab iot
ictendto a ffectthe appetite ofthe
fish andsh rimp .

The bio
ticfacto
rs include:
1. Foodavailabi
lityandfoodd istr
ibu t
ion
, which maybeseasonalinna ture
;
2. Compe t
ition
. Mo re aggressivefishin astock a
ffec
tthe appet
ite and or
feedingandhencetheg rowtho fsubordinates
. Genera
lly
,sizedif
ferences
 CHAPTER 3 Feed
ing Hab
itsand D
iges
tive Phys
iologyo
fFishes 93
ofthe s tock and dom inance cause
inc reasedvar ia t
ionin bodys i
zes . To
avo id such occur rence , pe r
iod ic s ize
grad ing,toensuretha t nof ishis 1 .5
timesla rge rthan ano the r, shou ld be
done to prevent cann iba l
ism . In
add it
ion , increas ingthef requencyo f
feed ingo rbe tte rd ispe rsa lo ffeed may
helpthein fe r
io rfishtoob ta infood .The
presence o f compe t
ing spec ies may
en tai
l a sh iftto o the r“less des irab le
food ”;
3
. Thep resenceo fpreda tors mayinh ibit
feed ing;
4
. Phys iologica l cond i tion such as
starva tion o r mo t ivationleve l, and
circad ian rhy thm o fthe an ima l may
dictatefeed ingt ime and amoun to f
foodinges ted;
Figure 3.11
5
. Selec t
iono fprey
Feedingp rocessinfish.
a) Cho ice o f prey o rfood ea ten ,in Source: Mod i
fiedf
rom Knigh
t 1985

terms o fsize andfo rm ,isl im ited

bythe mou thgapeo fthef ish .
b) The“op t
ima lfo ragingtheory ”s tatestha tthe na tura lse lec tionfavo rs
thosef ishtha t max im izee fficiencyo fpreycap tu re. S inceene rgyis
spen tinsea rching ,pu rsu ing, hand lingandd iges tingp rey,anop tima l
foragings tra tegy mus tba lancethesefac to rssuchtha ttheto talene rgy
expend itureis m in im ized andthe ne tene rgyga inf romthecap tu red
preyis max im ized .
6
. Hand l
ingcausess t ressinf ish. Brownt routre fusetofeedfo rth reedays
afterhand ling wh ilera inbowt rou t,m ilkfish
,o rseabassresumefeed ing
the nex t day .

Theab ioticfac torsinc lude:

1. Disso lvedoxygen(DO )isoneo fthe mos timpo rtan tab io
ticfac to
rsa ffec ting
feed ingbehav ioroffishes
. Low DOleve lsleadtodec reasedfeed ingac t
ivity.
Milkfishandshr impsra isedinpondsceasefeed ing when DOleve lsd rop
below 1 .5and 3 .0 ppm ,respec t
ive ly.
2. In mos tf ish species, daily andseasona l wate rtempe raturef luctua t
ion
affectfoodin take. Fo rmilkfish, at ropicalspec ies, amb ien ttempe rature
does no ta ffect feedingac tivi
tyinponds , howeve r,ex tremetempe rature
fluc tua tionsa f
fectfeedingac tiv
i tyo fm ilk
fish. Sh rimpsa realsoa ffected
byex tremetempe raturef luctua tions .
3. Ligh tin tensity alsoin f
luencesfeed ing ac t
ivity of mos tf ish, espec ia l
ly
la rvalf ish, wh ichre lyonv is
iontorecogn izefood pa r
ticles.
4. Un ion izedammon ia(NH3 )levelso fabou t0.1ppminh ibitsg rowthin mos t
spec ies. Howeve r, appetite could beinh ibited a t ammon ialeve ls h igh
enoughtocausese rious phys iologicaland mo rpho logical damage .
94 CHAPTER 3 NUTR
ITIONINTROP
ICALAQUACULTURE

A
rousa
landsea
rch
Fish on afixedfeed ingschedulecanlea rnto antic
ipatefeedingt imes but
arousaliso f
ten duetoPav lov
ian conditioning
. Thes ightorsoundo ffa rm
workers abouttog ivefeedcans t
imu latefeeding. Sound(200 -700 Hz) has
been usedto cond itionred sea breamto cometo afeed ing spo tin a sea
ranchingsys tem. Theinfluenceofsocialinteract
ionamongf ishinas tockis
alsoimpo rtant. Arousal andfeeding of one or afew usua llyleadtothe
stimulat
iono fothers.

Loca
tionandiden
tif
ica
tion
Thechem icalandphys ica lcharacte r
isticso ffoodpa r
ticlesareimpo rtan tto
considersotha tfishcansuccess ful
lyloca tefood . Makingfood moreobv ious
tofish w i
ll makethem eas iertoloca teinthe wa te
rco lumn . This can be
done by add ing chem ica la t
trac tan ts, prov id
ing co lor and con trast,
maintaining desirable s i
ze and shape o f pa rt
icles
, and man ipula
tingl ight
intensityandtu rb
idi ty.V is
ion,olfaction,andgus tat
ion(senseo ftaste using
surfacetas tebud )a rea llimpo rtantinf ind ingandloca tingfooditems . The
farme rcanfeedthef ishregu larlyin ase ta reaofthefa rm orthetank and
encou ragefishtoagg rega te.Itisausua lp rac t
iceinsh r
impfa rmstop rovide
feedingt rays a
tseve ralp laces whe resh r
impcancometoea t
.

Cap
ture
In carnivores, small part
icles can beinges ted who le by being suckedin to
the mou thf rom a distance. Cap tu
resuccessinthe w i
ld depends on p rey
size and avo idance behavior, bu tthisis no t applicablein cu lture using
comp lete o
r supp lemen taryfeed . Howeve r, movemen tin wa ter currents
maycause p rob lemsinge ttinga tthefood . Food pa rtic
les i
zeisafac torin
capturesuccess . Big par
ticles notea tenimmed iately wi
lldisintegrateand
be wasted. Howeve r
,ingene ral,somefishandsh r
imp w il
lcon t
inueton ibble
bigfood particles unt
ilinges tionis possib
le.

Tas
te Tes
ting
Oncethefoodiscap turedinsidethe mou th, thefish o
rsh r
imptas testhe
foodleadingtoswallowingorrejec
tion
. No tal
l materia
lstakenintothe mouth
areswallowed, they aretestedforsuitabil
ity bytasterecepto
rsinsidethe
mouth,gi
llarches,gil
lrakers
,andinthet issuesurround ingthepharyngeal
tee
th.

Swa
llow
ingo
r Re
jec
tion
Food wh ichtas tesgood and have op timal part
icles ize, shape , andtex ture
wil
lbeswa l
lowedbyf ish. Mo rere jectionsandd isin tegrationoccurinpe llets
tha ta re ha rd, ab ras ive and a re muchlonge rthanthe ird iame ter. La rge
particles a re usua llye jectedthroughthe mou th w ith a “cough ing ”ac tion.
The refore,feedsshou ld havetheco r rectsizeandtex turefo rthespec iesin
cultu re. De laysintheinges tiono ffood duetorepea tedsp i
ttingou toffood
particlesbythef ishleadtoinc reasedene rgyexpend itureandfood was tage.
The ove ra l
lp racticalimp l
ica tions o fthe know ledge onfeed ing p rocessto
fishcu lturea re:encoun terratesshou ld be maxim ized byconcen tra t
ingfood
availabilityint ime andspace ;foodsupp ly mustbe ma tchedtoappe tite
; and
chem icalandphys icalcha racteristicso ffoodpa r
ticlesneedtobere latedtof ish
spec ies,s i
ze, andsenso ryabili
tiestoa idlocation,iden tif
ication,andcap tu re.
 CHAPTER 3 Feed
ing Hab
itsand D
iges
tive Phys
iologyo
fFishes 95
Summa
ry
Thera tea twh ichf ishd iges tthe i rfoodiso fp rima ryimpo rtanceinde term in ing
frequency ,feed ingra tes , andra tions ize.F ishescanbec lassifiedacco rd ing
tothe i rd ie to rfoodtheyhab i tua llyea tashe rb ivores ,ca rn ivores ,omn ivo res ,
plank tivo res ,andde tritivo res . Ac lass if
ica tionbasedonthe manne ro ffeed ing
orfoodge ttinginc ludesp reda tors,g raze rs,s t rainers ,sucke rs,andpa ras ites.
Fish adap ttothe i rfood d i
ffe rences by ana tom ic as we ll as behav io ral
means . Thereis as t rongco rrela tion be tween ana tom ica ls truc tu re o fthe
diges t
ivet rac t andthefeed ing hab its o fthef ish andthefood ea ten . The
mou th has avar ie ty o fadap tationsfo rcap tu ring, hand l
ing and so rting o f
foodbe fo reen tryin tothes tomach .F ishhavetee ththa tva ryintype , numbe r,
and a r rangemen t. The a rrangemen tands truc ture o fthetee th a rere la ted
tothe k indo ffoodtha tthef ish no rma llyea t.
Thes tomacho ffishes hasacon figura tiono rshape wh ichisconven ien t
forcon tain ingfood . The s i ze o rcapac ityo fthes tomachinre lationtothe
body we igh tvar ies be tween spec ies andis usua l
lyre la tedtothein terva l
betweenfeed ingsandtothes izeo ffood pa rtic les.A llf ishes havein tes tines .
Theleng tho fthein tes tinesvar iesf romaslowas 1/5toas h ighas20t imes
the bodyleng th . Carn ivo res have are la tive ly simp le and sho rt gu t,w i
th
thick mucosafo r abso rp t
ion wh ile he rb ivo res have along andth in gu tto
inc reasegu tre ten t
iont imeandenhance d iges tion andabso rp t
ion .
Diges tionisap rocess wherebyinges tedfood nu trien tssuchasp rote ins ,
lipids,andca rbohydra tesa reb rokendownin to uni tstha ta resma llenough
to be abso rbed ac rossthegu t wa ll
. The p rocessis accomp l
ishedth rough
the ac tion o fd iges t
iveenzymes . Enzymestha ta idin d iges tion o fp rote ins
are knownasp ro teases . Fo rca rbohyd rate d iges t
ion ,theenzymesinvo lved
are ca rbohyd rases .L ip id d iges tionisfac ili ta ted by enzymes l ipases and
phospho l
ipases .
The ma in d i fference be tweenf ish andc rus tacean d iges t
ionistha t,in
thela tte r
,enzymesa resec re tedbythehepa topanc reas . Thehepa topanc reas
alsoac tsass to rageo rganfo rg lycogen ,fat,andcon tainsenzymesfo rsec retion
ofb ile ac ids . The c rus tacean hepa topanc reasto ag rea tex ten tfu lf
illsthe
roleo fthel ive rinve r teb rates .
Ana lys is and measuremen to fs tomach con ten ts g ivesin fo rma t
ion on
thefeed ingp re fe rencesas we l
lasf requencyo ffeed ingo fapa rticula rspec ies.
Diges t
ib il
ity measuremen tscanbedoneth rough inv ivoandinv i
tro me thods .
Know ledge o ffeed ing processinf ish and sh rimpis use fulto max im ize
foodloca tion ,cap tu re , andinges tion .

Gu
ide Ques
tions

1
. Wha
tco
rre
lat
ionex
istsbe
tweenana
tomyo
fthe d
iges
tivet
rac
tand
feed
ing hab
itso
ffishes?
2
. Wha
tarethefourtypeso
ffeed
ingbehav
ioro
ffishinthe
ir na
tura
l
hab
ita
t?
3
. Wha
tarethe d
iffe
ren
tpa
rtsandfunc
tionso
fthe d
iges
tivesys
tem o
f:
a
)fish
b
) c
rus
taceans
96 CHAPTER3 NUTR
ITIONINTROP
ICALAQUACULTURE

4
. Wha
tarethefou
rgene
ralcon
figu
rat
ionso
rshapeso
fthef
ishs
tomach?
5
. D
iscussthere
lat
ionsh
ipbe
tweengu
tleng
thandfeed
inghab
itso
ffishes?
6
. D
iffe
ren
tia
tethefourtypeso
fep
ithe
lia
lce
llsfoundinthe
hepa
topanc
rea
tictubu
leo
fthec
rus
taceans
.
7
. De
fineandexp
lain d
iges
tion
.
8
. Desc
ribethes
tepsandenzymesinvo
lvedin
:
a
) pro
tein d
iges
tion
b
)lip
id d
iges
tion
c
) ca
rbohydra
te d
iges
tioninf
ishes andc
rus
taceans
9
. D
iscussthe me
thods usedto measures
tomachcon
ten
tso
ffishes
and d
iges
tib
ili
tyo
ffeeds
.
10
. D
iscussthefac
torstha
t maya
ffec
tdiges
tionandabso
rpt
ioninf
ishes
.
11
. Exp
lainthe d
iffe
ren
tstagesinthefeed
ing processinf
ish

Sugges
ted Read
ings

ADCP
. 1980.FishFeedTechnology
. Aquacul
ture Development Coo
rdination
Prog
ramme , Food and Agr
icul
tural Organiza
tion. Un
ited Nations,
Rome. ADCP/REP/80/11 , 395 p
.

Ash R
. 1985. Pro
tein d
igestionandabsorp
tion.In: Cowey CB
, Mackie AM
,
Be
llJG,(eds)
. Nu t
rit
ionandfeed
inginfish. AcademicPress
.London.
p 69
-93.

Cecca
ldi HJ . 1997. Anatomy and physiology o
fthe digest
ive sys
tem.In:
D’AbramoLR , Conkl
in DE,Akiyama DM(eds )
. Crus
tacean Nutr
ition
,
Vol.6, Wor
ld Aquacu l
ture Soc
iety
, Ba ton Rouge
, Louis
iana, USA.p
261-291.

Ben
itezLV,TiroLB. 1982
.Stud
iesonthed igest
ivep
roteaseso
fthe m
ilk
fish
,
Chanoschanos. Mar
.Bio. 71
:309
-315 .

Bo
rlonganIG. 1990
.Studies onthe d
igest
ivel
ipases o
fmi
lkf
ish
, Chanos
chanos
. Aquacu
lture 89:315
-325.

Ch
iu YN
, Ben
itez LV
. 1981
.S tud
ies onthe ca
rbohydrasesinthe d
iges
tive
t
racto
fthe m i
lkf
ishChanoschanos. Mar.Bio
l. 61
: 247
-254.

Da
ll W
. 1992
. Feeding,digest
ionandass imi
lat
ioninPenae idae.In:Al
lan GL,
Da
ll W.(eds).Proceedings ofthe Aquacu lture Nutr
ition Workshop,
Sa
lamande rBay, 15-17April 1991
. NSWF isher
ies,Brackish Wate
rFish
Cul
ture Resea
rch Stat
ion, Salamander Bay, Aust
ral
ia.p57 -63.

Fange R, Grove D
. 1979. D iges
tion
.In : Hoa
r WS, Randa
ll DJ
, BrettJR
(eds)
.F ish Phys
iology
, Vol
.VIII
, Academic P
ress
,Inc.
, New York
.p
161-260.
 CHAPTER 3 Feed
ing Hab
itsand D
iges
tive Phys
iologyo
fFishes 97
Kn
igh
tB . 1985. Feed
ingbehavio
randf
ishcul
ture
.In: Nutr
itionandfeeding
infish. Cowey CB, Mack
ie AM
, Be
llJG(eds
). Academ ic P
ress
,Inc.
,
London.p 223 -24
.

Lege
rC. 1985.Diges
tion,absorp
tion
,andt ransportofl
ipids
.In
: Cowey CB
,
Mackie AM, Bel
lJG(eds )
. Nutr
ition andfeed
inginfish
. Academic
Press
, London.p 299-332
.

McLaughlin PA. 1983

.Interna
l Anatomy.In: Man
telLH(ed
).The B
iologyo
f
Crustacea
. Vol.5
. Academic P
ress,Inc
, NewYork
.p 1-52
.

Sm
ith LS. 1989
.D iges
tive Funct
ionsinTe
leostFishes
.In: HalverJE(ed)
.
Fish Nut
rit
ion, 2ndedi
tion
. Academ
ic P
ress,Inc, New York.p 331
-
421.

Vonk HJ
. 1960. Digest
ion and metabol
ism.In
: Wa terman TW(ed)
. The
Physio
logy o
f Crustacea
. Vo
l.1, Academ
ic P
ress,Inc
. New York.p
291-316.

Wee KL
. 1992. Anove rv
iewo ffish d
igestive phys
iologyandthere levanceto
theformu lat
ion ofa rt
ific
ialfishfeeds .In: A l
lan GL, Da l
l W(eds ).
Proceedingso ftheAquacu lture Nutr
ition Workshop, Sa
lamande r Bay
,
15-17 Apr il 1991. NSW F ishe r
ies, Brackish Wa terF ish Culture
Research S ta
tion, Sa
lamande r Bay, Aust
ralia.p 17-24.
 CHAPTER

Fo
rmu
lat
iono
fAqua
feed
s
MAE R
. CATACUTAN
4
In
troduc
tion
The deve lopmen to f afeedtha tis bo th e ffective and econom icalfo r an
aquacu lture spec iesin a llitslife stagesis a con tinuous e f fort. Aqua feed
deve lopmen ts tar ted when na tura lfood sourcesin cu l ture sys tems
becameinadequa te and hadto be supp lemen ted w ith p repa redfeed . As
fish s tock ing dens it
iesin cu ltureinc rease , supp lemen ta lfeed ingis no
longersu f
ficient. A comp letefeedtha tcon tains allthe necessa ry nu trien ts
in su fficient amounts to br ing about good growth , surv iva l, and
reproduc tionis needed . Feeding red ien ts gene rally comef rom an ima lo r
plan t sources and some a re by-produc ts o fthefoodindus try. The reis no
sing lefeed ingred ient orfeeds tuf ftha t con tains a ll the nu trien tsin
adequa te amoun ts . Thus,d i
fferentfeeding redien ts arecomb inedto make
afeedtha t hasthe des i
red compos ition and nu trientleve ls.Incomb in ing
var iousfeeding red ients,i tisimpo rtan tto know how much o feachfeed
ingred ien t shou ld be usedto produce a cos t-e f
fec t
ive aqua feed .
W
iththe g row th and expans ion o faquacu lturein to a ma jorindus try,
seve ra lf ish spec ies a re be ing cu ltured ; thus , the deve lopmen to f mo re
efficien t aqua feed formu lations shou ld con tinue . In deve loping cos t -
effect ivefo rmu lated d iets, manyimpo rtan tfac tors haveto be cons ide red.
Th is chap terd iscussesthesefac to rs andthe ma thema tica l ca lcu lations
info rmu lating afeed .Ita imsto enab le s tuden tstofo rmu la te d iets us ing
pur ified and prac t
icalfeedingred ien ts , and a lso toformu late e ffec tive
supp lemen tal and comp lete dietsfor aquacu lture spec ies .

FeedIng
red
ien
tsfo
rAquacu
ltu
re

Feeding red ients orfeedstuffsfor aquacu l

ture comef rom many d if
ferent
sources and a re used because of
ten t
imesthey a re notu ti
lizedfo r human
consump tion. The cho ice of afeedingred ientin aqua feedformu lation
depends on : content o f essent ia l nutrients ; d igest ib i
lity and
bioava
ilab il
ity; absence o f ant
inu t
ritiona
lfac tors andtox ic subs tances;
commerc ial availabil
ity; and cost
.
Anima l and p lan t materia
ls arethe mos t commonfeeding redients.
Some was tes and by -products ofthefoodindus try arealso u t
ilized. The re
100 CHAPTER 4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Tab
le4
.1 Somesourcesofpro
tein
,lip
id,andca
rbohyd
ratein a refeedingred ients tha t areind igenousin
aquacu
ltu
refeeds some areas and may no t be ava ilab le in
Pro
tein
L
ipid
Carbohydra te
c omm ercial q ua n
tit
ies. Som e f eed in gre dien ts
for aquacu lturefeeds a rel istedin Tab le 4 .1
B
lood mea
l Bee ftallow Breadf lour
Cop
ra mea
l Corn o i
l
a n d show n i n Figure 4
. 1
.
Cassava s tarch
De
fat
ted soybean mea
l Codl ive roil Corn mea l
Feedstu f
fs o fan ima lo r
igin usua llycon tain
F
ish mea
l Coconu to i
l Corn starch high amoun ts ofp rotein with good am ino ac id
Mea
t and bone mea
l Cottonseed o il Finerice bran pro file (Tab le 4 .2). A prote in source is
Sh
rimp head mea
l Menhaden o il Sago pa lm starch cons idered goodi fthe am ino ac id pro fileis
Sh
rimp mea
l Pollackl ivero i
l Seaweeds closetotha to fthe spec ies be ingfed . Some
Squ
id mea
l Tunal ive roil plant prote in sources have h igh prote in
T
rashf
ish Squ idlive roil con ten t bu t they o ften havein ferior am ino
Yeas
t Soybean o il
ac id pro fi
les compared w ith pro tein f rom
an ima l sources . Feeds tuffsf rom p lan ts a re
good sou rces o fca rbohyd rate, and some a re ofgood p rotein qua l
ity such
as soybean mea l. Some p rotein sou rcesl ikelegumes and nu ts a re a lso
sou rces o
fene rgy bu ta re mo reexpens ivethanthe common ca rbohyd ra te
sources w ith h ighleve ls ofd igestible carbohydra te such as sago pa lm
sta rch and b readf lour. Gene ral
ly, carbohyd rates a re cheape r sources o f
ene rgy (F igu re 4 .2 )thanl ipid o rfat sou rces. They a re a lso used asfeed

F
igure4
.1
Somefeedstu
ffsfo
raquafeeds
,fisho
ffa
l(A)shr
imp mea
l(B
)an
ima
l mea
t was
te(C
) mea
tand
bone mea
l(D)soybean mea
l(E)andyeast(F
).
 CHAPTER4 Fo
rmu
lat
ion o
f Aqua
feeds 1
01
T
abl
e4.2 Am
inoac
idcompos
it
iono
fsomef
ish mea
ls(FM
),lea
f mea
ls(LM
)ando
the
rpro
teinsou
rces
Amoun
t(g
/100gp
rote
in)

Am
ino ac
id Wh
ite Pe
ruv
ian Sapsap Tuna Tabagak Shr
imp Squ
id Soybean Kangkong Camote Ip
il
-ip
il Acac
ia
FM FM FM FM FM meal meal mea
l LM LM LM LM T
ama
r
ind
LM

A
lan
ine 6
.0 6
.1 6
.7 5
.0 5
.5 5
.5 5
.8 4
.0 5
.0 5
.2 6
.2 3
.2 3
.6
Arg
inine 6
.2 5
.5 4
.0 5
.4 4
.7 7
.1 7
.5 6
.0 3
.3 3
.7 5
.2 2
.2 2
.4
Aspa
rtic ac
id 9
.4 8
.7 9
.5 8
.5 8
.1 8
.3 10
.1 11
.1 8
.7 10
.2 11
.4 6
.3 7
.6
Cys
tine 0
.7 0
.9 0
.8 0
.7 0
.9 0
.6 0
.9 1
.2 0
.5 0
.3 0
.6 0
.5 0
.5
G
lutam
ic ac
id 13
.8 13
.2 14
.9 11
.2 11
.3 12
.4 15
.8 18
.7 8
.8 10
.2 11
.2 6
.6 7
.6
G
lyc
ine 7
.0 5
.4 6
.4 5
.2 5
.3 4
.9 5
.7 4
.0 4
.3 4
.7 6
.0 3
.2 4
.0
H
ist
idine 2
.2 1
.8 3
.9 5
.6 2
.3 2
.1 2
.2 3
.2 2
.7 2
.8 1
.4 3
.9 4
.0
Iso
leuc
ine 3
.9 4
.1 4
.5 3
.9 4
.2 3
.8 4
.2 4
.5 3
.4 3
.7 6
.6 2
.4 3
.0
Leuc
ine 7
.4 7
.1 7
.5 6
.5 6
.6 6
.7 7
.7 7
.2 6
.5 7
.9 6
.6 4
.8 5
.7
Lys
ine 7
.7 7
.4 7
.3 6
.2 7
.3 6
.3 7
.8 5
.8 4
.6 4
.4 6
.1 2
.9 3
.4
Me
thion
ine 3
.0 2
.8 2
.9 2
.3 2
.8 2
.4 2
.8 1
.4 1
.5 1
.8 1
.2 0
.9 0
.9
Pheny
lalan
ine 4
.3 3
.8 3
.8 3
.8 3
.5 3
.8 3
.4 5
.2 5
.7 6
.5 3
.9 3
.8 4
.2
Pro
line 4
.3 4
.2 3
.5 3
.7 3
.0 3
.2 3
.8 5
.5 3
.9 3
.8 5
.5 3
.6 3
.8
Ser
ine 4
.4 3
.6 3
.4 3
.3 3
.1 3
.1 3
.9 5
.2 3
.7 4
.3 4
.4 3
.2 3
.8
Threon
ine 4
.2 3
.9 3
.9 3
.6 3
.7 3
.6 4
.1 3
.8 3
.9 4
.4 5
.1 2
.7 3
.6
Tryp
tophan 1
.0 0
.7 1
.2 0
.8 0
.8 1
.1 1
.0 0
.4 NA NA NA NA NA
Ty
ros
ine 3
.8 3
.2 4
.4 2
.6 3
.3 3
.3 3
.1 2
.7 4
.1 6
.5 3
.4 3
.5 3
.0
Va
line 5
.1 5
.2 5
.2 4
.5 4
.9 4
.3 4
.3 4
.4 5
.3 5
.8 6
.3 3
.5 3
.9
% Pro
tein 70
.5 70
.5 66
.0 77
.2 78
.4 71
.8 78
.7 43
.8 31
.9 30
.8 27
.8 25
.3 15
.1

Value
sgivena
re mean
s; NA
,no
tana
lyzed
Sou
rce
: Peña
flo
rida 1989

F
igu
re4
.2
Somesou
rceso
fdie
taryene
rgya
reco
rn(A
),r
iceb
ran(B
),cas
sava(C
).

binders. As idef rom carbohydra tes,l ip ids a re also used as sources o f

energy. They a lso provide essentialfat ty ac idsinf ish die ts
. Examp les of
common ly used d ietarylipid sources a ref ishl iver o ils and p lan to i
ls
such as soybean o il and com o i
l.
The cho ice o fafeeding redientis ma in ly dependen t onthe amoun to f
essentia l nutr ients thati t con tains. Tab le 4 .3 g ives the prox ima te
compos i
tion(c rude protein,c rudefa t,d iges tib
le carbohydra te o r NFE ,
crudef iber, and ash)o fsomefeeding red ients. Feeds tuffscon taining high
amoun ts ofp ro
tein with good amino ac id p rof
ile are usua l
ly expens ive and
their useis cons t
rained by cost. The b ioava ilability of nu t
rien ts presen t
in afeeds tuffvariesfo rd i
fferent aquacu l
ture spec ies and w i l
lin f
luence
theleve lofinc lusiono fafeeds tuffinthefeedfo rmu la.A lthoughfeeds tuffs
may con tain the same amoun to f nu t rien t
, for examp le pro tein, the
feedstu ffw ith more d igest
ible pro te
in shou ld be chosen ove rtha tw i
th
102 CHAPTER 4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Tab
le4
.3 Proximatecompos it
ionofsomefeeding
redien
tsanalyzedbythe Cen
tra
lized
Analy
tical Labora
torya
tSoutheas
t As
ianFisher
ies Deve
lopment Cen
ter
,
Aquaculture Depar
tment*

Amoun
t(% d
ry ma
tte
r)

C
rude
Mois
ture Crude Crude NFE** Ash
F
ibe
r
Pro
tein Fa
t

An
ima
l:
F
ish mea
l(FM
,loca
l)(6
) 10
.3 64
.1 6
.5 0
.8 8
.5 20
.1
FM
, Ch
ilean (27
) 8
.4 70
.1 8
.5 0
.5 4
.1 16
.8
FM
, Dan
ish(2
) 9
.5 73
.9 9
.4 0
.3 2
.4 14
.0
FM
, Pe
ruv
ian(30
) 8
.3 68
.3 5
.9 0
.8 7
.7 17
.3
FM
, Pe
ruv
ian(26
) 7
.1 67
.9 10
.0 1
.3 4
.1 16
.7
FM
,tuna(9
) 9
.4 65
.4 8
.0 0
.8 8
.8 17
.0
FM
, wh
ite(11
) 7
.2 69
.0 7
.6 0
.6 4
.8 18
.0
P
rawn head mea
l(35
) 6
.5 51
.2 5
.2 13
.3 5
.3 25
.0
Sh
rimp mea
l, Ace
tes sp
.(60
) 8
.2 68
.6 3
.9 3
.6 7
.6 16
.3
Squ
id mea
l(60
) 6
.9 78
.5 5
.5 1
.3 6
.7 8
.0
Squ
id mea
l, sc
rap (4
) 5
.5 74
.1 7
.1 0
.9 8
.1 9
.8
F
rog mea
l(2
) 7
.6 62
.5 1
.7 1
.2 4
.7 29
.9
B
lood mea
l(2
) 6
.3 87
.7 3
.0 0
.4 3
.3 5
.6
Mea
t and bone mea
l(19
) 5
.6 46
.8 9
.6 2
.0 7
.5 34
.1
P
lan
t:
Acac
ia Lea
f Mea
l(LM)(2
) 4
.4 25
.7 5
.6 21
.2 41
.7 5
.8
A
lfa
lfa LM 7
.2 17
.2 3
.0 27
.7 42
.9 9
.2
Camo
te LM
,(7
) 4
.5 29
.7 4
.9 10
.0 43
.2 12
.2
Cassava LM
,(8
) 5
.9 22
.1 9
.3 12
.4 49
.2 7
.0
Ip
il-
ipi
l LM
,gian
t(14
) 7
.8 25
.1 6
.8 10
.6 44
.0 13
.5
Ip
il-
ipi
l LM
, na
tive (6
) 10
.3 29
.3 8
.8 11
.5 43
.5 6
.9
Kang
-kong LM (6
) 5
.7 28
.5 5
.4 10
.5 43
.6 12
.0
Ma
lunggay LM (7
) 3
.5 30
.4 8
.4 8
.3 43
.7 9
.2
Papaya LM (10
) 5
.4 20
.7 11
.6 11
.2 42
.6 13
.9
Cop
ra mea
l(10
) 7
.9 22
.0 6
.7 17
.3 44
.3 9
.7
Cowpea (7
) 8
.0 23
.0 1
.3 4
.1 67
.5 4
.1
Cowpea
, dehu
lled (2
) 7
.7 25
.4 0
.9 1
.4 68
.3 4
.0
Mungbean
,green (5
) 7
.1 23
.2 1
.2 3
.1 68
.7 3
.8
Mungbean
, ye
llow (5
) 7
.7 24
.1 1
.1 3
.8 67
.1 3
.9
R
ice bean (2
) 5
.0 26
.5 0
.8 4
.0 64
.6 4
.1
Co
rn mea
l(10
) 8
.4 7
.8 4
.7 2
.6 83
.1 1
.8
Co
rns
tarch (5
) 11
.9 0
.4 0
.2 1
.1 98
.2 0
.1
F
lou
r,b
read (40
) 12
.1 12
.9 1
.2 0
.3 84
.9 0
.7
F
lou
r, who
le whea
t(15
) 11
.3 15
.3 1
.7 0
.8 81
.1 1
.1
Whea
t, Po
lla
rd (4
) 9
.5 15
.4 4
.5 10
.3 64
.0 5
.8
Ge
rm, whea
t(2
) 6
.0 27
.8 4
.3 3
.4 59
.6 4
.9
G
luten
, co
rn(5
) 7
.3 62
.6 7
.7 2
.2 25
.9 1
.6
G
luten
, whea
t(6
) 8
.9 80
.7 1
.4 0
.4 16
.4 1
.1
R
ice b
ran (78
) 9
.2 13
.3 14
.1 8
.5 53
.4 10
.7
R
ice b
ran
,tik
i-t
iki (5
) 10
.7 18
.0 2
.0 8
.0 62
.4 9
.6
R
ice hu
ll(7
) 7
.0 3
.3 2
.0 32
.4 41
.6 20
.7
Soybean mea
l, asis(21
) 5
.6 35
.8 19
.8 4
.9 33
.9 5
.6
Soybean mea
l, de
fat
ted (108
) 8
.4 43
.6 1
.5 5
.5 41
.7 7
.7
Other sources
:
Case
in (11
) 7
.2 89
.7 0
.1 0
.3 8
.9 1
.0
Crab mea
l(2
) 4
.2 37
.9 4
.1 10
.7 8
.9 38
.4
Ge
lat
in (6
) 7
.9 94
.4 0
.0 0
.1 5
.1 0
.4
Musse
l mea
l,g
reen (30
) 5
.9 64
.6 8
.6 3
.0 12
.5 11
.8
Oys
ter mea
l(6
) 4
.4 54
.6 9
.4 4
.0 20
.1 11
.9
Sca
llop mea
l(2
) 7
.3 65
.2 10
.9 1
.4 8
.8 13
.7
Sna
il mea
l, kuho
l(5
) 4
.0 52
.1 1
.8 2
.1 15
.7 28
.3
Yeas
t, Brewe
rs(2
) 7
.2 49
.4 1
.6 2
.4 34
.5 12
.1
 CHAPTER4 Fo
rmu
lat
ion o
f Aqua
feeds 103
Tab
le4
.3 (con
tinued
)

Amoun
t(% d
ry ma
tte
)
r

Mois
ture Crude Crude NFE*
* Ash
Pro
tein Fa
t

Yeas
t, Cand
ida (3
) 8
.3 55
.2 0
.8 1
.7 35
.1 7
.4
Na
tura
l Food
:
Aca
rtia sp
.(copepods
) 7
.8 71
.2 8
.3 5
.4 9
.9 5
.2
Ar
tem
ia (37
) 8
.0 55
.5 6
.8 11
.3 15
.0 11
.4
Azo
lla (2
) 8
.0 27
.2 3
.4 12
.9 36
.5 20
.0
Brach
ionus sp
.(5
) 8
.1 51
.9 10
.4 3
.5 15
.3 18
.9
Chae
toce
ros ca
lci
trans (7
) 7
.6 24
.4 7
.1 2
.5 26
.7 39
.3
Ch
lore
lla
, ma
rine (3
) 10
.1 35
.1 4
.2 5
.6 27
.7 27
.4
Isochrys
is ga
lbana (2
) 10
.4 33
.6 18
.1 4
.4 23
.0 20
.9
Mo
ina mac
rocopa (3
) 8
.5 57
.8 7
.6 8
.4 17
.2 9
.0
Sa
rgassum (2
) 10
.4 9
.0 0
.8 9
.6 46
.4 34
.2
Ske
letonema sp
.(4
) 10
.4 24
.7 2
.6 0
.7 20
.2 51
.8
Sp
iru
lina (2
) 8
.0 56
.7 2
.8 0
.6 28
.1 11
.8
Te
trase
lmis sp
.(4
) 5
.5 49
.1 10
.7 2
.1 19
.0 19
.1
D
igman (4
) 9
.8 20
.6 3
.3 16
.4 35
.9 23
.8
En
teromo
rpha (
lumo
t) (15
) 15
.2 13
.8 1
.9 9
.3 36
.9 38
.1
Grac
ila
ria sp
.(18
) 7
.0 10
.2 0
.4 5
.8 44
.8 38
.8
Kappaphycus sp
.(10
) 6
.1 5
.4 0
.8 6
.1 57
.3 30
.4

*Va
lue
sa re meansforthenumberofsamplesgiveninpa
ren
theses
. Notal
lfeeding
red
ientsareava
i
lab
lein
commerc
ialquant
it
iesbu t maybeused whe
retheyarecommonlyfoundinla
rgeamounts.
*
*NFE- Nit
rogen-
free ex
trac
t

poor prote
in avai
lab
ili
ty. The diges
tib
ili
ty of pro
tein (expressed in
percen
tage)in somefeedstu
ffsfor some aquacu
lture spec
ies a
relisted
in Table 4
.4.

Tab
le4
.4 Apparentpro
teindiges
tib
il
itycoe
ff
icien
ts(APDC
)in %o
fsomefeeds
tuf
fsfo
r
aquacul
turespecies
Aquacu
ltu
re Spec
ies Feeds
tuf
fs APDC % Re
ferences

Sh
rimp
T
ige
rsh
rimp F
ish mea
l 61 Ca
tacu
tan 1997
Soybean mea
l, de
fat
ted 93
Squ
id mea
l 96
Shr
imp mea
l 95
Shr
imp head mea
l 89
Mea
t and bone mea
l 74
Yeas
t Cand
ida sp
. 93
Cop
ra mea
l 75
F
ish
M
ilk
fish F
ish mea
l 45-81* Fe
rra
ris e
tal
. 1986
De
fat
ted soybean mea
l 45-94*
Carp Wh
itef
ish mea
l, 95 NRC 1977
mechan
ica
l ex
trac
ted
Soybean seed mea
l, 81-96
so
lven
t ex
trac
ted
Red sea b
ream Wh
itef
ish mea
l, 61-87
mechan
ica
l ex
trac
ted
Channe
lca
tfish Soybean seed mea
l, 72-84
so
lven
t ex
trac
ted

*tes
ted a
tdi
ffe
ren
tsa
l
ini
ties
104 CHAPTER 4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Many o the
r componen ts a
re addedinthefeedfo rmu la as
idef romthe
major sources o
f nut
rients.In comp le
tefeedfo rmulations, micronutr
ients
are addedin sma l
l amoun tsinthefo rm ofvi
tam in and m ineral mix
tures
.
Examp les ofthese mixesforc rustaceans andfishes are shownin Tab les
4.5,4 .6, and 4.7. For econom ic reasons,o ther subs tances are also

Tab
le4
.5 V
itam
inand m
ine
ralm
ixtu
resfo
rcrus
taceans(A
)andt
ige
rsh
rimpjuven
ile(B
).
mg/100 g d
ry d
iet

A* B**

Th
iam
ine HC
l(B1
) 1
.26 4
.0
Py
ridox
ine HC
l(B2
) 3
.70 12
.0
R
ibo
flav
in (B6
) 2
.57 8
.0
Cyanocoba
lam
in (B1
2) 0
.025 0
.08
N
ico
tin
ic ac
id 12
.62 40
.0
Fo
lic ac
id 0
.25 0
.8
B
iot
in 0
.13 0
.4
Pa
ra am
inobenzo
ic ac
id 3
.16 10
.0
Ca
lcium pan
tothena
te 18
.93 60
.0
Inos
ito
l 126
.18 400
.0
Na
-asco
rba
te (C
) 630
.92 2000
.0
Cho
line ch
lor
ide 189
.27 600
.0
β
-ca
rotene (A
) 3
.03 9
.6
Ca
lci
fero
l (D
) 0
.38 1
.2
α-
tocophero
l(E
) 6
.31 20
.0
Menad
ione (K
) 1
.26 4
.0

TOTAL 1000
.00 3170
.08

g
/100 g d
ry d
iet
MINERALS
A* B*
*

K2HPO4 2
.339 2
.000
Ca3(PO4
)2 3
.181 2
.720
MgSO4
.7H2O 3
.556 3
.041
NaH2PO4
.2H2O 0
.924 0
.790

TOTAL 10
.000 8
.551

*Sou
rce
: Tesh
imaand Kanazawa1982

includedinthefeedformu lato reducefines dur ingfeed manu fac

ture,
and storagelosses duetofeed degrada t
ion and spo ilage
, andimprove
feed durab i
lity during handl
ing and wa ter s
tabil
i ty. These substances
include synthe t
ic binders
, antioxidan
ts, and mo ldinh ib
itors
. To make
feed more a ttract
ive tofish,p igments and att
rac tants may be added
(Table 4
.8)
.

FeedFo
rmu
lat
ion

Feed formu lation is a process thatinvolves comb in

ing various feed
ingred ients
, wh ich con ta
in dif
ferent amoun ts o
f nu tr
ients
, sotha tthe
resultingcompos i
tion would mee tthespec
ificrequi rementofthecu ltured
species. The nu tr
ien tleve
ls offeedingredientsto be used a re ba
lanced
mathema t
icallyin o rde
rto come up w iththe des iredfina
l compos it
ion.
 CHAPTER 4 Fo
rmu
lat
ion o
f Aqua
feeds 105
Tab
le4
.6 Recommendedvi
tamin m
ixtu
refor wa
rmwa
terf
ishessuch
as m
ilk
fish
,seabass
,andcatf
ish*
V
itam
in Amoun
t(pe
r kg
)in d
ry d
ieta
Supp
lemen
tal Comp
lete

V
itam
in A ac
tiv
ity 2
,000 IU 5
,500 IU
V
itam
in D3ac
tiv
ity 220IU 1
,000IU
V
itam
in E 11IU 50IU
V
itam
in K 5 mg 10 mg
Cho
line 440 mg 550 mg
N
iac
in 17
-28b mg 100 mg
R
ibo
flav
in 2-7b mg 20 mg
Py
ridox
ine 11 mg 20 mg
Th
iam
in 0 20 mg
Ca
lcium pan
tothena
te 7-11b mg 50 mg
B
iot
in 0 0
.1 mg
Fo
lac
in 0 5 mg
V
itam
in B1
2 2
-10 mg 20 mg
Asco
rbic ac
id 0
-100b mg 30-100b mg
Inos
ito
l 0 100 mg

a Theseamoun
ts dono ta
llowforprocess
ingo rsto
ragelosses.
b Highes
tamounts a
reapp rop
ria
te when“stand
ingc rop
”offishexceeds500kg
/hec
tare
.
* Source
: NRC 1977 .

Tab
le4
.7 Mine
ralm ix
turesfo
rpu r
ifiedand T
able4
.8 O
the
rfeedadd
it
ives
p
ractica
l warmwaterf
ish diets
*
Feed B
inde
rs
Minera
l D
ry D
iet(g
/100g
) aga
r
a
lgin
ic ac
id
Prac
tica
lDie
ts
as
tarch
CaCO3 0
.750
ben
ton
ites
MnSO4
.H2O 0
.030
ca
rboxyme
thy ce
llu
lose (CMC)
ZnSO4
.7H2O 0
.070
car
rageenan
CuSO4
.5H2O 0
.006 ge
lat
in
FeSO4
.7H2O 0
.050 g
rac
ilar
ia (d
ried and g
round
)
NaC
l 0
.750 hem
ice
llu
lose
K
IO3 0
.0002 l
ignosu
lfa
tes
CaHPO4
.2H2O 2
.00
An
tiox
idan
ts
Pur
if
ied D
iets
v
itam
in C
CaHPO4
.2H2O 2
.07
bu
tyla
ted hyd
roxyan
iso
le (BHA
)
CaCO3 1
.48
bu
tyla
ted hyd
roxy
toluene (BHT
)
KH2PO4 1
.00 e
thoxyqu
in (1
,2 d
ihyd
ro-6
-ethoxy
,
KC
l 0
.10 -2
,2,4-
trime
thy
lqu
ino
line
)
NaC
l 0
.60
Mo
ld inh
ibitors
MnSO4
.H2O 0
.035
ci
tric acid
FeSO4
.7H2O 0
.05
sod
ium
, ca
lcium
,or po
tass
ium so
rba
te
MgSO4 0
.30
KIO3 0
.001 A
ttrac
tan
ts:
CuSO4
.5H2O 0
.003 Co
lor
ZnCO3 0
.015 caro
teno
ids
CoC
l2 0
.00017 xan
tho
ter
in (
red and ye
llow xan
thophy
lls
)
NaMoO4
.2H2O 0
.00083
Feed
:
Na2SeO3 0
.00002
be
taine
g
lutam
ic ac
id
Sou
rce
: NRC 1977
taur
ine
106 CHAPTER 4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Ita lsotakesin to accoun tthe cos to f ma te r

ia ls , accep tab ility, and ease
of prepa ra tion .
Thetwo mos timpo rtan tfac to rstocons ide rinfo rmu lating afeedfo rany
aquacu ltu re spec ies a re nu t rien trequ iremen ts andfeed ing behav io r. Othe r
fac tors to cons ider a rethe s tagesinthel ife cyc le( la rval, grow-ou to r
broods tock ) andthetype o fcu l
tu resys tem .L ike anyo the ran ima l,f ish and
crus taceans need enough p rote in ,lip id, ene rgy ,v itam ins , and m ine ra ls .
These nu trien ts shou ld be presen tintheformu la tedfeedin adequa te
amoun ts . Excess ive amoun ts o f nu trien tsis was te fu l
, wh ileinsu ff
icien t
leve ls w i
llresu ltin s low g row th . An a l
lowanceis madeinthefo rmu lafo r
nu trien tlosses du ringfeed manu fac tu re.
Thefeed ing hab itand behav ioro fthean ima lisa lso animpo rtan taspec t
to cons ide rinfeedfo rmu lation . Fo rexamp le,c rus taceans such as sh r
imps
andc rabs a res lowea te rs andtheytakesomet imetoconsumethe pe l
le ted
feed .In con tras t,f ish consumethefeedimmed ia tely and ,o ften times,the
feedis ea ten be fo reittouchesthe pond bo t
tom . Thus ,c rus taceansrequ i re
a mo re wa te rs tab lefeedthanf ish . Afeed b inde r, usua lly a ca rbohyd ra te
sou rce,is addedtothefo rmu la tionto makethe pe lletedfeed s tayin tac t
muchlonge rinthe wa te r
. Seaweed ex trac ts such as kappa -ca rrageenan
and a lg ina tes a re common ly used as b inde rsin m ic roboundlarva ld ie t
prepa ra tion . La rva ld ie ts may be p repa redin suspens ionfo rm , co l
lo idfo rm ,
in so lu tion ,o rin d ryfo rm enc losed w i
th in a m icrocapsu le o r m icrocoa ted
with a b inde r. Some t
imes an a ttrac tan tis addedtothefeed sotha tthe
pelle tcaneas ily beloca ted and qu ick lyconsumed bythef ish. Mos taqua tic
spec iesinthela rvals tage s tayinthe wa te r column and g radua lly become
bottom dwe lle rs. Thus ,the p refer redfeed a tthe ea rlyl ife s tagesis afeed
tha t does no tse ttleinthetank bo ttom bu ts taysinthe wa te rco lumnfo ra
longt ime . Some aqua tic spec ies may p refe r a spec ificfeedfo rm ,type ,o r
sizetha t shou ld be cons ide redinfeedfo rmu la tion .
The requ i remen tfo r nu trien ts and ene rgy va riesfo rb roods tock ,g row
ou t
,andla rva ls tages ;the refo re,feedfo rmu la t
ion wou ld a lsova ryin nu tr
ien t
con ten tdepend ingontheg row ths tages .A lthoughfeed has beenfo rmu la ted
andtes tedfo ra llthel ife stages o f m ilkfish ,t ilap ia , and sh rimp andfo r
var ious s tagesin some spec ies(see Chap te r7 ),these a re con tinuous ly
beingre fined based onrecen tresea rchf ind ings . Recen t da ta on nu trien t
requ iremen ts, and ava ilability o f new , cheap bu t good p rotein sou rces a re
impo rtan tin fo rma t
ioninre fining afeedfo rmu la .
The p rocessedfeedisin fluenced bythe qua lity o fthefeeding redien ts .
Somefeeds tu ffs show w idef luc tua t
ionsinthe ir nu t r
ien t con ten t dueto
seasona l and geog raph icalva r
ia tions . Feeds tu f fs shou ldf irst be ana lyzed
fo rthe i rp rox ima tecompos ition , bu ti fth isis no t poss ib le,feedcompos ition
tab les(FCT ) can be used as gu ideinfo rmu la tingfeeds .In some FCTs ,the
mois tu re con ten tisinc luded andthe amoun ts o f nu t r
ien ts a re exp ressed
eithe ron a d ry ma t
terbas is o ron asrece ived bas is.Info rmu la t
ingfeeds ,a
un ifo rm se to fva lues shou ld be cons isten t
ly used . Va lues a re exp ressed
eithe ron a d r y ma t
ter bas is,o rasrece ived bas is . To conve rtthe amoun to f
nu trien tsf rom d ry ma tte r bas isto asrece ived bas is, usethefo rmu la:

(100- % mo
istu
re)
% nu
trien
t(asrece
ived
) = % nu
trien
t(d
ry ma
tte
r bas
is)x
100
 CHAPTER4 Fo
rmu
lat
ion o
f Aqua
feeds 107
The reis no de f
initefeedfo rmulafo r any spec ies because o fthe many
environmen tal and phys io
logica
lfac torsinvo lved as we l
l as dif
ferences
in the ava i
lab i
lity offeed ingred ients f rom one loca lity to another.
Subs ti
tution o fafeeds tuf
fo rfeedstuffsinthefo rmu lais possible p
rovided
thatthef ina lformu lationis s imi
larin nu trient con tent andthe re are no
nega t
ivee f
fec ts on grow th and surviva l andinc reaseinfeed cos t
. Feeding
expe rimen ts have showntha tfavorab le resu l
ts a re ob ta
ined aslong as
theamoun tso fnu t
rien ts p
resent do no texceedtherecommended nu tr
ient
levels. Excess o f nutrients can be expens ive, and can cause de ter
ioration
of thecu lturesys tem. Max imumleve ls o finco rpora t
ion o fsomefeeds tuffs
in theformu lafor carn ivores, omn ivores , and herb ivores arelistedin
Table 4.9

T
able4
.9 Recommended max
imuminclus
ionleve
ls(%
)ofsome ma
jorfeeding
red
ien
tsina
p
ract
ica
ldie
tforf
ishandshr
imp

| F
ish Sh
rimp

Carn
ivore Omn
ivore
/ Carn
ivore Omn
ivore
/
Feeds
tuf
fs
Herb
ivore Herb
ivore

A
lfa
lfa mea
l 5 10 5 10
B
lood mea
l, sp
ray d
ried 10 10 10 10
Cassava
/Tap
ioca mea
l 15 35 15 25
Coconu
t mea
l 15 25 15 25
Co
rn g
rain mea
l 20 35 15
Corn g
luten mea
l 15 20 15 20
Co
ttonseed mea
l, so
lven
t ex
trac
ted 15 20 10 15
Co
rn d
ist
il
lers 10 15 10 15
D
ica
lcium phospha
te 3 3 3 3
Hydro
lyzedfea
ther mea
l 10 10 10 10
F
ish mea
l nol
imi
t nol
imi
t 20 35
F
ish p
rote
in concen
tra
te 15 10 15 15
Ground mea
l, so
lven
t ex
trac
ted 15 25 15 25
L
ive
r mea
l 50 50 25 20
Mea
t and bone mea
l, 20 25 15 20
Pou
ltry by
-produc
t mea
l 15 20 15 20
Rapeseed mea
l, so
lven
t ex
trac
ted 20 25 15 20
R
ice b
ran
, so
lven
t ex
trac
ted 15 35 15 35
Sh
rimp mea
l 25 25 nol
imi
t nol
imi
t
Squ
id mea
l nol
imi
t nol
imi
t nol
imi
t nol
imi
t
So
rghum mea
l 20 35 15 35
Soybean mea
l, so
lven
t ex
trac
ted 25 35 20 30
Soybean mea
l,fu
llfa
t 35 40 20 30
Whea
tgra
in mea
l 20 35 20 35
Whea
tbran 15 30 15 30
Whea
tglu
ten mea
l 15 15 20 20
Whea
t m
idd
lings 25 40 20 35
Whey 10 10 10 10
Yeas
t(Brewe
rs)
,dr
ied 15 15 15 15

Sou
rce
: Tacon 1988
108 CHAPTER 4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

There a re seve ra
l me thods o ffo rmu la
tingfeeds . These are Pearson’s
Square Me thod , A lgebra ic Equa tion, Tria
l and Error , and L inear
Program m ing. The Pearson ’s Square me thod is recom mended in
formu lat
ing a supp lemen talfeed w ith onlytwotofouring redients. The
Tria
l and Er ror me thodis gene rally usedin calcu la
ting aformu lafor a
comp lete d
ietw ith manying red
ien ts.Incomme rcialfeed product
ion whe re
cost is a pr inc ipa l cons iderat ion, a computer program (L inear
Programm ing ) is used to comb ine feed ingred ients that wil
lg ive an
ef
fectiveformu la t
ion w i
ththelowes t cos
t.
Ma thema tical calcula t
ions us ingthese me thods, exceptfor L inear
Programm ing ,a re g
iveninthefo llowing examp les
.

I
. Pea
rson
’s Squa
re and A
lgeb
raic Equa
tion Me
thods

Examp
le 1
. When on
ly twofeed ingred
ients are to be comb
ined
.

To determine the amount of each ingredientin a supp lementa

lfeed
that wou ld contain 25% protein using Fish Mea l (FM) and Rice Bran
(RB), wi
th protein contents of 63% and 10% , respect
ive
ly.

□ Pea
rson
’s Squa
re Me
thod
:

a
) Firstd raw a squa re and writethe des ired p roteinlevelo fthefeed
atthe cen tero fthe squa re
.
b
) Wr i
te the twoingred ients w i
th the ir respec t
ive pro te
in con ten
ts
on each comer o fthele fts ide o fthe squa re,theingred ientw i
th
higher protein onthe uppe r and w i
thlowe r pro te
in onthelowe r
left hand co rner.
c
) Sub tract the des iredlevelo f proteinf rom eachingred ient and
write the difference on the r igh t corner o fthe square tha tis
diagona l
ly oppos i
tethe pro teinleve lo f eaching redient
.
d
) Ge tthe sum o fthe numbe rs a tther ights ide o fthe squa re
.
e
) De terminethe pe rcentage of eachingred ien t neededforthefeed
formu la by d ivid
ingthe numbe rs wr i
tten onthe r ight hand side
bythe sum o fthe dif
ference mu lt
ipl
ied by 100 .

Abou t 283 g FM and 717 g of RB are requiredto make a kg o fthe

supplementalfeedthatwi
llcontain 25% pro te
in. To checkforthe amount
ofprote
ininthefeed ,the pro
tein content(in percent)o feachingredien
t
 CHAPTER 4 Fo
rmu
lat
ion o
f Aqua
feeds 109
is multip
lied bythe amountof FM and RB(283g and 717grespec t
ively
)
to be comb ined
. The amounto f pro
tein com
ingfrom eachingredientis
addedto ge tthetota
l amount
. Theto ta
l amoun
tis d
ivided by 1000 and
mul
tipl
ied by 100to give 25% p
rotein.

FM = 283gx 63% = 178

.3 g pro
tein
RB = 717gx 10% = 71
.7 g pro
tein
To
talp
rote
in 250
.0 g /kg o
r 25% p
rote
in

o
r (250 / 1000
)x 100 = 25%

□ A
lgeb
raic Equa
tion Me
thod
:

Le
t: x =g FM/ kgfeed
y =g RB/ kgfeed

sothat:
x +y = 1000gfeed (Equa
tionI
)
0.63 x + 0
.10y = 250 g pro
tein/1000 gfeed (Equa
tionI
I)

Mul
tip
ly Equat
ionI by 0.10
:
0.10x + 0
.10y = 100g (Equa
tionI
II)

Sub
trac
t EquationI
IIf
rom Equa t
ionII:
0 .10y = 250 g pro
.63 x +0 tein/1000gfeed
-(0.10x+0.10y = 100g)
.53x +
0 0 = 150
X = 150/0.53 = 283 g FM

Subs
titu
tein EquationI
:
283 + y = 1000
y = 1000- 283 = 717g RB

Examp
le 2
: Whenth
ree o
r mo
refeeding
red
ien
ts a
re ava
ilab
lefor use
.

Formulate afish d
iet to con tain 35% pro tein by comb
ining the
fo
llowingingred
ients wi
th their respect
ive pro
te in con
tent
:

FM - 60% prote
in
Soybean Mea
l (SBM
) - 45% prote
in
RB 8% pro
tein
Com Meal (CM) 12% pro
tein

Assumethe proport
ion o
f 1pa rt FMto 2 parts SBM or1 :2andequa l
parts o
f RB and CM or1:1
. FM and SBM a rethe ma in sources o
fp ro
tein
while RB and CM arethe ma in sources o
f carbohydrate or energy but
also conta
in pro
tein
.
110 CHAPTER4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

□ Pea
rson
’sSqua
re Me
thod
:

a
) Draw a square and wr
ite a
tthe cen
terthe des
ired p
rote
inleve
l as
inthefirst example
.

b
) Calculatethe p
rote
inlevelf romthe pro
tein sou
rces FM and SBM
.
accordingtothe spec
ified rat
io: 1
:2

FM : 1 x 60 = 60
SBM : 2 x 45 = 90
Ave
rage: 150 /3 = 50

c
) Ca
lcula
te the ene
rgy sou
rces
, RB and CM a
lso accord
ingtothe
ra
tio 1
:1

RB : 1x 8 = 8
CM : 1x 12 = 12
Ave
rage: 20 /2 = 10

d
) Write the ca
lculated average protein con
tent on the upperlef
t
hand cornerforthe pro
tein sou
rces andthe protein conten
tofthe
energy sou
rces onthelowe rleft cornerofthe square.

e
) Wr i
te the desi
red pro teinlevela tthe cen ter and subtract th is
valuefromthe p rote
in con ten
to f FM and SBM and p ro
tein content
of RBand CM . Wri
tethe numbe rdiagonallyoppos i
tetheing redients
or onthelowe rright hand co rnerfor protein sources and uppe r
right hand cornerfo r energy sources
. Proceed asin d ) and e)o f
Examp le 1
.

f
) Mu l
tip
lythefina
l percentage de
rivedforthe pro
teinsources by 1/3fo
r
FM and 2/3for SBM. Fo rtheene rgysources
, mult
iplyby 1/2eacho f
the RB and CMtof ind outtheexac t amounto feachingredien
tto be
usedintheformu la w
iththe desiredleve
lo f35% prote
in.
 CHAPTER4 Fo
rmu
lat
ion o
f Aqua
feeds 1
11
Pro
tein sources = 62
.5%
In % In g/kgfeed
FM = 62.5x 1
/3 = 20
.83% 208.3
SBM = 62
.5x 2/3 = 41
.67% 416.7

Energy sources = 37
.5%

RB = 37
.5% x 1/2 = 18
.75% 187.5
CM = 37
.5% x 1/2 = 18
.75% 187.5
To
tal 100
.00% 1,000

g
) To check tha
tak
ilo o
ffeed con
tains 35% pro
tein
, proceed as
fol
lows:

FM 208
.3 gx 60% pro
tein = 125
.0 g pro
tein
SBM = 416 .7 gx 45% pro
tein = 187
.5 g pro
tein
RB 187
.5gx 8% pro
tein = 15
.0 g pro
tein
CM 187
.5gx 12% p
rotein = 22.5 g pro
tein
To
tal pe
r 1000 gfeed 350
.0 g pro
tein/kg o
r 35%

□ A
lgeb
raic Equa
tion Me
thod
:

Separateingredien
tsin to prote
in and energy sources and calculate
average protein contribution of each group accord ing to specified
propor
tions asin Example 2let
ters a
),b) andc
) Pearson’s Squa
re me thod.
Then:

Le
t: x = g o
f FM and SBM ofp ro
tein sources/ kgfeed
y = g o
f RB and CM as energy sources/ kgfeed

X + y = 1000g feed (EquationI

)
0
.50 x + 0
.10y = 350 g pro
tein/ kgfeed (Equa
tionII
)

Mul
tip
ly(
I) by 0
.10: 0
.10 x + 0
.10y = 1
00 (Equa
tion I
II)

Sub
trac
t(I
II)from(II
):
0.50x + 0
.10y = 350
-(0.10x + 0.10y = 100
)
0.40 x + 0 = 250
X = 625

FM : 625x 1/3 = 208

.3 g
SBM : 625x 2/3 = 416
.7g

F
rom Equa
tion(
I)

y = 1000- X
y = 1000- 625 = 375

RB = 375/2 = 187
.5g
CM = 375/2 = 187
.5g
112 CHAPTER4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Examp
le 3
: When somefeed components aref
ixed
.

Prepare af
ish dietto conta
in 32 % prote
in us
ing FM(60% pro te
in)
and Copra mea l or CpM (20% prote in) as prote
in sources to be
inc
luded in the diet
:

Rice bran (10% pro

tein
) = 22%
Mineral mix = 2%
Vitamin mix = 1%

□ Pea
rson
’s Squa
re Me
thod
:

a
) Determinethe amoun tof FM and CpM per 1 kgfeed mix
tu re
, by
subtract
ing the amoun ts o
f RB, mineral m
ix and vitam in mix
from 1000.
1000- (220 + 20 + 10
) = 750g/ kgfeed

b
) Vitamin and mineral mixes do not contain prote
in, but 220 g RB
suppl
ies 22 g p
rotein. There
fore, subtract
ing 22 g prote
infrom RB
fromthe 320 g des ired prote
inleve l equals 298 g pro
tein, wh
ich
must comefrom 750 g o f FM and CpM .

c
) Conver
tto percentagethe amounto fprote
in(298 g/kg
) and so
lve
bythe Pearson’s Square method the amoun
ttha t shou
ld come
fromthe combinedlevels o
f FM and CpM(750 g)
:
298/750 x 100 = 39 .73%

d
) Fol
low steps a
)to e
)in Examp
le 1 and checkforf
ina
l pro
teinin
one kg d
iet
:

e
) Atota
lo f 370 g FM (222 g p
rote
in) and 380 g CpM(76 g protein)
g
ives 298 kg p
rotein
. Th
is amoun tofprotein p
lusthe 22 g p
ro te
in
f
rom 220 g RB makes ato talof 320 g pro
tein o
r 32% of one kg
d
iet
.
 CHAPTER 4 Fo
rmu
lat
ion o
f Aqua
feeds 113
□ A
lgeb
raic Equa
tion Me
thod
:

Befo
re proceedingtoformulatethe necessary equat
ions,f
irs
t do
s
teps a
) and b) desc
ribedinthe Pea
rson’s Square method
.

Le
t: x = go
f FMrequ
ired
y = go
f CpMrequi
red

x +y = 750g (Equa
tion I
)
0
.60x + 0
.20y = 298g p
rote
in (Equat
ionII
)

Mu
ltip
ly (
I) by 0
.2:
0
.20x + 0 .2y = 150 (Equa
tionI
II
)

Sub
trac
t(I
II
)f rom (II
):
0
.60x + 0 .20 y = 298
-(0 .20 x + 0.2 y = 150)
0
.40 x + 0 = 148
x = 148 /0
.4 = 370g FM

Subs
titu
te(
I)
:
370 + y = 750g
y = 380g CpM

I
I.T
ria
l and E
rro
r Me
thod

Forthe Tr
ial and Error method
, a workshee
t and atab
le o
f prox
ima
te
composi
tion are necessa
ry.

Examp le 1
: Formu late a diettha tw i
ll conta
infish mea l(FM )
, defat
ted
soybean meal(DSBM ), mea t and bone mea l(MBM )
,r ice bran(RB ), and
vitamin and mineral mixes
. Codl ivero i
l(CLO )isthel ipid source
,b read
flour(BF)isthe sou rce ofcarbohyd rate and b
inder
, andl imeis used as
fil
ler
. Thefinished d ie
t shou ld conta
in 35% crude pro tein, 12% crude
fat,w i
th a d
ietary energy o
f 340to 400 kca l/100 g d
ie t
.

The prox
ima
te compos
ition o
fthesefeeding
red
ien
ts a
re:

Ingred
ien
ts g/100g
Protein Fat Fiber Ash NFE
FM 66.70 9
.11 0
.59 13.36 10
.24
DSBM 50.34 1
.45 8
.59 7.64 31
.98
MBM 46.91 10
.90 1
.11 36
.10 4
.98
RB 14.80 12
.66 4
.17 8
.92 59
.45
BF 14.17 1
.54 0
.56 0
.68 83
.05
114 CHAPTER 4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Thefo
llow
inging
red
ien
ts a
reinco
rpo
ratedinf
ixed amoun
ts:

FM = 10%
SBM = 18%
BF = 5%
V
it/min mix = 3%
To
tal = 36%

The remain
ing percen tage
, wh ichis 64%,w i
ll be suppl
ied by MBM, RB,
CLO, andthefil
ler(l
im e)
. About 58 g wi
llcomef rom MBM and RB and 6 g
froml ip
id andf i
ller
. The c losest es
timates tothe requi
red values a
re
obta
ined bytrial and error
.

1
.Fil
linthe requ
iredingred
ients and the correspond
ing amoun
tsin
Co
lumns 1and 2inthe Worksheet(Appendix A).

2
. Fi
llin the respective nutrient content based on the proximate
compos
ition ofeachingredient onthe upperleft hand comer o
feach
box(Worksheet1).

3
. Calculatethe amounto f nu
trien
tf rom eachingred
ient by mu l
tiply
ingthe
speci
fied amoun toftheingredient(column 2 ) bythe percentages ofthe
nutr
ien t(co
lumn 3 )toobtainthe amoun tingram s
.S tartwiththe p ro
tein
sources.
Fo rexamp le
, 10g FM with 66.7% prote
incon tr
ibu tes 6
.67gp rotein
tothe d ie
t. Add upthe proteinfrom FM , SBM , and BF(Wo rksheet
1
, column 3 )
. Dothe samefo rtherest.

4
. Calcu
latethe amoun tofp roteintha tshou ld comef rom MBM and RB
bysubtrac tingthe known p roteincon tribut
iono fthe othering
redients
(FM+DSBM+BF )fromtheto tal prote
in requ i
red(35% ).
35 g protein-(6 .67 + 9
.06 + 0 .71) = 18.56 g p
rote
intocomef rom
MBM +RB
Th
is va lue should comef rom 58 g o f MBM + RB , whichis 18.56/
58 = 32% p rote
in

5
. Use Pearson
’s Squa
reto ca
lculatethe exac
t amoun
tof MBM and RB
to be usedinthefeedformulat
ion:
 CHAPTER 4 Fo
rmu
lat
ion o
f Aqua
feeds 115
6
. F
ill
-up Workshee
t2 w
ith ca
lcu
lated amoun
ts o
f MBM(31
.07 g
) and
RB(26.93 g
).

7
. Ca
lcu late nu trient contributionf rom eachfeeding redient.Afterthe
amoun ts ofp rote
in sou rces have beenca lculated
,p roceedtoca lculate
thel ipid sources andl ipidlevels.
Fo rlipids, the partial amoun tfrom o thering redientsis equalto
8.05% , therefore
, to mee tthe requ i
remen to f 12% , CLO wil
l be
added a t3 .95% (co lumn 4 ). To make the formu lat
ion 100% ,
calcu late allthe othe r nu tr
ients and de termine how muchf i
ller
orl imeto add by sub tract
ing 97 .95from 100 equa ls 2
.05
.

8
. Calcu
latethe sum o ffibe
r, ash and NFEto have acomple
tereco rd o
f
the major nut
rients. Indicatethese va
luesin columns 5,6 , and 7,
respect
ively
. These can be compared to the ana lyzed proximate
composit
ion of processedfeed.

9
. To de
termine the d
ie tary energy/100 g d
iet use the phys
iolog
ica
l
va
lues o
f4.5,8 ,3.3 asfollows
:

Prote
in = 35 g x 4
.5 kca
l/g = 157
.5 kca
l
Fat = 12 g x 8
.0 kca
l/g = 96 kcal
Carbohydra
te = 28
.5g x 3
.3 kca
l/g = 94 kca l
347
.5 kcal/100g d
iet

W
ork
she
et1
Ca
lcu
lated nu
trien
tcompos
it
ion o
ffeed
Fe
edformu
lat
ioncode:M-1
Da
tecomputed:Ma
rch8,2000
Fo
rmula
tedby: Mary C
ruz

Ing
red
ien
ts g
/100g Protein% Fat% Fibe
r% Ash% NFE%
(1
) (2) (3) (4) (5
) (6) (7
)
66.7 9
.11 0
.59 13
.36 10
.24
F
ish Mea
l 1
0 6
.67

Mea
tandBone M
eal

50
.34 1
.45 8
.59 7
.64 31
.98
So
ybean mea
l(de
fat
ted
) 1
8 9.06

R
iceb
ran

14
.71 1
.54 0
.56 0
.68 83
.05
B
readf
lou
r 5
0.71

V
itam
in/m
ine
ralm
ix 3

Codl
ive
roi
l

L
ime

35.00
16.44
18.56
TOTAL 36
116 CHAPTER 4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Wo
rkshee
t2
Ca
lcu
lated nu
trien
t compos
it
ion o
ffeed
Feedformu
lationcode :M-1
Datecomputed: March 8. 2000
Formula
tedby : Mary Cruz

Ing
red
ien
ts g
/100g P
rote
in% Fa
t% F
iber% Ash% NFE%
(1
) (2) (3
) (4
) (5
) (6) (7)
911 .59 1336 1 0.2 4
F
ish M
eal 1
0
6
.67 0
.91 0
.06 1
.34 1
.02
46
.91.
1
0
9 1
.11 36
.1 .4
8
9
Mea
tand Bone M
eal 31
.07
14
.57 3
.39 3
.34 11
.22 1
.55
50
.34 1
.45 8
.59 7
.64 31
.98
So
ybean mea
l(de
fat
ted
) 1
8
9
.06 0
.26 1
.55 1
.38 5
.76
14
.8 12
.66 4
.17 8
.92 59
.45
R
iceb
ran 26
.93
3
.98 3
.41 1
.12 2
.40 1
6.0
1
14
.17 0
.68 83
.05
B
readf
lou
r 1.54 .0
6
5
5
0
.71 0
.08 0
.03 0
.03 4
.15

V
itam
in/m
ine
ralm
ix 3

100
Codl
ive
roi
l 3
.95
3
.95
100
L
ime 2
.05

TOTAL 100 35
.0 1
2

Examp le 2
: Formu late a d
ie tthat would conta
in no
tlessthan 40% pro tein
wi
th dietary energy content of no
tlessthan 340 kca l/100 g d
ietw ith the
follow
ing feed ingredients. Cons ider a
lso the d
igestib
ili
ty coef
fic
ients o f
the ma jor nutr
ients, protein,lipid
, and carbohydrate

Use FM and SMin a 1 :1 ratio

Lip id sourceis codliver o
il, with to
taldietaryfa
t notto exceed 15%
Carbohydrate sourceis bread f lour
Vitamin mix 2% , Mineral mix 1% , and Lecith
in 0.5%
Filler is ce
llu
lose

The prox
ima te analys
is and diges
tibi
lity coeff
icients of pro
tein
,lip
id, and
carbohydrate sources on a dry mat
ter bas is
:

% Nu
trien
ts % D
iges
tib
ili
ty

Pro
tein L
ipid Carbo­ Crude Crude Pro
tein L
ipid Carbo­
hydra
te f
iber ssh hydra
te

Fish meal 65
.8 5
.9 7
.8 0
.8 19
.7 80 93 94
Shrimp meal 68
.6 3
.7 7
.5 3
.6 16
.6 75 96 97
Squid meal 78
.5 5
.5 6
.8 1
.2 8
.0 88 91 89
Breadflour 13
.9 1
.2 83
.9 0
.3 0
.7 75 90 95
 CHAPTER 4 Fo
rmu
lat
ion o
f Aqua
feeds 117
Thefixed amountfrom v
itamin and m
inera
l mixes andlec
ith
in,is 3
.5%.
The remaining amoun
tis equalto 96.5% and wil
l be suppl
ied bythe
othe
ring red
ients
.In Worksheet3:

1
. Fi
llinthef
ixingred
ien ts andtheco rrespond
ingamoun tsin Workshee
t
3
. Then,try a 1
:1 ratio of 25 g each of FM and SM and wr
itethese
numbers onthe wo rksheet.

2. Fi
llinthe nut
rientcompos i
tiono fFM and SMin Wo rksheet3
,co lumns
3to 7 onthele ft hand boxfo r each co
lumn cor respondingto each
nutr
ient
. Calcula
tethe nu trient contr
ibut
ion based onthe p roximate
composition and digest
ibil
i ty coe
ffic
ien
ts andf i
llin the respective
box.
Compu te the protein
,l ip
id , and carbohydra te contributed by a
feedstuff usingthefo rmula:

We
igh
toffeeds
tuf
fx %P
rote
in X %D
iges
tibi
lity
100 100

F
rom the prox
imate compos
itiontab
le,
FM: 25g x 0.658 x 0.80 = 13.16 g prote
in
25g x 0.059 x 0.93 = 1 .37gl ipid
25g x 0.078 x 0.94 = 1 .83 g carbohyd
rate
Dothe samefo
r SM
.

3
. Ca
lcu
latethe amoun
tofp
rote
into be con
tribu
ted by SqM as
:

13
.6g(FM
) + 12
.7g(SM
) = 26
.3g

40g (
tota
lpro
tein
)- 26
.3g(FM + SM
) = 13
.7 g SqM p
rote
in

To ca
lcu
late we
igh
tof SqMto be usedinthefo
rmu
la:

We
igh
tof SqM = ( SqM p
%Pro
ro
te
te
in
in/100 x)
(
% D
iges
tib
100
ili
ty
)
13
.7g
= = 19
.8g SqM
(0
.785 x 0
.88
)

4
. Fi
llinthe ca
lcula
ted amounts o
fp ro
tein,l
ipid
, and ca
rbohyd
ratef
rom
19
.8g SqM(usingtheformu
lains tep 2
).

5
. Sum upthe d ietarylip
idfrom allthe prote
in sources and subtract
thevaluetoca lcu
latethe amountof CLOto be added. Assume d
ietary
l
ipidto be about 12%[12-(1 .37 + 0
.89 + 0
.99
) =8 .75].

6
. Sum upthe d ie
tary energy(use energy valuesfor each nu t
rientin
the prev
ious example
)a tthis po
in tto be ab
letocalculatethe amount
ofenergyto be contr
ibu ted by breadflour(340 kcal- 292 .5 kcal=
47.5 kca
l):
118 CHAPTER 4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Prote
in : 40g x 4
.5 kca
l/g = 180
Lip
id : 12g x 8
.0 kca
l/g = 96
NFE : 5g x 3.3 kca
l/g = 16.3
292
.5 kca
l

7
. Breadf louris about 90% carbohydrate with a d
igestib
ili
ty o
f about
95% , so tha
t 47.5 kcaldivided be 3.3is abou t 14.4 g
.S ince the
digest
ibil
ityisless than 100% , the amoun to f breadflour maybe
increasedto 16 g. Ca
lcula
tethe nu tr
ients con
tributed by 16g bread
f
lour(asin s tep 2
).

8
. Dete
rminethe sum o fing
redients used(25+25+19 .8+16+3+0
.5+8
.75
=98.05
) andsubtrac
ttheva luef
rom 100 .The d
iffe
renceistheamount
o
fthefil
ler
, ce
llu
lose(1.95)to maketheto ta
l equa lto 100
.

9
. Add upthe nu
trien
ts and ca
lcu
latetheto
tald
ieta
ry ene
rgy /100g d
iet
.

Prote
in : 41
.5 gx 4.5 kca
l/g = 186
.8
Lip
id : 12
.67g x 8
.0 kca
l/g = 101
.4
Carbohydra
te : 17
.83g x 3
.3 kca
l/g = 58.8
347.8 kca
l/lOOg

10. Compu tethetota

l ash andfiberconten
tinthefeedfo rmu la(co
lumn 5
and 6). The min
imum dietaryleve
lforasho rfiberisabout 10%because
higherlevelscancausepoorg rowthandsu rv
ival
. Comparethecompu ted
values to the actual resu lts ofthe chem ical analys
is to de
tect
discrepancies
.

Wo
rksh
eet3
Ca
lcu
lated nu
trien
t compos
ition o
ffeed
Fe
edformu
lat
ioncod e:M-1
Da
tecomput
ed:March 10. 2001
Fo
rmula
tedby: Marv Cruz

Ing
red
ien
ts g
/100g P
rotein% Fat% Fibe
r% Ash% NFE%
(D (2) (3) (4) (5
) (6) (7
)
65.8 5
.9 0
.8 19
.7 7
.8
F
ish Mea
l 25
13
.2 1
.37 0
.20 4
.92 1
.83
68
.6 3
.7 3
.6 16
.6 n\5“
Sh
rimp mea
l 25
12
.9 0
.89 0
.90 4
.15 2
.0
78
.5 5
.5 1
.2 8
.0 6
.8
Squ
id mea
l 19
.8
13
.7 0
.99 0
.24 1
.58 1
.2
13
.9 1
.2 0
.3 0
.7 83
.9
B
readf
lou
r 16
.0
1
.7 0
.17 0
.05 0
.11 12
.8

V
itam
in/m
ine
ralm
ix 3
.0

100
Lec
ith
in 0
.5
0
.5
100
Codl
ive
roi
l 8
.75
7
.5
Cel
lulose 1
.95

TOTAL 100 41
.5 12
.67 1
.39 10
.76 17
.83
 CHAPTER 4 Fo
rmu
lat
ion o
f Aqua
feeds 119
I
II.L
inea
rProg
ramfo
r Leas
t-Cos
t Fo
rmu
lat
ion

Linearp rogramm ingis used when manying red

ien tsa retobecomb ined
for aleas t cos t feed formu la. Th is me thod is espec ially use fulin
comme rcia lfeed manu facture whe re
inla rge quan t
ities o ffeeding redients
are used . A compu teris necessa ryinth istype o ffo rmu lat
ion . When one
or mo reing redien ts are no t available,otherfeeding redients a re ut
ilized
as subs ti
tu tesin order to come up w ith the samefeed qua l
ity. The
quan t
i ties o f subs t
itu teingred ients are de term ined us ing thel inear
programm ing me thod . In forma t
ion onthe amoun ts o f nutrientsin each
ofthefeedingred ientsto be usedis essen t
ialinth is me thod . This can
be obta ined f rom afeed compos i
tion table or chem ical ana lysis.A
compu ter prog ram w il
ll ist several combinationsto come up w ith almos t
simi
larfeed qua l
ity a t alesse r cos t
. Examp les o fcompu terp rogramfo r
least cos tformu lation a rethe s imp lelinear programm ing spreadshee t
andthe soph is t
ica ted Br il
l Formu lation Package.

Pu
ri
fiedD
ietFo
rmu
lat
ion

In nu tr
itional requiremen ts tudies, purif
ieding red
ien ts are usedinthe
formu lat
ion o fdiets. Th
isis necessa ryto s tudythe op timumleve lofone
speci
fic nu t
rient,e.g.essential am ino ac
id. The useo fpu r
if
ieding red
ients
enables oneto varytheleve lo fthe nu t
rien t whose dietaryrequ irement
isbeings tudied whi
le keepingtheo ther nutrientleve
lscons tant
. Examples
of purif
iedingred ientsinc lude case in and ge latin as pro te
in sources,
dextr
in and suc rose as ca
rbohyd rate sources, and cellulosefo rthe non­
nutri
tivef il
ler.

Examp le: Formu late a pur

ified dietto contain about 50% pro te
in us ing
case in and ge latin at 1:1 ratio, cod liver oillevela t 12% . and v i
tamin
and minera l mixtures a t 3% each , with a dietary energy o
f 380 kca l/100
gd iet. Assume case in and ge latin to contain 93% pro tein with apparent
prote in digest
ib i
lity of 96% , wh ile dextrin is 100% d igestible
. Inc lude
feed add itives such as a ttractant a t 1% and b inder at 2% .

Feedingred
ien
ts Percen
tage

Gelatin
Case in
Dex t
rin
Codl ivero il 12
Vi
tam in m ix 3
Minera l mix 3
Feed b inder 2
Attractant 1
120 CHAPTER4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Fol
lowingthecompu tat
ionin Examp le 2intheTrial and Erro
r Me
thod
sect
ion,the amoun tofcasein and ge
lat
in would be 28 g each
. Thetota
l
die
tary energy con
ten
to fthe above die
tis:

50 g prote
in x 4.5 kca
l/g prote
in = 225
12gl ipid x 8.0 kca
l/glipid = 96
Total 321 kca
l/100 g d
iet

The amoun to fdextrin wi

ll make uptherema in
ingene rgyrequ i
rement.
Ifthetotaldietaryenergycon tentis 380 kcal/100g d iet
,thenthe amoun t
of dex
trinto be added shou ld supp lythe ene rgy di
fference wh ichis 380
kcal- 321 kca l = 59 kcal.S ince a gram ofca rbohyd rate has an energy
contentof3 .3 kca
l,the refo
re . 59 kca
ld iv
ided by 3.3 kca l/g dextr
inis 18
grams. To maketheformu la 100% , afil
ler, such as ce llu
lose, can be
used. Thefinal compos it
ion wou ld be:

Feedingred ients %
Gelatin 28
Case in 28
Dex t
rin 18
Codl ivero il 12
Vitam in m ix 3
Minera l mix 3
Feed b inde r 2
Attractan t 1
Cellulose( fi
lle
r) 5
Total 100

Summa
ry

Feedis a veryimpor tant componen tin the success o f an aquacu lture

venture . W ith increased s tock ing densities, na tural food in cu lture
systems hasto be supp lemen ted withformu latedfeeds .Itisimpo rtan t
to know the nu tr
ients requ ired by the aquacu lture species as these
would bethe s tar
ting po intinfeedfo rmulation. As inglefeeds tuff does
not have alltherequ i
red nu trientsfo rgrowth, su rv
ival, andreproduc tion .
Hence,i tis necessary to know the nutr ien t conten t and levels in
common ly usedfeeds tuffs so tha tthese can be comb ined to come up
wi
th the des i redformu lation by ma thema t
ica l ca
lculations.
Seve ral ma thema tical me thods can be usedinfeedfo rmulation: the
Pearson 's square techn ique , the a lgebraic equa tion me thod, the t rial
and e r
ro r me thod, andl inear p rogramm ing . Thef i
rsttwo a re simp le,
theth ird uses a wo rkshee t, andthefou r
threqu ires acompu tersoftwa re.
In balanc ing ara tion,p ro tein wh ichisthe ma jor componen to fthe d ie
t
is compu tedf irst
,the ene rgy( lip
id and ca rbohyd rate)levels ofthe d ie
t
arethen ad jus tedtothe des ired dietarylevel.
 CHAPTER 4 Fo
rmu
lat
ion o
f Aqua
feeds 121
Gu
ide Ques
tions

1
. Name some me
thodsinformu
lat
ing d
iets
. G
ivethe advan
tages and
d
isadvan
tages o
f each me
thod
.
2
. Whyisi
timpor
tan
tto knowthe nu
trien
t con
ten
toffeeds
tuf
fs and
the
irleve
ls be
fore one can s
tar
tfo
rmu
lat
ing afeed?
3
. Whyis pro
tein thef
irs
t nu
trien
tto be cons
ide
redinformu
lat
ing a
d
iet?
4
. D
iscussfac
torsto be cons
ide
red andthe
irimpo
rtanceinfo
rmu
lat
ing
afeedfo
r a spec
ific spec
ies
.
5
. G
ive a
tleas
t 3feeds
tuf
fstha
tare good sou
rces o
f:
a
) pro
tein
b
)lip
ids
c
) carbohydra
te
6
. Ca
lcu
latethe amoun
toff
ish mea
landr
ice b
ranina d
iettha
tcon
tains
30% pro
tein
. The pro
tein con
ten
ts o
ffish mea
l and r
ice bran a
re,
60% and 15%
, respec
tive
ly.
7
. Use theingred
ien
ts in Examp
le 1in the Tr
ial and Error Me
thod
sec
tioninformu
lat
ing a d
iettha
t w
ill con
tain no
tless than 35%
pro
tein
,die
taryl
ipid o
f no
t mo
rethan 10% and d
ietary ene
rgy no
t
lessthan 350 kca
l/100g d
iet
.

Sugges
ted Read
ings

Baut
ista MN , Millamena OM , Kanazawa A . 1989. Use o
f kappa-
carrageenan m
icrobound d
iet(C-MBD
) asfeedfo
rPenaeus monodon
la
rvae . Mar
. Bio
l. 102
:169-174
.

Bo
rlonganIG, Mar
te CL, Noci
llado JN
. 2000
. Developmentoflarva
ldie
ts
for m
ilk
fish(Chanos chanos).J. App
l.Ichthyo
l. 16
:68-72
.

Ca
tacu
tan MR . Coloso RM . 1995
.E f
fectofdie
ta ry p
rote
intoene rgyrat
ios
on growth, survival
, and body composition ofjuven
ile Asian sea
bass
, Lates calcari
fer. Aquacul
ture 131:125-133.

Ca
tacu
tan MR . 1997.Protein and dry ma
tterdigestib
ili
ty offeeds
tuffsin
comple
te dietsfor Penaeus monodon. In: Zhou Y, Zhou H, Yao C
,
Lu Y
, Hu F , Cui H and Din F(eds )
. Proceedings of The Fourth
As
ian Fisher
ies Forum , Be
ij
ing, China, 16-20 October
. 754 p.

Cuzon G, Guil
laume J . 1991. Recom mendat
ions for pract
icalfeed
formulat
ion.In
: Castel
l JD and Co
rpron KE(eds)
. The Crus
tacean
News
letter
, Lou
isiana
, Wo r
ld Aquacul
ture Soc
iety
. p 52
-53.

Euseb
io PS . 1991
. Effec
to f dehulling on the nut
rit
ive value o
f some
legum inous seeds as prote
in sourcesfort iger prawn,Penaeus
monodon juveni
les. Aquaculture 99
:297 -308
.
122 CHAPTER4 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Euseb
io JS
, Eusebio PS. 1984
.Effec
tofprocessing onthe p
rote
in qua
lity
o
f mungbean(V ignaradia
ta).Kim
ika 3 :1-9
.

Feed Deve
lopment Sect
ion. 1994. Feeds andfeeding o
f m
ilk
fish
, N i
le
ti
lapia
, As
ian sea bass andtiger sh
rimp. SEAFDEC Aquacul
ture
Departmen
t,Tigbauan,Iloi
lo, Phi
lipp
ines
. 97 p.

Feed Technology and Nu tr

ition Wo rkshop
. 2001
. In: RA Swick and D
Ghazalay
-De la
ine (eds). Malaysia
, May 27 -30
. 2001. American
Soybean Associa
tion. p 30
.

Fe
rrar
is RP, Catacutan MR, Mabeline RL, Jazul AP. 1986. Diges
tib
ili
ty
in m i
lkf
ish, Chanos chanos (Forsskal)
:e ffec
to f pro
tein source,
fish s
ize
, and sal
ini
ty. Aquaculture 59:93-105.

Ha
lver J
. 1976. The nutr
itiona
l requirements o
f cult
iva
ted warmwater
and coldwater fish spec ies
. Paper No . 31, FAO Technical
Confe
rence on Aquacultu
re, Kyoto
, 26 May-2 June 1976
. 9p
.

He
rtrampJW , Pascua
lFP . 2000
. HandbookonIngredien
tsforAquaculture
Feeds
. K luwer Academic Pub
lishe
rs, The Ne
therlands
, 624 p
.

La
llS
. 1991 . Conceptsinthefo rmula t
ion and prepa ra
tion of a comp lete
fish die
t.In: De Silva SS
. (ed).Fish Nutrition Researchin As ia
.
Proceedings o
fthe Four th As
ian F ish Nutr
ition Wo rkshop . Asian
Fisher
ies Society Spec
ial Publicat
ion 5. Asian F isher
ies Soc iety,
Manila
, Phil
ippines.p 1-12.

L
ienerIE
. 1962
. Toxicfactorsin ed
iblelegumes andthe
ire
lim
ina
tion
.
Amer
.J.C l
in. Nu
tr. 11:281
-298.

L
iene
rIE . 1980
. Tox
ic cons
tituen
ts o
fplan
tfoods
tuf
fs. Academ
ic P
ress
,
New York
, 171 p
.

McE
llh
iney, RR(ed
). 1994. Feed manufac
turingtechnologyIV
. Ame
rican
FeedIndustry Assoc
iation
, Inc
. Ar
lington
,V i
rginia.

M
illamena OM , Primavera JH , Pudadera RA, Caba l
lero RV. 1986. The
ef
fecto fdiet onreproduc t
ive per
formance of pond-reared Penaeus
monodon Fabr icius broodstock. In
: Maclean JL , Dizon LB, and
Hosillos LV (eds ). The First Asian Fisheries Forum Man ila
,
Phil
ipp ines. Asian Fisher
ies Society
. p 593-596.

M
illamena OM ,Tr
iñoAT. 1997
. Low-costfeedfor Penaeus monodon rea
red
in tanks and under sem i
-intensive andintensive cond
itionsin
brackishwate
r ponds. Aquaculture 154:69
-87.

Na
tiona
l Research Council
. 1977
. Nutrien
t requi
remen ts o
f warmwa ter
fishes
. Nat
ional Academy o
f Sc
iences, Washing
ton DC . 78 p
.
 CHAPTER 4 Fo
rmu
lat
ion o
f Aqua
feeds 123
New MB. 1987
. Feeds andfeeding o
ffish and sh
rimp
. A manual onthe
prepara
tion and presen
tation o
fcompoundfeedfor shr
imps and
fishin aquacul
ture ADCP/REP/87/26 , UNDP/FAO
. 275 p.

Pena
florida VD. 1989. An eva
luat
ion ofind
igenous protein sources as
po tent
ial componen tin the d
ietformulation for t
iger prawn ,
Penaeus monodon, us ing essent
ial amino acid index (EAA I)
.
Aquacu lture 83
:319
-330.

San
tiago CB ,A
ldaba MB , Laron MA. 1983.Effec
tofva ry
ing d
ietarycrude
p
roteinleve
ls on spawn ingfrequency and growth o
f Saro
therodon
n
ilo
ticus breede
rs.F ish. Res
.J . Phi
lipp.8:9-18.

San
tiago CB,Aldaba MB, Abuan EF, Laron MA
. 1985 .Effec
ts ofart
ific
ial
d
iets on f
ry production and growth o f Oreochrom is n
ilo
ticus
breeders
. Aquacul
ture 47:193-203.

San
tiago CB, Reyes OS,Aldaba MB, La
ron MA . 1986. An eva
luat
ion of
formu la
ted die
tsfo
rN i
let
ilapiafinge
rlings
.F ish
. Res.J. Phi
lipp
.
11
:5-12.

Sumagaysay NS, Bor

longanIG. 1994.G rowth and product
ion ofmilkf
ish
(Chanos chanos)in b
rackishwater ponds:effec
ts o
fd ie
tary pro
tein
andfeedinglevels
. Aquaculture 132:273-283.

Tacon A
. 1988 . The nutr
ition andfeeding o
ffarmedfish and shrimp -
Training manua l3. Feeding Methods.Fie
ld Documen t No
. 7/E.
,
FAO -
Italy
. 208 p.

Tesh
ima S, Kanazawa A. 1983
. E f
fec
ts ofsevera
lfac tors on g
rowth and
surviva
lo fthe prawnlarvae reared w
ith m icroparticu
late d
iets
.
Bul
l. Jpn. Soc
. Sc
i.Fish. 49
(12)
:1893-1896.
 CHAPTER

5
P
rocess
ingofFeedstu
ffs
andAquafeeds
NELSON V
. GOLEZ

In
troduc
tion

Feed manu factur ing and qua litycon tro lofthef inishedfeed a reimpo rtan t
to success ful aquacu l
ture . The techno logy o ffeed process ing has
unde rgone cons ide rab leimp rovemen tsthroughthe yea rs. P rocess ing o f
feeds tu ffs and aqua feeds has progressedf rom s imp le m ixing o f seve ral
ingred ien ts by hand to mechan ica l m ix
ing , to con t
inuous m ixing , and
recen tlyto compu ter -con trolled process ing . In sp ite o fth is , the bas ic
concep to f mix ing theingred ien ts toge ther to ob tain a nu tritiona lly -
balancedfeed rema ins unchanged . Infeed process ing , seve ra lfac tors
haveto be cons ide red .
The produc tion o fgood qua l
ityfeedsrequ iresthe use o fing red ien ts o f
high qua l
ity. Thereis a lso a need to avo id the cond itions tha t cou ld
cause de teriora tion o fthefeeds tu ffsf rom harves tingto process ing and
storage . Manyfeeds tu f
fs con tain an t
inu tritiona lfac tors tha t preven t
uti
lizat ion o f nu t r
ien ts(pa r
ticu la rly p ro te
ins ) byf ish and o the r cultu red
anima ls. Fo r examp le, the seeds o f manylegumes con tain subs tances
tha tinac tiva te tryps in and chymo trypsin , the enzymes for pro te in
diges t
ionin an ima ls. The nu trit iona l quality o ffeeds tuffs dependsin
partonthe process ingthey unde rgotoremovethe an tinu tritiona lfac to rs.
Some o fthese fac tors are eas ily des troyed by hea t wh ile o thers are
resistan tto hea t.
Grow th and produc tivity o fthe aqua tic an ima ls grea t
ly re ly onthe
qua li
ty o ffeed g iven tothem . Feeds represen t a ma jor por tion o fto tal
cos tsin an ima l produc tion . Thus , process ing o ffeeding red ien ts,feed
prepara tion , and s torage are ma jor cons idera tions in mak ingfeeds .
Howeve r, prope r me thods app lied dur ing process ing and p repa ra t
ion w ill
be use lessi ftheend p roduc tis no tca re ful
ly hand led and p rope rlys to red .
Infeed prepara tion , the ma in ob jec t
iveis no tjus tto m ix wha tever
ing red ients a re ava ilab le bu tto p repa rethefeed us ing p rope rly p rocessed
ingred ien ts. O ften times , be fore these raw ma teria ls can be used , they
haveto be processedtoremove subs tancestha t may p reven tthe irp rope r
util
izat ion bythef ish . Thefeed prepa ra
tion p rocessis mo re conce rned
with the phys ica l convers ion o fag ivenformu la tionin to compounded ,
nutr itiona l
ly-e ffec tive d ie t. Feed s torage mus ta lso be g iven equa l
impo rtance asinfeed p repa ration and p rocess ing becausei ta f
fec tsthe
she lf
-life o fthef inished p roduc t.
126 CHAPTER5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

This chapterw il
l helpthereade r unde rstand and app reciatethe basic
principles ofp rocessing,p reparation,s torage, and qua l
ity controlinthe
prepara tion of aqua feeds. The ma ter
ialin th is sectionis presen tedin
sequence beg inning w ith the processing o f basicingred ients to remove
an t
inu trit
ionalfac tors,followed by s tepsinfeed prepara tion,from the
easiesttothe mo recomp lex processes, ands torage. Thischap terp resen
ts
methods and equ ipmen ttha ta re use ful not on lyforfeed m i
llers, but
alsofo r extension wo rkers andf ishfa rme rs.

Feeds
tuf
fProcess
ing

Feedstu ffprocessingre fersto all ope ra

tions necessa ryto ob tainthe highes t
nutri
tiona l value o f afeeds tuff andthe bes t econom icre tu rnsf romthe i
r
use. The re are various ways o fp rocess ingraw ma terialstha t can destroy
some an t
inu tr
itionalfac tors presentinfeeds tuffs
. Theya resoak ing, heat
ing ,
chem icalt reatmen t such as ex t
rac t
ion w ith o
rgan ic solvent, and dehu ll
ing .
The k ind o ftreatmen tw il
l depend onthe subs tances p resen tinthefeed
ingred ient
. Table 5.1l istssomean tinutrit
ionalsubs tances andthe irremova l
orinac t
ivation.
Some a re na tural componen to ffeeds tuf
fs such as gossypo lin co tton
seed and th iam inasein rawf ish . O thers may or igina tef rom na tura l
contam ination o fthe raw ma terials such as a flatoxins produced bythe
mold Aspe rgil
lusflavus, o r man-madecon taminan tssuchaspo lychlor
ina ted
bipheny ls(PCBs )
, pes tic
ides, he rbicides, and hyd roca rbons .

T
able5
.1Va
riousan
tinu
tri
tiona
lsubs
tancesinsomefeeds
tuf
fsandthe
irremova
lorinac
tiva
tion
Subs
tances Adve
rse ac
tions Occu
rrence Remova
lor remedy

Hea
tLab
ilesubs
tances
Tryps
in inh
ibi
tor B
indst
ryps
intofo
rm aninac
tive Soybean and o
the
rlegumes D
ry hea
ting a
t 175
-195°C
,or cook
ing
compound fo
r 10 m
in.
Lec
tins Des
troythered b
loodce
lls Soybean and o
the
rlegumes Bo
il
ingin wa
tero
rau
toc
lav
ingfo
r30 m
in.
Go
itrogens Inh
ibi
tthe up
take o
fiod
ine by Soybean and o
the
rlegumes S
team and
/or au
toc
lave
thethy
roid g
land for 10to 30 m
in
An
ti-v
itam
in D B
inds V
it.D
, mak
ingi
t Soybean and o
the
rlegumes Au
toc
lav
ing o
r bo
il
ingfor 30 m
in.
unava
ilab
le
An
ti-v
itam
in E Con
tribu
testo V
it. E de
fic
iency Soybean and o
the
rlegumes Au
toc
lav
ing
Th
iaminase Promo
tes des
truc
tion o
f Raw and spo
iledf
ish
, Au
toc
lav
ing
, hea
ting
, and cook
ing
th
iam
in(V
it
. B1
) musse
ls,c
lams
, and soybean
Hea
tres
istan
tsubs
tances
Es
trogens (
iso
flavones
) In
ter
fere w
ith rep
roduc
tive P
lan
tglycos
ides So
lven
t ex
trac
tion
per
formance
Gossypo
l B
inds phospho
rus and some Co
ttonseed mea
l Add
ition o
firon sa
lts o
r phy
tase
p
rote
ins
Tann
in B
inds p
rote
in andinh
ibi
t Beans and o
the
rlegumes Dehu
lling
t
ryps
in d
iges
tion
Cyanogens Re
lease po
isonous hyd
rocyan
ic Cassavaleaves Soak
ingin wa
terfo
r 12 h
ac
id
M
imos
ine In
ter
feres w
ith enzyme syn is Ip
thes il-
ipi
lleaves Soak
ingleavesin wa
terfor 24 h
inthel
ive
r; des
troys hepa
to­
panc
rea
tic ce
lls o
f sh
rimps
Pe
rox
ides B
ind p
rote
ins and v
itam
ins Poo
rly s
tored and Prope
rsto
rage
unp
rotec
ted o
ils
Phy
tates B
ind p
rote
ins and m
ine
rals Co
ttonseed mea
l, ce
rea
l hu
lls Dehu
ll
ing
and reducesthe
ir ava
ilab
il
ity soybean
, and o
the
rlegumes

Sou
rces
:Liene
r 1962
, 1980
; Euseb
io and Euseb
io 1984
; Euseb
io 1991
 CHAPTER 5 Process
ing o
f Feeds
tuf
fs and Aqua
feeds 127
Feeds
tu f
fs are processedto :
• remove an t
inu tr
itionalfactors andtox ins
• increase pa latabil
ity,digestibi
lity
, and nu tr
ient ava
ilabi
lity
• ad justfeed pa r
ticle s
izeto su i
tag iven spec
ies and s
ize
• reducefeed was tage
• max im ize pro
fitthrough op timum process ing o
ffeeds
• lowe rthe mo isture contento ffeedstuffto 10% orless
Ip
il
-ip
illea
f
D
if
feren
t me
thodso
ffeeds
tuf
fprocess
ing

Soak
ing
Somefeeds tu f
fs haveto besoakedin wa terfo r6 -24 hatroomtempe rature.
Soak ing is some times done w ith hea tto so ftenthe g rains tha t swe ll
duringthe process andfac i
litate remova lo fsometox ins(an tinutritional
factors)insomefeeding redien tssuch asip il-
ipi
lLeucaenaleucocepha lalea f
meal, cassavalea f mea l,e tc. Soak ing Leucaena leaves wh ich arer ichin
protein and some m ineralsre leases m imos ine, a non-protein am ino ac id
tha tis stableto hea t
. M imos inein terferes with enzyme p roduction and Cassavalea
f
destroysthe hepa topanc reas o f shrimps andl ivero ffish. Soak ingthe
Leucaena leavesfo r 24 hreducesthe m imos ine contentto an accep table
level. Incorpo rat
ion o fLeucaena lea f mealin sh rimpfeed shou ld no t be
more than 5%to avo id tox icity. Un treated cassavalea f mea l con tains
poisonous hyd rocyanic ac id. Cassavalea f mea l when soakedfo r6 -12 h o r
blanched w ith boiling wa terre leasesi ts cyanogens and p roduces a sa fe
cassavalea f meal.

Hea
ting and Cook
ing
Heatt reatmen tis app l
iedto d ryfeeds tu ffstoinac t
ivatesome antinu
tri
tional
facto rs andinc rease u til
iza t
ion o f nu tr ien ts. These an t
inu t
rit
iona
lfac to
rs
presen tinrawsoybeansando therlegumescan ma rked
lyaffec
ttheintest
inal
tracto fthe an ima ls and p reven td iges tion and
util
izat ion o f many nutr ients par ticu lar ly
proteinsandv itam ins.Thet imeandtempe ratu re
of hea ting afeeds tu ffis ofg reatimpo rtancein
obta ining a good qua l
ityfeed . In mo is t hea t
treatmen t, the samp leis bo iledfo r 30 m in a t
100 °C,p ressu recookedfo r 10 m in a t 120 °C ,o r
steamedfo r 30 m in.In d r
y hea tt rea tmen t
,the
samp leis sun -dr
iedfo r6 -12 h , oven -d riedfo r
12 h a t 60°C ,o rroastedfo r2 -5 m in at 250 °C
Soybean(1 )is ave ry good sou rce o fp ro tein
buti t con tains an an tinut r
itivefac tor ca lled
tryps ininh ibitor.T rypsin,anenzymetha td iges ts
proteinis des troyed bythet ryps ininh ib i
to rin
raw soybean mea l. Sun f
lowe r mea l(2)is a lso a
poten tialp rote in sou rce bu t con tains p rotease
inh ibitortha tis des troyed by hea t. Legumes ,
like mung beans(3 )
,and cowpea(4 )
, peanu ts(5)
and sesame seeds(6 )
,and ce reals such as com
kerne l(7 )
,r ice gra ins(8 ), and sorghum(9 )a re
poten t
ialingred ien ts bu t may con tain non -
128 CHAPTER 5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

nutritive componen ts i f no t proper ly

processed .
Pro tease o rtryps ininh ibi tors b ind
tryps intofo rm aninac tive compound .
Soak ing and dry hea ting a t 175 -195 °C
for 15 m in or pressure cook ingfor 10
min a t 120 °C inact ivates these
inhib itors . Lectinsinrawsoybean mea ls
are des troyed o rinac tivated by bo il
ing
in wa ter o r au toclavingfor 10 m inu tes
at 120 °C resu lting in a d igest ib le
soybean mea l. There are an ti-vitam in
D and E subs tancesin raw soybeans
thatb ind these v itam ins . Au toclav ing
or boilingfo r 10to 30 m inu tes des t
roys
these factors mak ing the v i
tamins
availab letof ish.
Th iam inase , an enzymein raw and
spoiledf ish( 1
0
), musse ls(11
), oys ters(1 2)
,
and c lams ( 13
) destroys the v itamin ,
thiam in andisinac t
iva ted by bo ilingin
water or au toclavingfor 30 m inu tes .
This me thod makesraw and spo iledf ish,
mussels, andc lams safefo ru se
.
Mo is t hea tis mo re effectivethan d ry
heatforlegume seeds . Mo ist hea t trea tmen ta lso improves the
digestib
ili
ty and nutr
itionalvalues o fg rains .I tis also usedto ex trac t
oilf rom o i
lseeds. L ikewise, dry hea t treatment increases the
nu t
ritional va
lue offeeds tuff bu t such trea tmen t makes hand ling
and pe lle
ting eas
ierthaninthe mo istt rea tmen t

Dehu
lling
Tannins a refound mos tlyin seed coa ts or hu
lls of colored beans(1 )
.
Tannins and phy tates bind proteins, m inera
ls, and v itam ins thus
reducingthe ava ilabil
ity ofthese nutr
ients. Dehull
ingremoves mos tof
thetann ins by split
ting or pee
lingthe softseed coat using a dehu l
lerto
separatethe hulls(2). This methodis done by useo fasp litt
ing machine
orbysoak ing 1pa rtoftheseedsbyvo lumein 3 pa rtso f waterfor 4to 6
h,followed by sundry ingfor 12 h o r oven drying a t 80 °Cfor 12 h .
Subsequen tly, sp
littingis done by us ing an ordinary co rn m i
llo r by
hand pressing. Ano ther wayisbysoak ingtheseedsin wa tero r 12to 48
h and allow ingthem to ge rm ina
te. The germ inated seeds a rethen
dehulled and sund ried.

Ex
trac
tion w
ith o
rgan
icso
lven
t and chem
ica
ltrea
tmen
t
Organic solvents are a lso used to remove the ant inutrit
iona l
componen ts o
fva riousfeeds tuf
fs. Ex t
raction o
flipidfromlegum inous
seeds such asrape seed(1 )
,and b lack beans(2)with a
lcohol and wa
ter
el
imina testhe beanyf lavor. Extract
ion ofo i
lfrom cottonseed meal(3
)
wi
th hexane dec reasesthetox in gossypol
.
 CHAPTER5 P
rocess
ing o
f Feeds
tuf
fs and Aqua
feeds 129

Feedp
repa
rat
iontechn
iques

To achieve goodfeed charac

ter
ist
ics
,feed mus
t be prepared bythe
fo
llowing procedures
:

1
. G
rind
ing increasesthe sur
face a
rea o
fing
red
ien
ts. Th
isimp
roves
mixing,d iges tibil
ity, pe lletability, and wa ter s tab i
lity offeeds tuf
fs .
The gr ind ing equ ipmen t var ies w ith the na ture and tex ture o f
mate r
ials andthe des ired pa rticle s ize. Feed componen ts no rma lly
comein d iffe rent particu late s izes and shou ld be un iform lyg round .
Mate r
ia ls w ith a w ide range o f pa rt
icle s izes do no t m ix we l
l and
produce s t ruc turally weak pe lle ts
.
The re a re a numbe ro fg rind ingequ ipmen t usedfo rsizereduc tion
offeeds tu f
fs .The hamme rm i
llisacommon lyused mach ine. Bas ically,
hammer m il
ls a reimpac td is in tegra t
ion mach ines composed o fa
high speed ro tat
ing sha ftw ithf ree sw ing ing hamme rs. The s ize o f
the g round ma ter
ialis con tro lled bythe s i ze ofthe sc reens o r me tal
bars moun ted ontheex it open ing usua llyfound a tthe bo ttom o fthe
mach ine . O the rg rinding mach inestha t can be usedfo rg r
ind ing Hammerorgr
inde
rsha
tte
ring
and s izereduc tion offeed componen ts a re:
a. Sw ingtype hamme rm ill-animpac tg rinde rw ith
sw ing ing or stat ionary s teel bars forc ing
ingred ients aga ins t ac ircu lar screen or so lid
serra ted sec tion des igna ted as a s t rik ing p late.
Mate rials a re heldinthe g rind ing chambe r un ti
l
theya rereducedtothes izeo fthesc reenopen ings.
This type o f hammer m ille ff
icien tly gr inds dry
andlow - fating redien ts.
b. Attr ition m i
ll - gr ind ing pr incip le is through
sha tte ring byimpac t. Howeve r,ita lsoimpa rts a
shear ing and cut ting ac t
ion . Gr ind ing is
accomp lished be tween two d iscs equ ipped w i
th
replaceab le wea ring su rfaces . Th istype o fg r
inde r Swingtypehamme
rmil
l
is usedfo rb lending and smoo thingou ting redients
ora m ix tu re(con tain ingl iqu id)tha t have c lumps .
c. Ro l
le r m ill-comb ine cu tting ,a ttr
ition , and c rush ing p rocesses .It
has smoo th o r corruga ted ro llers seta tap re-determ ined d is tance
apa rtro ta tinga tthesamespeedand w iththe ma terialpass ingbe tween
thero l
le rs . Anadd itionaltea ringac t
ion maybep rovidedbythebo ttom
rol
lla te ralco rruga t
ions o r by ope rat
ingthero l
ls atd ifferent speeds .
130 CHAPTER 5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

A
ttr
it
ion m
i
ll Ro
lle
rmi
ll

d
. Cu
tte
rs -reduce dry par
tic
le so
lids ma
inly by shea
ring w
ith kn
ife
edges against astr
iking plate. The mil
l cons is
ts ofaro tating shaf
t
w
ithfou r para
lle
l knives a
ttached and asc reenoccupy ingonefou rth
ofthe 360°rotat
ion. The m illisbes
tsu itedfo rcracking wholeg ra
ins
w
ith a min imum amoun to ffines
. Cu t
te rs should not be used as a
f
inalp rocessforreduc ingthe s i
ze o
fing redients usedin aqua feeds
.

2
.Sizeg
rad
ing o
rsiev
ing screens ground ma
ter
ialsto ob
tain aspec
ific
Cu
tte
rmi
ll
particle size o
fthefeeds tu ffs. The s ize ofthe screen opening w i
ll
determ inethela rgest par
ticle s izethatw il
l passthroughthe sc reen
whichis usua lly expressedin mesh numbe r. A No .40 mesh sieve
(425 m icrons)is o ften used . V ibra
ting screens operate wi
th h igh
frequency andlow amp litude . Some mach ines bu i
lt wi
th gas or
airt
ight cas ings and suc tion fans are used for screen ing dus ty
materialslikerice b ran. Coa rse ma ter
ials(big par
tic
les)that do not
passth ruthe sieve a rere
tu rnedtothe g rinder andreground unti
la l
l
materials passthroughthe s ieve.

3
. We
igh
ing o
fal
lingred
ien
ts. We
ighingis a veryimpor tants tepin
S
iev
ing feed prepara t
ion. The feed componen ts as prescr
ibed in the feed
formu la
tion mus t be accurate
ly weighed. An accura te balance with
ata r
ing deviceis best becauseit can be adjus
tedto zero and a l
lows
weighing w
iththe con ta
ine r
. Thetype o f we
ighing mach ines depends
onthe amoun t and kind offeedingredientsto be weighed.

4
. M
ixing o
f we
igheding
red
ien
ts. Mix
ingisthe process o f sca tter
ing
dissimila
r pa
rtsinto a blend. Feeding redients are sequen t
ia l
ly added
and m ixedfo
ra tleas
t5 m into produce a we llblended m ixture. M ix
ing
ofthefeedcomponen tstofo rmthe mea la re donein ba tches . There a re
two m ix
ing opera t
ionsinvo lved in the
process:p remixing ofm icronu tr
ien ts, and
blend ing o fthe bu lk d iet componen ts.
Feed componen ts are sequen t
ia l
ly added
alittle at at ime or by ba tch. Theth ree
basic mechan ismsin m ix ing are:t rans fer
of ad jacen t part
iclesf rom oneloca t
ion
in the massto ano the r,d istribu t
ion o f
par t
ic les over a fresh ly deve loped
su r
face , and slipping o f particlesinthe
We
igh
ingo
ffeeding
redien
ts
m ass
.
 CHAPTER 5 Process
ing o
f Feeds
tuf
fs and Aqua
feeds 1
31
There are three genera
l types o
f m
ixers usedin thefeed m
ill
ing
indus
try
:

□ Ho
rizon
talr
ibbontype m
ixe
rs (ba
tch
and con tinuous). No rmally, mixingis
doneth rough a ho r
izonta
lr ibbontype
mixer when m ixing ingred ients o f
dif
ferent part
icle s
ize with somel iqu
id.
The des ign and con f
iguration o fthe
mixingribboninre lat
iontothe k ind of
materia
l being m ixedis essential
.
Ho
rizon
talm
ixe
r
□ Ve
rtica
l boo
tload
ing m
ixe
r. M
ixe
rs
shou ld be constructed for easy
load ing
, mea ldischarge as well as
cleaning o
fthe m ixer un
itaf
ter eve
ry
mixing operat
ion
. Residual mea
l can
cause qua li
ty problemsin the nex t
batch orload.

□ Osc
illat
ing sc
rew m
ixe
r. M
ixing
,
particularly when work ing w ith dry Ve
rt
ica
lmixe
r
powders in the absence o f dos ing
liqu
ids, cancausef inefeed pa r
tic
lesto
escapef romthe m ixer unless proper
covers are prov
idedforinthe m ixing
vessel
. The power and speed o fthe
mixerdrivesshouldbedes ignedto mee t
the needs ofthe m ixe
r whenloaded a t
a max imum ma terial dens
ity.

Horizon tal m ixers ® a re p refe

rredinthe
indus try because hand ling and m ix
ing o f
bulk and h igh dens ity ma terials are
Osc
il
lat
ingsc
rew m
ixe
r
eff
icien tly achieved. In ve r t
ical mixe rs ©
high dens i
tying redientstendto se t
tle at
the bo t tomleav ingl ight we ight(powde ry
)
ing redientsto escape du r
ing m ixing.A fte
r
thefeeding redients are m ixed,the mea l
requ ires cond it
ion ing before pellet
izing.
132 CHAPTER5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

5
. Cond
ition
ing isthe process o
f add
ingl
iqu
idstothe mea
l, knead
ing
ofthe mealandsteam ingtoa l
lowsomeo fthes tarch ando the rb inde rs
to ge
lat
inize
. The mea lis usua lly steamed a ta p ressu re o f abou t 10
to 25 pounds/ inch2 (ps i
) w i
th enough t ime
(approx ima te ly 1 -2 m in )a llow ing a s tarchy ma te rial
toform a ge l. Con tro lo fs team tempera ture and
pressureis ve ry essen tialin o rde r no tto a f
fec t some
ofthe hea t-sens itiveing red ien ts.
No rma lly , pe l
le t
ing mach ines a re p rovided w iththe
necessa rysou rce o fs team , me te ring and dos ing pump
forthel iqu id add itives , and a var iab le speed d r
ive .
The speed d rive a llowsfo rthe ad justmen to fthe mea l
cond ition ing t ime to ob tain op timum pe llet qua lity .
In conven t
iona l cond i t
ion ing , abou t 30 -35% o fs ta rch
isge la tin ized . Howeve r,the amoun to fs tarch andtype
ofing red ien ts used may no t be su ff
ic ien tto p roduce a
wa ter s tab le pe llet. There fore , for pe lleted shr imp
feedi tisrecommendedtha ta tleas t 50% o fthe s ta rch
shou ld be ge la tin ized . Th isis done byinc reas ingthe
steam p ressu reto 28 ps i
.
Mos t pe lle t m ills a re equ ipped w i th one to three
directs team cond itione r ba rrels whe re s team can be
injec tedin tothe mash o rin tothe cond itione rjacke ts
to par tia lly ge lat inize s tarch or d ie tb inders (see
Chap ter 4 ). Increas ingthe re ten t
ion t ime or us ing
mu lt
ip le cond it
ioners and con trolo fs team pressure
caninc rease s tarch ge latinizat ion. Thefeed m ixture
intheth ird cond i
tione rreaches atempe ra ture h ighe r
than 90 °C be fore en tering the pe lle td ie. S team ing
imp rovesthe wa ter s tab ili
ty and d iges tibility o f the
feed and k ills mos t ha rm ful bac teria . In sma ll-sca le
feed p repa ra t
ion ,s team ingis necessa ryfo rsh rimpfeeds
but may no tbe necessa ryforf ish d ie
ts. S teamed pe llets
are s tab le in wa ter for 4-12 h depend ing on the
ef
fec tiveness o fthe b inde r
. Uns teamed pe llets © b reak
up w ith in 30 m in ® .

6
. Pe
lle
tingand Ex
trus
ion
. Pe
lle
tinginvo
lvesfo
rcing
the m ixture through holes of a me tald ie pla te to
transfo rm a softfeedinto a hard pel
let. The p rocess
is accomp l
ished by compress ion, ex trusion , and
adhes ion . The primary objective of pe l
leting is to
compac t thefeedf romthe m ixed powderform o r meal. Fo r sh r
imp
feed, pellet
ing preven ts thefeedf romimmed iate dispersion in the
pond wa ter which resu ltsinfeedloss and wa ter po
llution. Theloss
of no
t mo rethan 10% o fthe pe l
let we
ightafter a 10 m inimme rsion
in wateris cons idered accep table.
 CHAPTER5 P
rocess
ing o
f Feeds
tuf
fs and Aqua
feeds 133
The more commontypes of pe
lle
ting machines a re:
□ Ring die pe
lle
t mi
ll(1
) □ D impled ro
lls pel
letizer
□ Flatdie pel
let m
ill(2
) □ Sc rewtype pelle
t ex t
ruderp
ress
□ Scheulertype pel
let

Thering andf la
td ie pel
le tm i
lls a
regene ral
lyemp loyedin aquacul ture
feed mil
ls. Bo th are capable ofp roducingfairly small pe
lle
ts sui
tab lefo r
aquaticfeeds.
Inthe ex t
rusion p rocessthe mea lis heated, kneaded, and m ixed. A
pressure is increased to 20 to 70 bars depend ing on the produc t
formulation
. The sudden p ressure drop a
tthe ou tle
to fthe dieresu ltsin
mater
ial expansion. Theleng th ofthe die channe lp lusthe numbe ro f

In
terna
lpa
rtso
far
ingd
iepe
lle
ting mach
ine

Pel
lets
Knives
Pe
llet
ing mach
ine drum w
ith bu
ilt
-in ro
llers Rol
lers

Pel
leted feeds
Unpel
leted
forced through
mater
ial
the die plate

Pe
lle
ting mach
ine

d
ies greatlyinfluencethe outcome o
fthefinalproduct
. Unl
ike a pe
lle
t
m
il
l, the extruder can shapethe ma
teria
lin a
lmost anyfo
rm.

Thetwotypes o
fex
trus
ion a
re:

Dry extrus
ion -the mea lis
extruded w i
th wa ter only
.
Usua l
ly the pe llets are of
burned tas te andlow bu lk
density. Abou t 30 -35% of
the vitamins a re damaged .
Thef inal produc t need not
be dr ied bu t coo ling is
necessary.

Wet ext
rusion -the mea
lis Expans
ionpe
lle
tmi
ll
extruded w ith steam and
cooked atlowertempera
ture
than in dry extrus ion.
Vi
tamin losses are on ly
abou t 10%.

Ex
trus
ion mach
ine In
terna
lpa
rtso
fanex
trude
r
134 CHAPTER 5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

The productis pas

te-
like be
forei
t en
tersthe d
ie p
late
. Some o
fthe
possib
le shapes are
:

Ex
trus
ion vs Pe
lle
ting
The dec
ision whethe
rto use anex
trus
ion o
r a pe
lle
ting mach
ine depends
on manyfac tors
.
Ex
trus
ion Pe
lle
ting
1
. H
igh
lyf
lex
ibleinfeedfo
rmu
lat
ion
. 1
. Lessf
lex
ibletofeedfo
rmu
lat
ion
.
2
. Can p
rocess mea
lwi
th 55% mo
istu
re. 2
. Mo
istu
re con
ten
t upto 17% max
imum
.
3
. Cook
ing o
fthe mea
lis ach
ieved a
t90% 3
. Cook
ing o
fthe mea
lisach
ieved a
t50% on
ly
o
rhighe
r. 4
. Can hand
lefa
tleve
ls upto 8%l
ipid on
lyin
4
. Can hand
lefa
tleve
ls upto 22%l
ipidin thefo
rmu
lat
ion
.
thefo
rmu
lat
ion
. 5
. Requ
iresf
ine
ly g
round ing
red
ien
ts.
5
. Can b
ind coa
rse
ly g
rounding
red
ien
ts. 6
. Bac
ter
ia may s
ti
ll be p
resen
tinthef
ina
l
6
. Bac
ter
ia a
re e
lim
ina
tedinthe p
rocess
. p
roduc
t.
7
. H
igh p
roduc
t du
rab
il
ity and wa
ters
tab
il
ity
. 7
. Low p
roduc
t du
rab
il
ity
.
8
. H
ighe
r accep
tab
il
ity o
ffin
ished p
roduc
t. 8
. Produc
tarerewo
rkedth
rough sys
temto
9
. P
roduces ve
rsa
ti
le(
floa
ting
,sink
ing and reducef
ines
.
s
low s
ink
ing)feeds
. 9
. Produces on
ly s
ink
ingfeeds
.
10
. H
ighe
rcap
ita
l and p
roduc
tion cos
ts. 10
. Lowe
r cap
ita
l and p
roduc
tion cos
t.

Extrus ion is more e ff

icient in
producing var
iedfo rms o f aquafeeds(1)
,
human(b reak
fast cereals, snackfoods ,
textured vegetab
le pro te
ins (TVP ) and
starch-basedindus t r
ial produc ts) and
premium pe tfoods(2 )
.

Bene
fits o
f Pe
lle
ted Feeds
□ Feeds are mo restable andcan be s to
redfora t mostthree months wi
th
minimal nutrient degradat
ion.
□ Pel
lets tab
il
ityis ma intained duringfeeding
.
□ Feed consump t
ionisinc reasedthe rebyreducingtheleach ing o
f water-
soluble nutr
ien ts
.
□ Salmonella ando therharmfulandd iseasecausingorgan
ismsa redest
royed.
□ Finished produc tis hand led eas
ily.
 CHAPTER 5 Process
ing o
f Feeds
tuf
fs and Aqua
feeds 135
Phys
ica
l cha
rac
ter
ist
ics o
fagood pe
lle
t

Theidea
l pel
letis smooth, has uni
formlength and s
ize(1
), wa
terstab
le(2
)
,
w
ithou
tcolor(3
) and size var
iat
ion(4 )wi
thin a batch
, does notclump(5)
,
wi
thoutfractures(6
), and not wet and mo
ldy(7)
.

7
. Pe
lle
t Coo
lingand D
rying
.The
pelle
tsleav ingthe d ie a
re stil
l
hotand mus tbecoo ledtoallow
the binde rsto set and harden.
The coo l
ing process is
accomp lished by b lowing cool
air in a counter current
direct
ion allowingthe binderto
set as the pe l
lets are cooled.
The vertical or hor
izontaltype
pellet coolers are com mon ly
used.

Hor izonta l coo ler uses a Ho

rizon
talcoo
ler
number o f hor izon tal mov ing
perfora ted t rays w ith the ho t
pel
le ts mov ing coun tercur ren t
to the coo ling a ir. One
disadvantage o f a hor izonta l
pel
le t cooleristha ti t occup ies
a larger f loor area than a
vertical coo ler o fsim ilar s ize .
Ofthe two pe l
let coo lers, the
vert
ica ltypeis common ly used
for sma l
l capac ityfeed p lan ts
together w ith the pe llet
crumb ie r
,ins tal
led a tthelowe r
exit end o fthe pe l
le t cooler.A
cyclone co l
lector is a lso
necessaryto recoverf inefeed
particles tha t may be car ried
136 CHAPTER5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Componen
tsofavert
ica
lpe
lle
t by the coo ling a ir asi t passes
coo
ler through the pe llets. Coo lers
extract mo isture and hea t and
separate thef ine par ticlesf rom
the pel lets. F ines are removed
fromthe coo ling a i
r by a cyc lone
collector and re turned to the
meal cond itione rfo r re
-pe lle
ting.
Itisimpor tan ttha tthe recyc led
mealfines a re removedf rom the
system when thereis a change
foreve ry new ba tcho ffeed . Pellet
tempe ra ture shou ld no t be mo re
than 4-5°C h igher than the
Ve
rt
ica
lcoo
ler amb ien t or room tempera ture .
The coo ling process shou ld have
ah igh a
irf low, andlong re tention timeinthe coo ler whe re pellets
are no
trubb ing against each o
the r
. Anine fficien t cooling sys tem w ill
produce a pelle
tw ith high moisture content,h igh pe rcentage o ffines,
and poor wa ter s
tabil
ity.

8
. Pe
lle
tCrumb
ier
. The s
ize o
fthe pe
lle
tstha
t can be
ef
ficien tly produced by a pe l
le t
ing mach ineisl im i
ted
to a m in imum d iame tero fabou t2 .5 mm . Th isfeed
si
zeisra the rla rgefo rf ryo f upto 3gs izeto hand le
.
The mos te f
fective wayto p roduce sma l
ls i ze offeed
fortheyoungf ryistoreducethe pe lletbyc rumb ling.
Crumb ledfeeds a rerhomb ic andi rregula rin shape
because they are produced by break ing the
cyl
indr ica l pellets .
The c rumb ieris composed o ftwo c lose ly spaced
rotating s tee
l ro llerstha tc rush the pe llets asthey
pass through the ro llers. Mos t crumb lers are
designed sotha tthe su r
faces o fthetworo llersrun
B
reak
ingofcyl
inderpel
letsin
toc rumb
led ats ligh tly dif
feren t speeds . Th is gives a shear ing
feedsusingro
tat
ingrolle
rs act
ion onthe pe llets asthey a re c rushed be tween
thero lle rsto y ield a 0 .5to 1 .0 mm c rumb le size
.

9
. Pe
lle
t and C
rumb
led Feed C
leane
r. F
ines p
roduced a
fte
r pe
lle
ting
mus t be removed by s i
fting or by using af ine separatorin o rderto
producef ine-freefeeds . To separa tefinesf rom the pe l
le ts, mu lti­
decked vibra t
ing sc reens o rrotarytype sifters are used.
The advan tages o fthe ro tary si
fter areits compac ts ize,h igher
ou tput
, and e f
fective separat
ion offines. Howeve r,the d isadvan tage
istha tthe sc reens haveto be changedfo r each desired s
ize o f pellets
orc rumb les. Inthe mu lt
i-decked vibra
ting screen a numbe ro f pellet
sizes can be ob tainedin one ope rat
ion duetothe d i
fferents izes ofthe
screensins ta
lledinthe mach ine.

S
if
ter
/Finessepa
rato
r
 CHAPTER 5 Process
ing o
f Feeds
tuf
fs and Aqua
feeds 137
10
.Produc
t Packag
ing and S
torage
. Packag
ing
ands torageo fape l
le tedfeed p layve ryimpo r
tan t
rolesin ma in taining goodfeed qua l
ity. Feeds
usua lly have al im ited she lfli
fe (3-4 mon ths
withou t an tioxidan ts). Packag ing ma terialssuch
as po rous sacks andloose lysewed open ings can
shor ten she lf
-li
fe.
Freshness o fthe feed can be ma intained
when packed in bags made o f lam inated
polypropy lene (PLP )o r paper bags w ithl ine rs.
The PLPisl ight, very s trong ,inexpens ive, and
provides good wa ter proo fing. Paper bags a re
prone to tear ing dur ing hand l
ing and are no t
recommended when transpor tingfeeds tofar
places. Un l
ikefeedsforl ivestock, aqua feeds a re packedin sma ller
sizes of5 , 10 , and 25 k ilogramsto avo idtea ring o fthe bags .
Pack ing o f pel
le tedfeeds requ ires ca re and a t
ten t
ion to preven t
break ing o fthe pe lle ts and produc tion o ffines . The very essen tial
componen ts o ffeeds such as v itam ins, m inera ls, andl ipids may be
adve rsely a ffected by pro longed exposu retol igh t, excessive moistu re
,
poo rven t
ila tion orh ightempe rature. Inthefa rms , pelle
ts are s
to red
int igh tly cove red p las t
ic bucke ts orjars a f
te rd ispens ingf romfeed
bags .
The s teps to befo l
lowedinlarge -sca le, sma ll-scale
, andlarva l
feed p repa ra t
ion and a gu idetothetype and k ind offeed at various
sizes and ages o f aqua t
ic an ima ls are presen ted.

A
.Stepsin La
rge
-Sca
le Feed P
repa
rat
ion

Sophist
icated andla
rge equ
ipmentare usedin afeed mil
l.
Pel
leted feeds can be made much better and inlarger
amoun ts(as much as 1
-20tons o
ffeed a day)
. The steps
involved are
:

G
rind
ing O
A
lldrying red
ients a
reground
separately with a ham mer
mi
llorc rusher
138 CHAPTER5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

2 S
ift
ing ands
iev
ing

Coa rse materials are separated by s ieving

andre turnedtothe g rinde r unti
la ll pass
through the s ieve. By means o f a screw
conveyor ,s ifted ma terials pass to the
pulve r
izerandg roundto un i
forms ize. The
sieveding redien ts a
re blendedfo r 5to 10
minto ob tain a homogenous p roduct. The
ground ingred ient shou ld be p laced in
tigh t
ly covered con tainers, and s toredin
an adequa te
ly ven ti
lated room .

3 We
igh
ing

Al
ling red
ients a
reindividua
lly we
ighed as
accurately as possible by using a top
load ing we
ighing balance with a tar
ing
device.

4 M
ixing

Theformu lat
ionisthen m ixed by batchin
a large m ixer. The ma jor ingredients
(prote
insou rceslikefish meals,leaf meals,
etc
.)a re pouredin to the m ixer one ata
time and m ixedfor 5 m in. The othe rd ry
ingredien
ts a re addedin to the mixer one
at atime and m ixedfor anothe r5 m in.

5 Ba
tch m
ixing

In a separate conta iner, the micro­

nutrientslike vi
tam ins and minera
ls a re
comb ined w ith a sma ll amoun to f dry
mixed ingredients and par tofthel ipid
sourcesinthefo rmulais addedtothe d ry
ingredients
,tho roughly mixed
, and poured
intothe m ixer. Th
is ensures good mixing
of smallfeed componen ts withthe ma jor
ones.

Note
:
Never p
lace micronur
ien
ts
in an empty mixer

When al
l dryingredien
ts inc lud
ing the
m
icronu
trien
tshavebeen mixed
,addtherest
oftheo
iland mixfo
r 15 min.
 CHAPTER 5 Process
ing o
f Feeds
tuf
fs and Aqua
feeds 139

Cond
ition
ing and ge
lat
iniza
tion 6
The s tarch or b inder is ge latinized by
introducing s
teamin tothe pellet condit
ione r
.
The amoun tof water and hea t required to
cookthe d r
ied mixedingredientis equ ivalent
to the amount o fs team in jected. Th is is
determined by wa
ter stab
ili
ty and pa latabi
lity
tests
.

Pe
lle
ting and ex
trus
ion
7
The cond i
tionedfeed m ixis pel
letizedtothe
desired s
ize. Asthe m ixture passesth rough
the die hole(2-22 mmin d iameter
), arotat
ing
blade cutsi ttothe des
i redlength(1 .5- 2.5
times the d iamete
r).

D
rying and coo
ling
8
The pe
lle
ts drop au
toma t
ical
lyinto a coo
ling
chamber wi
th aira t ambienttemperature.

Packag
ing and s
torage 9
Specia lfeed fin ishing such as coa t
ing and
glaz
ing may be adop tedtoimp rovethe qua l
ity
and s trength o fthe pellets
.
Thef inishedfeed a re packedo rbagged and
labe led proper ly inc luding the da te o f
manu facture. Feeds a re norma l
ly good on ly
forthree mon ths. Bags shou ld be p i
led(no t
morethan 5 bags h igh) on a p lat
form 12 -15
cmf rom thef loorin a we ll-ventilated room .
Thefi rst bagstha tcomein shou ld bethef irst
onesto go ou t
.
140 CHAPTER 5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

B
.Stepsin Sma
ll-Sca
le Feed P
repa
rat
ion

Mix
ing ofloca
lly ava
ilab
lefeeding
redients a
t homeis a p
rac
tica
l way o
f
produc
ingfeedsfor sh r
imps andfishes
.

2 Sc
reen
ing

Thegrounding redients a
resieved o
rsif
ted us
ing
a No
.40 sieve or a nylon ne
tw ith mesh s
ize o
f
425 mm.

3 We
igh
ing

Al
l ingredients are weighed or measured
accura
telyin a bow
lor bas
in.

4 M
ixing

A
ll dryingredients are m ixed thoroughly. The
v
i tamins and m
ine ra
ls are mixed sepa
rately wi
th
the lipid source (oil
) and added to the dry
mixture
. The m ix
tureis thorough ly mixed for
another5 m in
.

5 Ge
lat
iniza
tion o
rcook
ing

To cookco rn s
tarch,b readf lour, and binders
,1
par
ts tarchis addedin to 4 parts of water(50 g
in 200 ml waterfo r 1kg o ffeed)in a saucepan.
The sta
rchis cookedin to ajel
ly-like consis
tency
in a double boi
ler
.
 CHAPTER 5 P
rocess
ing o
f Feeds
tuf
fs and Aqua
feeds 1
41
Knead
ing 6
The gelat
inized binderis removedf rom
heat and a l
lowed to coo l. The cooled
gela
tinized s tarch or binderis added
into the dry m ixture and m ixed for 5
minutes unti
la s t
iff doughisfo rmed.

Pe
lle
ting 7
The dough is squeezed and passed
througha mea tg
rinder
.Theapp ropria
te
dyeis usedfo
rthes i
zeo ff
isho rshrimp
to befed(Table 5
.2)forva r
ious si
zes.

S
team
ing 8
The pe l
leted feedis spread even ly on
wire ne ts that fi
t ins ide the me ta l
holders ofthe steame r
. Wa teris boi
led
(5-8cm deep )inab igpotandthepe l
lets
are s
teamedfo r5 m in. Overcook
ing o r
steam ing can causeloss o f nutr
ien ts
and makes pe l
letstickto each other.

D
rying and coo
ling 9
The s teamed pe l
lets are a
ird r
iedfo ra
few minutes w i
th an e lectric fan,
transfe rredtoanovenandle ftove rn
ight
orfor 8 -12 h at 50-60
°C. Pel
letsshou ld
not be d ried unde rthe sunto p revent
dest
ruc t
ion o fsomev itamins and other
light-sens it
ivefeed componen ts
.

Cu
tting
, packag
ing
, and s
torage 0
1
Afterthe pelle
ts are removedfromthe
oven,theya recooledfor30 -60 m
in.The
dried ex
truded or pel
letedfeeds a
re cut
tothe desiredlengths,placedincovered
plasticjars, and storedin a cool dry
place.
142 CHAPTER 5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

C
.Stepsin La
rva
l Feed P
repa
rat
ion

1 Weighing
A
llgroundingredients a
re measured o
r
we
ighed accura
tely.

2 Mixing
Al
ld ryingredien
ts are mixedthoroughly.
Iflarge batches areto be prepared
, the
dryingredients can be mixedin ala rge
cake or cement mixer
.

3 Blendingofoilandlecithin
The o il and lec ithin are b lended
separately and added to the dry
ingredients gradually with continuous
mix
ing. When al
lthe oil andlec
ith
in have
been added , mixing is continued for
another5 m in.

4 Cooking ofcarrageenan
The carrageenan is cooked to gelat
in­
like cons
istencyin a wate
r ba
th at 80
°to
100°Cthen s l
ight
ly cooled
.

5 Microbounding
The cooked ca
rrageenanis addedtothe
dry mixture and blended well unt
ila
comp lete
ly homogen ized mixture is
obta
ined.
 CHAPTER5 P
rocess
ing o
f Feeds
tuf
fs and Aqua
feeds 143

F
lak
ing 6
The so
ft massis passedthrough a d
rum
dryerorflak
ing mach ine andthe br
itt
le
flakes a
re col
lec
ted .

G
rad
ing/S
iev
ing 7
The flakes are ground gen t
ly using a
mortar and pest
le or mea tgrinde
r and
sievedto uniform part
iculate s
ize us
ing
varied mesh size s
ieves.

D
ifferen
t par
tic
le sizesforlarva
lfeeds
a
re 25, 50
-60, 125
, and 250 pm.

Tab
le5
.2 Gu
idetotypesandk
indso
ffeedfo
raqua
tican
ima
lsa
tva
riouss
izesandages

F
ishsize Typeof Feeds
ize
, Leng
th
approx w
t(g) feed dia(mm) (mm)

0
.35 g andless S
tar
ter 1
.0 –

2-5 Growe
r 2
.0 -
5 –12 Growe
r 3
.0 2-3
12 –20 F
inisher 5
.0 3-5
20- 30 F
inishe
r 7
.0 5-7

Sh
rimpage Shr
imp Feed Feed Feed Feedleng
th
(days
) s
ize(g
) type form s
ize(mm ) (mm)

PL15– PL3
5 <1
.0 S
tar
ter F
ine c
rumb
le 0.6 –1
.0 –

PL36– PL5
5 2-3 S
tar
ter Coa
rse c
rumb
le 1.0 –2
.0 -

PL56– PL7
5 4-7 Growe
r Pe
lle
t 2.0 –2
.2 1.2 –3
.0
PL76– PL9
5 8 –14 F
inisher Pe
lle
t 2.0 –2
.5 2.2 –5
.0

PL96– PL1
15 14- 22 F
inisher Pe
lle
t 2
.2-2.5 2.2 –5
.0
PL116– PL1
35 23- >30 F
inisher Pe
lle
t 2
.5- 3
.0 4.0 –8
.0
144 CHAPTER5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Qua
li
tyCon
tro
l

Feed quality cont

rolis one ofthe ma jor keysto a success ful culture by
selec
ting
, eva lua
ting, and mon i
toringfeeding redients during processing,
manufactu re
, and un t
ilstorage of aquafeeds
.S torage condit
ions shou ld
be optima l sotha tthe nutrientsinfeedingred ien ts andfeeds do no t
deter
iorate and econom iclosses o ffeed m i
llers andf ish culturists are
Sto
ckingof10bag sperpal
let
avoided. The fo llowing pointers shou ld be fo llowed to avo id rapid
is prac
ticed whenfeeds are deter
ioration offeeds.
sto
red fo r sho r
t periods
(usual
ly3to5da ys
).
□ mo isture con ten t shou ld no t be more than 10% and re la tive
hum id ity notg rea te rthan 65%to p reven tfunga lgrowth andinsec t
in festa tiontha tg rows bes ta t 26 °-37 °C.
□ tempera ture shou ld no t be too h igh to preven t destruc t
ion and
reduc tion o f nu trien t.
□ exposu reto oxygen shou ld bel im i
tedto p revent oxidativeranc id i
ty
par ticu larlyinfeeds o fh ighl ipid con ten t. The perox idesfo rmed
froml ip id perox ida t
ion may b ind w ith pro teins or vitam ins and
reduce the ir ava ilab ility.
Feed shou ld be:
Indiscr
iminates
torageoffeed
cantriggerrap
iddeter
iora
t
ion
□ freef rominsec ts and roden ts and spo i
ledfeeds shou ld not be
of nut
rien
ts. used o rrecycled .
□ s to redin d ry, coo l, and we ll ven t
ilated a rea and no t unde rd irec t
sun l
igh t
. Spo ilage w il
l occurimmed iatelyi ffeeds become we t.
□ p laced on wooden pa lle ts and no td irec t
ly onf loors with no t mo re
than 5 bags pe r pa lleti fintendedfo rlong s torage. Feeds shou ld
be s to red awayf rom wa l
lsto avo id mo isture accumu lation dueto
hea tt rans ferf romthe wa lltothefeed dur ing suddentempe rature
changes .
□ used w ithinth ree mon thsf romthet ime o fp rocess
ing and p roduction .
Thef i
rs t onetha tiss toredshou ld bethef irstto be used. Thef i
rst -
in
first-ou t policyo fs torageshou ld bes trict
lyobse rved.

Feed m
i
llsan
ita
tionand ma
intenance

Ef
fective managemen t and prope r ma in tenance o fthe s torehouses
andfeed m i
ll he
lp ensu re good qua li
tyfeeds . Sanitation p revents
infestat
iono finsects,fungi, andbac teria. Infested ma te
r ialsshould
be removed andthein fested areas shou ld be d is
in fected . On ly
af
terd isin
fection can a new ba tch o ffeeds tuffandfeeds be s tored.
To prevent and con tro
linsec tin festation andfunga l and bac ter
ial
contam inat
ion par t
icularlyAspe rgi
llusflavus (1 ) and Sa lmone lla in
feed m i
lls
, several measures shou ld be s tr
ictlyfollowed .

□ feeds andfeeds tu f
fs shou ld be dr
iedto a mo is ture conten tof
10% o rless
.
□ feedstuff
'sshould bep rocessedtoensu recomp lete destruction
of bacter
ia, fung i
, and v iruses,o r reduc tion to accep table
levels
.
Aspe
rgi
llus sp
.oncon
tam
ina
tedfeedpa
rt
icles
.
□ whenever poss ible, variet
ies offeeds tu f
fs tha ta re resistant
tofung iandtox insshou ldbe used. Feeds tuf
fssuch as ma ize,
copra, and peanu ts are eas
ily contam inated w ith aflatoxin.
 CHAPTER 5 P
rocess
ing o
f Feeds
tuf
fs and Aqua
feeds 145
□ envi
ronmen t-f
riendly chem ica ls such as vinegar, ammon ia
, andlysol
should be usedto d is
infec tthe m i
ll
.
□ moving pa rts such as gea rs, roller chains
,e tc. should belub r
icated
atleast once eve ry 4 hfo r every con t
inuously m ixed batch.
□ atthe end o f each m i
lling pe riod, thefeede r
, conditione r
, and the
main dr
ive o fthe pelle
ting mach ine shou ld be always cleaned with a
bag ofrice bran(o rdinary grade )o r sawdus twith cooking oil(100 ml
ofoi
lfo r every 2 kg ofrice b ran).

Summa
ry

In p rocess ingfeeds ,theing red ien tsto be used shou ld bef reef rom mo lds
and/or o ther harm ful subs tances . Thus , the prepara t
ion o ffeeds tuffs
wil
l depend onthe presence o r absence o f harm ful subs tancesin the
ing redien tsto be usedin mak ingfeeds . Qua lity pe l
lets can be p roduced
when proper manu facturing prac t
ices a refo llowed . Among the bas ic
poin ts to cons iderto produce good and qua lity pe l
lets w ith m inima l
fines a rethe use o f:a) satisfac tory m ixing p rac tices w i
thther igh tk ind
of m ixerforthe type o f mealto be produced ,b ) the cor rec tk ind and
qua lity o f dosingl iqu id or b inders , for the s ize o f pellets and feed
ing redien ts used ,c) use ofco r rectd ies ize andth ickness ,d ) useo fp roper
pelleting p ressure andtempe ra ture,e) co rrecttype o f pel
let coo lingra te
and coo ling a irf low rates,f) co rrectc rumb lingro l
l settinginre la t
ionto
the des i
red crumb ledfeed s ize, and g ) pos t-pe lle t
ing ,c rumb l
ing , and
finish ing trea tmen ts to removef ines andf rac turesf rom thef inished
produc t. Atechn icallytrainedandsk il
ledfeed m i l
lope ratorshou ldove rsee
the manu facturing p rocess.
The s tepsto befo llowedinlarge sca le
, sma ll-scale, andlarva lfeed
prepara tion w ill guidethe reade r astothetype and k ind o f aqua feeds
forf ish and c rus taceans a t var ious stages and ages .

Gu
ideques
tions

1
. Wha
taretheimpo
rtan
tth
ingstocons
ide
rinthe p
repa
rat
iono
faqua
feeds?
2
. Why do we needto p
rocessfeed andfeeding
red
ien
ts?
3
. Whyis g
rind
ing o
fing
red
ien
ts a necessa
ry s
tepinfeed p
repa
rat
ion?
4
. Exp
laintheimpo
rtance o
f know
ingthe mo
isture con
ten
toffeeds
tuf
f
andfeed espec
ial
ly p
rio
rto s
torage?
5
. Wha
t are some typ
ica
l examp
les o
f an
tinu
tri
tiona
lfac
torsinfeed
ing
red
ien
ts andthe
ire
ffec
ts on aqua
tic an
ima
ls? How a
rethesefac
tors
e
lim
ina
ted o
r m
inim
ized?
6
. Wha
tisthe pu
rpose o
f pe
lle
ting and how does one makef
loa
ting and
s
ink
ingfeeds?
7
. Exp
lain br
ief
lythe bas
ic s
tepsin sma
ll-sca
lefeed prepara
tion and
compa
retheseto s
tepsin comme
rcia
lfeed p
repa
rat
ion(m
il
ling
).
8
. Wha
tarethe d
ele
ter
iou
sef
fec
tso
fpro
longedandimp
rope
rsto
rag
eoffe
eds?
146 CHAPTER 5 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Sugges
ted Read
ings

Ak
iyama DM , Dominy WG . 1989 . Penaeid shr
imp nutr
ition for the
comme rc
ialfeedindus try. Texas Sh
rimp Farm
ing Manual Volume
1. Co
rpus Chris
ti, Texas. 500 p.

Aquacu
lture Developmen
t and Coord
ina t
ion Program. 1980
.Fishfeed
techno
logy. ADCP/REP/80/11. FAO, Rome. 395 p
.

Aquacu
lture Developmen
t and Coordina
tion Prog
ram . 1983
.F ishfeed
techno
logy. ADCP/REP/83/18. UNDP/FAO , Rome. 97 p
.

Go
lez NV, Guanzon N, Mil
lamena OM . 2000
. Environmenta
limpac ts o
f
sem i
-intensive shrimp farming. A paper presen
ted a t the 4th
Sediment Quali
ty Assessment Symposium, November 22
-28, 2000,
Otsu
, Kyoto, Japan.

Go
lez NV
, M il
lamena OM , Hanssen OK . 1998. Evaluation o
f processed
soybean as pro te
in source for shrimp feed based on grow th
,
survival
, andfeed conversion of Penaeus monodon juven ile. A
paper presented atthe S
ixth Asian Fisheries Forum, November
11-14, 1998. Ch
iang Mai
, Thailand.

Kanazawa A. 1982. Penae id nutrit

ion. Proceed ings ofthe Second
Internat
ional Conferencein Aquacu lture Nutri
tion: Biochemica
l
and Physio
log
ical Approachesto Shel
lf
ish Nutri
tion, 27
-29 October
1981. Rebobo
th Beach , De
laware, USA. 87-105 p.

L
ienerIE. 1962
. Toxicfac
torsin ed
iblelegumes andthe
ire
lim
ina
tion
.
J
. Nutr
. 11: 281
-298
.

M
illamena OM, Golez NV. 2001
. Evalua
tion of processed meat so
lubles
as pro
tein sourcefor sh
rimp.Isr
.J . Aquacu l
t.-Bamidgeh. 52(3)
:
91-97

Mura
i TA, Suma lankay A, Pascual FP. 1983
. Supp
lemen tof var
ious
at
tractants to a pract
icald ietforjuveni
le Penaeus monodon
Fabr
icus.F ish
. Res.J. Phi
lipp.8:2-6
.

New MB. 1987

. Feeds andfeeding o
ffish and sh
rimp
. A manua l onthe
prepara
tion and presen
tat
ion of compoundfeedsfor shr
imp and
fishin aquacul
ture. ADCP/REP/87/26 , UNDP/FAO. 275 p
.

O
loghobo AD, Fe
tuga BL. 1983. Trypsininh
ib i
tor ac
tiv
ityin somelima
bean(Phaseo
luslunatus)va r
iet
iesasaffectedbydif
feren
tprocess
ing
methods
. Nut
r. Rep
.In t
l. 27:41-50.

Pascua
l FP, Rivera RV
. 1989
. Feed
ing prawnsin grow-out cu
lture
.
Aquacul
ture Extens
ion Pamph
let2
. SEAFDEC ,Tigbauan,Ilo
ilo
.
21 p
.
 CHAPTER 5 P
rocess
ing o
f Feeds
tuf
fs and Aqua
feeds 147
Peña
flo
rida VDP, Golez NV
. 1996. Useofseaweed mealsf
rom Kappaphycus
alvarez
ii and Graci
lar
ia heteroc
lada as b
indersin die
tsfo
rjuvenile
shrimp Penaeus monodon . Aquacu l
ture 143:393-401
.

Tacon A
. 1988. The nut
rit
ion andfeed ing o
ffarmedf ish and sh
rimp A
train
ing manual.3. Feeding me
thods. FAO F
ield Document,Project
GCP/RLA/075/TA .F ie
ld Documen t 7/E. FAO, Brasil
ia, Brazi
l,
208 p.
 CHAPTER

6
Eva
lua
tionofFeeds
tuffs
andAquafeeds
MYRNA B
. TERUEL

In
troduc
tion
The use o f qua l
ityfeedsisimpo rtan tinthe success o f any aquacu ltu re
venture . Feed qua l
ityis h ighly dependen t onthe qua lity o fraw ma terial
and the process ingtechn ique . Aformu latedfeed wh ich makes use o f
low qua lity raw ma ter ials w ill no tgive thef ishfarmer any s ign i
fican t
benefit. Feeds tuffs andf in ishedfeeds shou ld, there fore , undergo the
process o feva lua tion and qua litycon trolin o rde rto p roduce h igh qua lity
feed. Sys tema tic eva lua t
ion o ffeeds tu ffs and feed us ing phys ica l
,
chem ical, m icrob iolog ica l, and b iologica l me thodsis necessa ryto assu re
their e ffectiveness when fed to f ish . Th is procedure s tarts f rom
procurement o ffeeds tu f
fs and cont inues to feed process ing unt il
manufac ture and s torage o fthef in ished p roduc t. Thef in ishedfeed mus t
contain a llthe nu trien ts requ ired bythef ishin adequa te amoun ts and
proper propo rtions .
Differen t me thods a re used to eva lua tefeeds tuffs andfeed qua l
ity .
In carry ing ou ta ll these me thods , as tandard samp ling procedureis
necessaryto ob ta in a represen tative samp le . A spear probeisinse rted
diagona l
ly and hor izon tallyf rom one corne ro fthefeed bagtothe o the r.
Samp les a retakenf rom a ll bagsin case o f sma llerlo ts wh i
le on ly 10%
are cons ideredin b iggerlo ts. There a refour me thods o ffeeds tuff and
feed eva luation : phys ical, chem ical, m icrob iological, and b iolog ica l
.
Physica l me thodinvo lves senses o f sme ll, tas te ors igh tto de tectthe
presence o f adu lterants infeeds tuffs and feeds . Chem ical me thod
quant i
fies the amount o fg iven compound present in the feed .
Microb iologica l me thod invo lves the use o f m icroorgan isms in the
evalua tion o f nutr ien ts . B iolog ical me thod invo lves ac tua lfeed ing
experimen t. Th is me thodis mo reted ious and expens ivethanthef irst
three me thods bu tg ives a mo re accura te es tima te offeed u ti
lization .
Th is chap ter d iscusses how to eva lua tefeeds tuffs andfeeds . The
results o ffeed eva lua tion w i
ll be usedto ensu rethe p roduc tion o fh igh
qualityfeedsforf ish ,c rustaceans , and she ll
fish.
150 CHAPTER 6 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Phys
ica
l Eva
lua
tion

A
.Useo
ftheSen
ses

Rancidity and o ff
-odo rsinthefeed can be de tected by sense o f smell.
Of
f-f
lavo rs ofthe maining redien tscontainedinthera tioncan be de tec
ted
bysenseo ftas
te. The p resence o fex t
raneous ma ter
ialsl ikesma l
ls tones,
scrap me tal
,d i
rt,p ieces o f wood, and seedstha ta re addedtoinc rease
the weigh tofthe p roduct, as we ll as p
resence ofinsec ts and mo lds can
be detected by sight.F inally,the we tness,dryness o r ha rdness o
f afeed
orfeedingred ien t can be de tected bytouch . Feeds tu ffs andfeeds o f
acceptab le qual
ity mus t be d ry,freeflowing
, and un iformin appea rance.

B
. Feed M
icroscopy

The m icroscopeiden t
ifiesthe physical compos it
ion of afeedstufforfeed
ingred ient thate i
ther con f
i rms or den ies the presence of unwan ted
mater
ials. Ah igh-powe red compound m icroscope can detect eventhe
finestg round adu l
teran tsin a samp le. Th is me thodis mo re accurate
thanthe use o fthe sensesin check ing adulterat
ioninfeeds andfeed
ingred ients
.
There are two types o f microscope usedin feed m icroscopy: the
compound m icroscope , to identify the interna ls tructure o ffeed
componen ts
; andthe s tereomicroscope,toiden tifythe ex
ternalstructure.

Techniquesin samp le preparationfo rfeed m icroscopy

1
. Sc reen ing. Feeds tuffs or mixedfeed o fd ifferent par
ticle s
izes are
separa ted by hand screen us ing sieves o f no. 10, 20 , and 30
meshes . S ieving separa tes fine starchy dus tf rom thelarger
particlesinthefeedfo r bet
teriden tif
ica t
ion .
2
. F lotation. Feeds tuffo rfeed samp leis soakedin a so lvent(either
carbonte trachloride or chloroform)
,s t
i r
red , and al
lowedto se t
tle
to separa tethe organ ic portion(topf raction)from theinorgan ic
portion( lowe rfract
ion). Eachf ractionisremoved and p lacedin
a pe t
ri-dish and a l
lowedto d rya troomtempe rature.

C
. Measu
remen
tofFeeds
tuf
fsBu
lk Dens
ity

The bu lk densi
ty ofthe samp leis compa red withtha tofa pu refeeds tu f
f.
Ifcon taminants o r adul
te rants are presen
t, the bu lk density w i
lle ithe r
be h ighero rlowe rthanthe va lues ofthe pu refeeds tuffs
. Bu lkyfeeds
arelesse ff
icientin produc ingfishflesh. The useo fbu lkyfeeding redien ts
in afeed m ix
tureins tead o f heavier oneslowers the to ta ld igestible
nitrogeno fthe m ix
tu re
. Bu lkyfeedyieldslow biological
lyava i
lab leene rgy .
The bu lk densityis compu ted as we ighto f samp lesin gram perl i ter
afterthe samp le has been p laced and pou red offin a 1l cylinde r.
 CHAPTER 6 Eva
lua
tion o
f Feeds
tuf
fs and Aqua
feeds 1
51
D
.At
trac
tab
ili
ty

At
tractan ts areimportan t componen tsin anyfeedformu lation s
ince
they determine howfast and how much o fafeed w i
ll betakenin bythe
aquatic anima l. A wel
lformu la
tedfeed will be uselessifthe animal
does notaccep tit
.At
tractabili
tytests a
recarr
ied outw ith as
ingle an
imal
and effec
tivenessis measured usua lly w
ith a stopwatch based on how
fastthe an imalis a
ttractedtothe pe l
let
.

E
. Wa
terS
tab
ili
ty

Feeds shou ld be s tab lein wa terfor a cer ta in period toincrease the ir

avai
labilityto aqua t ic an ima ls. Thefeed hasto ma in taini tsin teg r
ityin
the wa ter sotha tthe en tirefeedis consumed . Th isis espec ial
lyt ruei f
the spec iesto befed a re s lowfeede rs such as c rus taceans . I ffeeds a re
not water s tab le
, they d isintegra te rapidlylos ing much o fthe nu trien t
content and resu l
tinginfeed was tage and deg radation o f wa ter qua lity.
The use o fe ff
icientb inde rs and p rocessingtechn iques , such as s team ing
and extrus ion ,p roduces a mo re stable pelle
t. As imp letes tto de term ine
whether afeedis wa ter s table or no tis by crumb l
ingthe pe l
le t by hand
or check ingfo rrough edges . Ano ther me thodfo rtes ting wa ters tability
is by de term ining we ightloss o fthe pe l
leta fteritis p lacedin wa terfo ra
speci
fic pe riod . Co rrec tin terpretation oftheresu ltsis necessa ryintes ting
otherb inde rstoimp rove wa ter stabili
ty. P rope rp rocess ing cond itionfo r
the formu la ted d ie t and the b inders used are equa llyimpor tan tin
achieving a wa ter s table pe llet. The h igherthe percen tage o f pe l
le t
dis
integ ration a tag ivent ime,the h igherthe we ightloss , andthelowe r
the pelle ts tabil
ity.

As
imp
le me
thod o
f de
term
ining wa
ters
tab
ili
tyisa
sfo
llow
s:

1
. Wire baske ts areto tal
ly oven -dried at 100 °C(1 -3 h), coo
ledin a
desiccator
, and we ighedto cons tant we ight
.
2
. A certain amoun toffeed(abou t 5g )w ith known mo isture content
isthen p lacedinthe w i
re basket.
3
. The w i
re baske ts withfeed a rethen a l
lowedto s tayinthe wa ter
under cond itions sim ilarto those o fthe exper imen taltanks a t
designatedt imes( 2 ,4,6 , and 8 h).
4
. The w i
re baske ts arethen oven -dried, cooledin a des iccator, and
weighedto cons tant we ight
.
5
. Percent dry weigh tlossis ca lculated a fter sub
tractingthe baske t
weight.
6
. Percent water s tabi
lityisthen compu ted as:

Fo
% Wa
terS
tab
ili
ty = x 100
Io

where
:Io = In i
tia
ld ry we
ightoffeed
Fo = f
inald r
y weightoffeed
152 CHAPTER6 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Examp
le: Compute the water s
tab
ili
ty o
f afeed

wtfeed (asis) = 5.26 g

dry matter (DM) = 95%
%DM
wtfeed (dry bas
is) = wtfeed (asis
)x
100

Io
,in
itia
l dry wt o
ffeed .26 x 95
= 5
100
Io = 5
.0 g

A
fter the ind
icated im mers
ion t
ime
:

wtfeed + baske
t = 12
.5 g
wt empty basket = 8.0 g

Fo
,fina
l dry wt o
ffeed = wt baske
t +feed - wt baske
t

Fo = 12.5-8.0
= 4
.5 g

% water stab
ili
ty = Fo x 100 =4
.5
x 100 = 90%
Io 5
.0

Chem
ica
leva
lua
tion

A
.Prox
ima
teana
lys
is

The proximatecompos i
tionis anindextothe nu tri
tiveva lue o ffeeds and
feeds
tu f
fs. This ana
lytica
ltechniqueis designedto d if
fe rentiate between
nutritive and non-nutr itive
components and ana lyze
moisture/dry mat ter content ,
crude p rote
in,c rudefa tore the r
-
extrac t
, crude f iber, and ash .
Nitrogen- free extrac t (NFE ) is
obtained by sub t
rac tingthe sum
ofthese cons t
ituen tsf rom 100 .
Al
though p roxima te ana lys
isg ives
a genera lind ica tion o fthefeed
value,i t does no t dea l w ith
specific nu tr
ien ts. Th is me thod
establishesthe ca tego ryin wh ich
afeeds tuffbelongs andisa use fu
l
descriptive devicein es tablishing
the character istics o f feeds
(Figure 6 .1)
.

F
igu
re6
.1
F
lowd
iag
ramfo
rthe p
rox
ima
te ana
lys
is o
ffeeds
tuf
fs andfeeds
.
 CHAPTER 6 Eva
lua
tion o
f Feeds
tuf
fs and Aqua
feeds 153
1
. Mo
istu
re/d
ry ma
tte
r con
ten
t. I
tisimpor
tan
t to
determ ine how much o fthefeed samp le we igh tis
actua lly made up o f dry mat ter espec ia lly in
formu lating rations us ing ingred ients w ith h igh
moisturecon tent
. Mo is tureis a d iluen to fthe nu t r
ient
in feeds tuffs and feed and has an e ffec t on the ir
stabil
ity and she lfl
ife. Feeds tu
ffs andfeeds shou ld
be dried to containlessthan 10% mo isture pr iorto
storage especia
llyint rop ical coun tries. H igh mo isture
content canleadto g row th o f mo lds dur ing s to rage o f
feedstu f
fs andfeeds .
Moisture content dete rm inationinvo lves d ryingthe
feed samp leto constan t we igh t using a d rying oven o r
a mo isture balance (F igure 6 .2)
. I tisimpor tan tto F
igu
re6
.2
interconve r
tfeed ana lys is da taf rom d ry-ma t
te r bas is A moistu
re ba
lance usedto determine
to as-rece ived basis for a more accura te feed ing mo
isturecontentoffeedstu
ffs and aqua
feeds
.
managemen t usingthefo llowingfo rmu lae:

% nu
trient(asrece
ived) = % nu
trient(dr
y bas
is)x(%d ry ma
tte
r)/100
% nutrient(dry basis
) = % nutrient(as received
) x 100
%d ry ma
tte
r

2
. C
rude P
rote
in. Crude pro
tein de
term
ina
tionis done us
ingthe
Kjeldahl me thod. The samp leis d igestedin concen trated su lfur
ic
acid, resultinginthe comp le te oxida t
ion o fallo rgan ic ma terials
andthe n itrogenisconve rtedtoammon iumsu l
fate
. Excesssod ium
hydrox ideisthen addedtol iberate ammon ia, whichis abso rbed
in bor ic acid and t i
trated w ith standard hydroch loric ac id
. The
procedureforp ro te
in analysis us ing K jeldah l methodisin Append ix
B1 . Ap icture o fthe K jeldah ld istilla t
ion- ti
tration (K jel
tec™ )
ins t
rumen t usedto ana lyzethe c rude p rotein contento ffeeds tuffs
andfeedsis shownin F igure 6 .3. Thete rm c rude p rotein means
tha to ther n i
trogenous ma terials, wh ich a re nott rue pro teins ,
such as urea , am ides, nuc leic ac ids, and am ino sugars may be
present. S incef ish has veryl imited ab i
lityto u t
ilize non -prote
in
nitrogen(NPN ), the measuremen to fthet rue amino ac id pro te
in
shou ld be done onfeeds tuffs.

3
. C
rudefa
tore
the
r ex
trac
t. The crudefa
t con
ten
toffeeds
tuf
fs F
igu
re6
.3
The K
jeldah
ldis
ti
lla
tion
-
and feedsis determ ined by extract
ion of ground samp les w ith
t
itrat
ion(Kje
ltec™) appa
ratus
ether. Ether-solub le mater ia
ls include a variety o f organ ic
usedto analyze c
rude
compounds bu t only afew have nu tri
tiona
ls igni
ficance such as prote
in.
thet ruefats
,fatty acid este
rs, compoundl ipids
, andfa t solub le
vi
tam ins o
r pro
-vitam ins such asthe carotenoids. Th is me thod
does notremove alllipids
, especia
llythe phospholip
ids and o the r
fats boundto proteins
.
154 CHAPTER6 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Ifthe e ther extract contains alarge percen tage

of fats and fat ty acid esters , th is me thod o f
evaluation is va lid because i ta ims to iso late a
fract
ion o ffeedstu f
f which has a h igh caloric value.
However,i fthe ex trac
t con tains alarge percen tage
ofp lant waxes , essent ialo ils, resins or s imi lar
compounds , th
is ana lysis has l it
tle mean ing, as
these compounds have l i
t t
le va lue to f ish. A
procedure for fa t analysis is shown in Append ix
B2. Sox tec™ appara tus usedin crudefa t analys is
offeedstu f
fs andfeedis shownin F igure 6
.4 .

4
. Ash o
r m
ine
ral ma
tte
r. Ash ortheto
tal m
ine
ral
F
igu
re6
.4 con tentis measured by burn ingthefeed samp lein a
The Soxtec™ appa
ratus usedfo
rcrude mufflefurnace a t 550-600 °C. Th is burns a l
lo rganic
fatanalys
is. matter,leaving ares idue o f ash o rino rganic mineral
salts. Excess ive ly high ash va lues may ind icate
con tam ination
, or d ilution o ffeeds tuffs with such
subs tances asl imestone and sa lt
. The con tento f
calcium , phospho rus
, magnes ium , and o therm inerals
infeeds andfeeds tu
ffs are de terminedf romthe ashed
samp le. Figure 6.5showsap ictureo fa mu ff
lefurnace
usedfor ash de termination.

5
.Crudef
ibe
r. Incrudef
ibe
r de
term
ina
tionthee
the
r-
extracted samp leis boi
ledin d i
lu te ac
id,thenin d ilute
base,d ried, and burnedin afu rnace. The d if
fe rence
in we ight be fore and after burningisthe c rudef iber
fraction. Th is me thod simu lates digest
ion occur ring
inthe gas tric stomach andinthe sma llin tes
tines o f
F
igu
re6
.5 thef ish. Crudef iberis made up pr imarily ofp lant
A muff
lefurnace usedto
struc tura l carbohydra tes, such as ce llulose and
analyzethe ashcontentof
hem icellulose , andi ta lso con tains somel ign in,a
feedstuf
fs and aquafeeds
.
highlyind iges t
ible materia
lassoc iated withthef ib rous
portion ofp lan ttissues. F igure 6 .6 shows a p icture
ofthe F ibertec™ , anins trumen t usedfor c rudef iber
determ ina t
ion .

6
.Ni
trogen
-free ex
trac
t(NFE
). NFEis der
ived by
subtractingthesumo ftheo therproximatecomponen ts,
crudep rote
in,crudefat
,ash,c rudefiberonad ry weigh t
basisfrom 100 . Itrepresents mainly s
tarch
, suga rs,
and othe rreadi
ly so
luble carbohydrates
. It may a lso
include hem ice
llu
loses and some o fthe more solub le
lignin
.

F
igu
re6
.6
The Fibe
rtec™ usedfo
rcrudef
ibe
r
dete
rm inat
ion
.
 CHAPTER 6 Eva
lua
tion o
f Feeds
tuf
fs and Aqua
feeds 155
B
. Me
thodso
fPro
tein Eva
lua
tion

. T
1 rue p rotein va lue de te
rm ina tion(Spec t
ropho tome tric methods )
.
a
. B iuret me thod . Th is methodis app licableto ex t
rac tedliquidf ish
protein al
iquo ts, w i
th a pro tein concen tration o f between 0 .1 to
0.5 mg N/m l. The me thodis based onthe reac t
ion o f Cu+2w ith
peptidesin alka line solutionto y ield a pu rple comp lextha t has a
peak abso rpt
ion a t 545 nm .
b
. Lowry me thod . Th is method es t ima testota lp rotein offeeds tuf
fs
andfeeds . I t uses Fo l
in -Cioca l
teu reagen t and has a b lue end
colorw i
th extinc tion at 660 nm . The p r
incip le behindth is me thod
isthereduc t
ion o fthe phosphomo lybd ic acid-phospho tungstic acid
(Folin-Ciocalteu) reagen t bytyros ine andt ryptophan res iduesin
the p ro
tein.

2
. Measu
remen
tofp
rote
in qua
lity
. The use
fulness o
ffeeds assou
rces
of prote
in depends pr imari
ly on the total concen trat
ion of pro
te in
andthecompos i
tion ofamino acids making upthe p rote
in. Imbalance
among am ino ac
idsin afo rmulatedfeedresu ltsininadequa te protein
nutrit
ion. Theimba lance decreases grow th andfeed e ff
iciency o r
mayresultinfish mortal
ity
. There lat
ive usefulness o
fthe prote
in ofa
par
ticularfeedin meetingthe needs o
fthef ishis known asi ts qua
lity.

a
. Am
ino ac
id compos
ition
. Am
ino ac
id ana
lys
isis done us
ing
either an am ino ac id ana lyzer(AAA ) or h igh per formancel iquid
chroma tograph (HPLC ) w ith f luorescence de tec tor. F
igure 6 .7
shows a p icture o f HPLC , anins trumen t usedto ana lyze am ino
acid compos ition o f a prote in. HPLC ut ilizes
o-ph thalaldehyde/N -acetylcys t
ine(OPA/AcCys ) and
Na hypoch lorite reagen t. Am ino ac ids reac t w ith
OPA/AcCystofo rm af luo rescen t subs tance w ith an
excitation wave leng th of 350 nm andf luoresces a t
450 nm . Prote
in samp les a re hyd rolyzed andin jected
into a co lumn w i
th ca tion -exchange res in . Bu f
fers
ofd ifferent pH and ion ic s treng ths are pumped
throughthe co lumnto br ing abou tthe sepa ration of
various am ino ac ids. The amoun to ff luorescen t
compoundis d irectly propo rtiona ltothe am ino ac id
concentrat ion in the e lua te. The am ino ac id is
ident if
ied in the resu lting chroma togram by the
F
igu
re6.7
retentiont ime o fthe peak and quan t
ified bythe a rea
The h
igh per
formancel iqu
id ch
roma tograph
unde rsuch peak . Inthe am ino ac id ana lysis by AAA , (HPLC) usedfor analys
is o
fam ino ac
id
ninhydr in so lu tion is used for co lor deve lopmen t composi
tion o
fa protein
.
instead o f OPA/AcCys .

b
. Chem
ica
lsco
resfo
rpro
tein
. Th
is me
thod cons
ide
rsthere
lat
ive
amoun ts of amino ac
ids presen tinthe pro te
in as determined by
chem ical analys
is. The qua li
ty ofp rote
inis a ffec
ted by arelative
de
ficiencyo fone o
r mo re essen t
ial amino acids. Wholeegg p ro
te in
is cons idered as standard or idea l in terms o f am ino ac id
compos i
tion andis common ly used as a“ reference pro
tein”
. The
156 CHAPTER6 NUTR
ITIONIN TROP
ICAL AQUACULTURE

am ino acid usua llyin greatest defici

tin a pro te
inis called the
“l
im iting” amino acid . The percen tage of am ino ac
id ofg reatest
def
icitissub tractedf rom 100tog ivethechem ica
lsco reforspec i
fic
protein source. For example:Ap ro tein withlysine contento f 40%
Tab
le6
.1 oftha tin egg pro te
in w i
ll have a chem ical score of 60
, provided
Essen
tia
laminoacidind
ices tha tthisisthe am ino acidin grea test def
ici
t.
(EAAI
)o fsomecommon
feeds
tuf
fsforsh
rimp c
. Essen
tia
l am
ino ac
idindex(EAA
I). The EAA
I me
thod o
f
Feeds
tuf
fs chem ical
lyevaluat
ing a p ro
tein conside
rs a l
lessen tial amino ac
ids
EAA
Iva
lues
ratherthan the mos tlim i
ting am ino acid w ith respectto some
Wh
itef
ish mea
l 0
.96
standard. EAA Iisthe geome tr
ic mean o fthe am ino ac idfound by
Pe
ruv
ianf
ish mea
l 0
.92 compa ringthe contento ftheten essen tial am ino acidsin afeed
S
lipmou
thf
ish mea
l 0
.94 prote
in w i
ththatfoundinf ishtissue protein. The c loserthe EAAI
Tunaf
ish mea
l 0
.92
value to 1.0the betteristhe pro tein qua lity
. Th is equat
ionis
expressed algebra
ically as:
He
rringf
ish mea
l 0
.95

Sh
rimp mea
l 0
.98

Squ
id mea
l 0
.96 …
Soybean mea
l 0
.87 I = 10√ 100ax 100b
EAA 100j
,
ae b
e je
Ip
il
-ip
illea
f mea
l 0
.54

Swee
t po
tato mea
l 0
.53

Sou
rce
: Pena
flo
rida 1989
in which a,b,...j arethe percentofessentia
l amino ac
idsinthe
food pro
tein and ae,be
,…jea rethe percen
to ftherespective amino
acidsin whole egg prote
in. Fo r computa
tion,itis convenientto
expressthe equa t
ioninloga r
ithm icform as
:

log EAA
I=
1
10
l
o (
g 100a+
ae
log100b +…
be
lg100j
o
je
.
)
Examp les o
fessential amino ac
idind ices o
ffeeds tu
ffs a
re
in Table 6.1
. Thus p rote
inf romip i
l-ip
illeaf meal and sweet
potato mealareofpoorqua li
tycompa redtofish meals
, shr
imp
meal
, squid meal, and soybean meal.

C
. Me
thodso
fLip
id Eva
lua
tion

1. Measuremen tofl
ipidqua lity
.Thequa li
tyo fl
ip idcon tained
in feedstu ffs and feeds is de term ined by i ts fatty ac id
compos i
tion wh ichis obtainedthru gas chroma tog raphic(GC )
analys
is(F igu re 6
.8)
. I tis atechnique o f separating samp le
into its cons t
ituents and then measur ing orident ifying
componen tsin same way . The separationtechn iqueinvo lves
F
igu
re6
.8 two phases , the sta
tionary and mobile.
The gasch
roma
tog
raph(GC
), anins
trumen
t The samp leis carried by the mob ile phase, usua lly an
usedfo
rana
lys
is o
ffa
ttyac
idcompos
it
ion
.
inert gas, through the s tat
ionary phase (co lumn ), where
 CHAPTER6 Eva
lua
tion o
f Feeds
tuf
fs and Aqua
feeds 157
separa tion takes p lace. S incethetechn ique
requ irestha tsamp les shou ld bevo latile,l ipids
are conver ted in to fa t
ty ac id me thy l es ters
(FAME ) beforein jection.
The measurement o f fatty ac ids uses
detec tors and the mos t popu laristhe F lame
Ion ization De tector. Samp lecomponen tse luting
fromthe co lumninthe gas s tream a reion ized
crea ting a curren t tha tis measured by an
electrome ter. The peaksind icate a measu re of
thefa tty acid concen trat
ioninthesamp le. The
peak re tentionst imes a re usedtoiden tifythe
componen tfa tty ac ids based on a s tandard
fatty ac id m ixture .
Samp le p repa rat
ioninvo lveslipidex t
rac tion
from the feeds orfeeds tuffs (Append ix B3 )
followed by sapon i f
ication and es terifica tion
processes , as desc r
ibedin Append ix B4 . The
output from th is GC is represented by a
chroma togram shownin F igure 6.9.

2
. Tests for l
ipid ranc
idity
. Oxidative
deter
io ration ofl
ipids has beenshowntocause
rancid f lavorin s tored feedstu
ffs andfeeds .
Lipid sources such asf ish and vegetable oils
which a re common ly usedinthe p reparation F
igu
re6
.9
A samp
le gas ch
romatog
raphic analys
is show
ing
offeeds a rerichin po lyunsatura
tedfa tty acids
re
tent
iont
imes ofvar
iousfa
tty acidsinafeedsample
.
and a re suscep t
ibleto ox idat
ion. The deg ree
of oxida tion in feedstuffs and feeds can be
determ ined through thefo l
lowing me thods:

a
. Pe
rox
ide va
lue(PV
). Perox
ide va
lueis de
fined asthe reac
tive
oxygen con tent expressed interms o f mil
liequivalen
ts (meq )of
freeiod ine pe r kg offat. Peroxides are precursors of breakdown
produc ts that cause ranc idflavorsinfa t. The concen tration of
perox ides ind icates ox idat ion dur ing ear ly stages o fl ipid
deter
iora t
ion . Th isindex becomesless re l
iable during thela ter
stage o f de
teriora t
ion, because perox ide degradationincreases .
PVis de term ined byt it
ratingtheiod ineliberatedf rom potassium
iodide w i
th sod ium thiosu l
fate solu
tion . Details o
fthe me thod o f
peroxide value de terminationis shownin Append ix B5.

b
. F
ree fa
tty ac
id va
lue (FFA
). Free fat
ty ac
id va
lues may be
de
termined us ingthet it
rime tr
ic me thod. Thef reefa tty ac
id value
is usual
ly calculated as ole
ic acid by divid
ingthe ac id value by 2.
The presence o fthese f ree fat
ty ac ids can speed up ox ida tive
dete
riorat
ion. O ils w
ith highcon tento ffreefat
ty acids will develop
undesirable color andflavor. Fo r de ta
ils ofthe procedures , see
Appendix B6.
158 CHAPTER6 NUTR
ITIONIN TROP
ICAL AQUACULTURE

c
. Th
ioba
rbi
tur
ic ac
id numbe
r(TBA
). Th
is me
thodinvo
lves hea
ting
the oilw ithth ioba rbituric ac idinthe p resence o fa st rong ac id . A
red colora tionis p roduced as a resu lto f areac tion be tween the
thiobarb ituric ac id and ma lona ldehyde. Thela t
te r may be p resen t
inthe o i
lo ri t may be p roducedf rom an ox ida t
ion p roduc t du ring
the course o fthe reac tion . The intens ity o fthe red co loris
proportiona ltothe concen tration o f malona ldehydeinthe m ix tu re
and can be measuredin a spec tropho tome ter. Resu lts may be
expressed as mg o f malona ldehyde pe r kg o fo il
.I tis necessa ryto
carry ou tthereac t
ioninthe absence o foxygenin o rde rto ob ta in
rel
iableresu l ts w ith ma rine o ils. Th isis a sens it
ivetes t and can
be correlated w iththe deve lopmen to fo ff-odo rs andf lavo rs. I tis
especially su i tedforthe de tec t
ion of ox idative ranc idityinl ipids
which a re unsa tu rated and con tain 3 or mo re doub le bonds . Fo r
detai
ls o fthe p rocedu re, see Append ix B7 .
The abso rbance o f a 1g samp lein 100 m l reagen t mu l
tip lied
bythefac tor 46isthe TBA numbe r,o rthe mg o f ma lona ldehyde
per 1000 g o fsamp le. Asthe amoun to freagen t usedis on ly 20
ml
,theresu lt mus t be mu l
tiplied by 0.2to g ivethe abso rbance o f
the samp lein 100 m l reagen t as specified bythe de finit
ion.

D
. Me
thodo
fVi
tam
in Eva
lua
tion

High Performance L iquid Chroma tography(HPLC )

. The HPLCis a
powe r
ful analyt
ical method ortoolw ith a wide range o
f applica
tions.
It provides h
igh resolu
tion, sensi
tiv
ity, and automa t
ic opera
tion, and
is eff
icientin de termining the amoun to fvitam ins contained in
feedstuffs andfeeds(F igure 6.10
).

F
igu
re6
.10
Asamp
le HPLCch
roma
tog
ram o
fav
itam
in m
ixtu
re.
 CHAPTER 6 Eva
lua
tion o
f Feeds
tuf
fs and Aqua
feeds 159
E
. Me
thodso
fMine
ral Eva
lua
tion

There a
re qual
ita
tive and quan
tita
tive me
thodsfo r de
term
ining m
ine
rals
infeedstu
ffs
. Some o fthe qua
lita
tive methods a
re:

1. Spottes tsforp resence o fsome m inerals

.The m inerals o
rino rganic
compounds used in an imal feed either occurin na ture or are
chemica l
ly compounded . To tes t for the presence o f minera ls
quali
ta t
ively
, a spo ttes tis usua l
ly applied. The m inera
lsin m ixed
feed are powde red ands ieved
. Thef ine po r
tionis placedinchlo roform
in a beake r
. Thef loating materialis poured off andthe res idueis
sprinkledinthef i
lter paperw i
th a sma ll spatu
lafo r spottesto fthe
fol
low ing m inerals.

. Coba
a lt, Coppe r, and I ron. To testforthe presence o fthese
minerals, af
ilter paperis mo istened w
ith solu tion A(100gsod ium
potassiumta rtrate dissolvedin 500 mld ist
il
led wa ter)
. Thesamp le
is sprinkledinthef ilte
r pape r and 2-3 drops o fsolution B(1g o f
1-nitroso-2-hydroxynaph thalene-3,6
-d isulfon ic ac
id) are added .
Thef il
ter paperisthen d ried. Cobaltg ives a p ink color; copper
gives alight brown co loredring; andi ron gives a deep green colo
r.

. Mangane
b se. Totes tfo rthe presence o f manganese ox
ide, sulfate
,
and carbona te
, af ilter paperis mo istened with solu
tion A (2 N
NaOH ). Thetes t samp leis sp r
ink ledinthe pape r and 2-3 d rops
ofso lut
ion B(0 .07 g benz idine dihyd rochlor
idein 10 m lofg lacia
l
acetic acidis d i
lu ted w i
th 100 m ld is
til
led water
). Manganese
oxide gives a dark b lue co
lor w i
th b lack center and manganese
sulfate gives alarge r spoto fl
igh terb lue which appears quickly.

. Iod
c ine , Magnes ium , and Z inc . Totes tforthe p resence o fiod ine,
magnes ium , and z inc, the same procedure as aboveisfo llowed ,
excep ttha ts tarch paper mo istened w ith brom ine so lution(1 m l
ofsa tura ted brom ine wa ter made upto 20 m lw ith distil
led wa ter
)
isthe one spr inkled w ith samp lefortheiod inetes t. Iod inethen
gives a b lue -purple colo r.
Inthe case o f magnes ium , so
lu t
ion A(1 N KOH ) and B(12 .7g
iodine and 40 g K Id issolvedin 25 m ld is
tilled wa ter and then
dilutedto 100 m l
)a re m ixedto g ive a very da rk brown co lor. A
sma ll par to fth isistaken and aga in 2 -3 drops o f solu tion Ais
added un ti
li tturns pa le yellow andth isis sp r
ink ledinthef ilte
r
papertoge ther w i
th the samp le. Magnes ium g ives ye l
low brown
spots .
For zinc , thef i
lter paperis mo istened w ith so lu tion A(2 N
NaOH ). The samp leis sp r
ink ledinthef i
lter pape r and 2 -3 d rops
ofso lu t
ion B(0 .1 g d i
th izonein 100 m lo fcarbonte trach lo ride)is
added . Z inc gives araspbe rryred co lor.
160 CHAPTER6 NUTR
ITIONIN TROP
ICAL AQUACULTURE

2
. M
ine
ral compos
ition
. Theresu
lting ashf
romthe c
rude ash ana
lys
is
is usedfor minera
l analysis o
fthe samp
le. The quan
tita
tive me
thods
for minera
l evaluation are
:

a
. T
itr
ime
tric me
thod
. In thet
itr
ime
tric me
thod
, permangana
te
ti
trationis done. In ca lcium ana lys
is, ca
lciumis prec
ipita
ted as
oxalate at pH 2.5-3,d issolvedin sulfur
ic acid
, andtheliberated
oxalic acidis titrated w i
th standard po tassium permangana te
solution
. Theleve lof phospho rusinthefeedis de te
rmined based
on M isson’sreaction.

b
. Spectrophotometr
ic me
thod
. The phosphorus present as
or
thophosphatereac ts wi
thvanada te-molybdatereagen tto produce
ayellow
-orange comp lex whichis measured spectrophotome tr
ical
ly
at400 nm. Calcium and phospho rus de
termina t
ionsa reconduc ted
onfeeds such asfish mea ls, bone meal
, calc
ium phospha te, and
calc
ium carbona te sources.

c
. A
tom
ic em
iss
ion spec
troscopy
. Th
is me
thod ut
ilizes h
igh
-
tempera ture atom ization sourcesto de te
rm ine the concen tration
of about 70 elemen tsinfeeds tuffs
. This operates onthe p rinciple
that when a subs tance is exc ited by a p lasma or e lectric
discharges,e lemen ts present em i
tlighta t wave lengths tha ta re
specificfor each e lemen t
. Thel ight em it
ted is dispersed by a
prism monoch roma tor
. The spec t
rallines produced a re recorded
on a pho tograph ic plate thatisl inked directly to a computer -
driven data processing system. The samp les are pre-concen t
rated
and mos telemen ts can be determined atthelow pa rts perb i
ll
ion
level.

d
.Flame spec
trome
tric me
thods
. The qua
lita
tive and quan
tita
tive
means o fthese me thods can be appliedto plant mate ria
ls ,plant
nutr
ien ts
, soi
ls, and otherbiolog
icalfluids
. The specificf requency
ofrad iation, emitted or absorbed, identi
fies the e
lemen t
. The
intensity ofemitted or abso
rbedrad iation a
tthe specificf requency
is propo rt
ionaltothe amoun to fthe elemen tpresent
.

e
. Atom
ic abso
rpt
ion spec
tromet
ry(AAS
). Th
is me
thod uses
combus t
ion and is used to observe the atom ic vapor tha
tis
produced when a samp le so
lut
ionis nebu l
ized and passedin to
theflamein an a tom ic abso
rpt
ion spect
rome te
r. Th isins
trumen t
consis
ts ofa cent
ralized hooded a
reain which ala rgeflame,o
ften
6in w ide by 6in high,islocated. This can be used toidenti
fy
mine
rale lements presentinfeeds andfeeding redien ts
.

F
. Me
thodso
fEne
rgy De
term
ina
tion

The ca
loric conten
to fthefeed sampleis dete
rminedthroughthe use o f
a bomb calorimeter
. The p r
incip
leinvolvedin bomb calor
imetryistha t
the heato f combus t
ionis measured, whenthe heat exchange process
takes p
laceinthe wa te
r conta
inedinsidethe calo
rimeterjacket
. Bomb
 CHAPTER 6 Eva
lua
tion o
f Feeds
tuf
fs and Aqua
feeds 161
calorime try genera lly measures BTU ’s, calories, orjou lesin so lid or
liquid fue ls and combus tible samp les as we ll. I tis a sys tem tha t
determ ines d irec
tlythe ca loric value o rg ross ene rgy o fthe samp le.
To de te rminethef rac t
ion o ftheg rossene rgytha tthef ish can actua l
ly
util
ize , a me tabolism tr ial is conduc ted to de term ine, d igestible,
metabo l
izab le
, and ne t ene rgyva lues . The use o f a bomb ca lo
r imeteris
necessaryto avo id diff
icu lties andinaccurac iesinhe rentin de term ining
the NFE . Subsequen tly,s im ilar de term inations on thefeca l residue
from such d iets makei t poss ibletoco r
recttheg ross calo r
icva luefo rthe
appa rentlossin d igestion and ob tain va luesfo rd igestible energy. Th is
is a bette r measurethan g ross ene rgyfo rexp ress ingthe use fulene rgy of
feeds tuffs.

1
. D
iges
tib
le ene
rgy(DE
). Aniner
t ma
ter
ial
, chrom
ium ox
ide
,is
addedtothefood a t alevelof0.5-1%. Thefish a
refedthe dietfor
seve
ral daysforfecal col
lect
ion, andthe ch
rom ium oxidelevels o
f
both thefeed andfeces a re determined
. The d iges
tible energy
(DE)is defined asfol
lows:

DE(kca
l/100 g d
iet
) =ene
rgy o
fdie
t – ene
rgy o (
ffeces x
mg C
r2O3
mg Cr2O3
/g dry d
iet
/g dryfeces )
2
. Me
tabo
lizab
le ene
rgy(ME
). The MEisthe por
tion o
fthe g
ross
energyconsumed bythef ishforg rowth,fa
ttening,o rheatp roduct
ion .
It does not appearinthefeces o rinthe u r
ine off ish
. The diges
tible
energy m inusthe ene rgyinthe u r
ine andfecesisthe ME , which
provides a bette
r measu re o
fthe ene rgyvalue o fthefeedthanthe
d
iges t
ible energy
. Thelarge dif
ferencesinthe e f
ficiency o
fu ti
liza
tion
of ME a re dueto wideva r
iat
ionsin hea tlosses.

3
. Ne
t ene
rgy(NE
). Th
is represen
ts the mos
t prec
ise measure o
f
the energy needs o
ffish andthe capac i
ty ofdif
ferentfeedsto meet
these needs. No t only do NE values a llowforthe ene rgylostin
the urine and feces, bu t they also takein to considerat
ion the
energylost as heat during nu tr
ien tu t
iliza
tion. Howeve r, ac
tual
NE values have been de termined for on ly alimited number o f
feeds
. Thus , most avai
lab le NE values are only es
timates
.

G
. Ana
lys
iso
fTox
insinFeeds

Somefeedingredien
ts con
tain natura
ltox
instha
ta re g
rowthinhib
itory
athighleve
ls and may be de
lete
rioustofish
. Methods o
f de
termining
thesetox
ins are
:

1
.Ureaseac
tiv
ity
. Theenzyme urease usua
llyfoundinrawsoybeans
producestoxic
itythroughthe hyd ro
lysis o
fureato ammon ia
. Hea t
treatmento f soybean meala t 120°Cfor 20 m inis adequate to
remove this enzyme. Quan t
itative analysisto dete
rm ine urease
act
ivi
tylevelinfeeds tu
ffsis pat
te rned after Chow(1980).
162 CHAPTER 6 NUTR
ITIONIN TROP
ICAL AQUACULTURE

2
. Gossypo
l. Th
isis an endogenoustox
in presen
tinthe g
land o
f
cot
tonseed which persis
ts during mealproduct
ion unlessremoved
byaspecia
lp rocess,or, un
lessthecottonseedisaglandlessvar
iety
.
The gossypo l level in a feed ingred ient is de termined
spect
rophotome tr
ically.

3
.Af
latox
in. Th
isis a c
lass o
f po
ten
ttox
ins produced bythe mo
ld
Aspergi
llusflavus andis usua lly presentinfeed ma ter
ials such
as groundnu t cake, copra, peanu ts, corn,rice
, and legumes
produced and s tored under ho t and hum id cond i
tions. The
chroma tographic me thods most w idely and routinely usedfor
af
latoxin ana lysis are one - and two-d imensiona l thin layer
chromatography (TLC ) and h igh per formance th in layer
chroma tography(HPTLC )
. In v iew ofthe po tentia
l deleterious
ef
fects ofthese an tinutri
tionalfac tors on the growth o ffish ,
analys
isshou ldbedoneonfeeds tu f
fsknowno rsuspectedtocon ta
in
these ma ter
ials.

M
icrob
iolog
ica
lEva
lua
tion

Am
ino ac
id compos
ition
. Another me
thod o
f de
term
ining am
ino ac
id
compos it
ionis through the m icrob iological me thod. Th is me thodis
valuablein ana lyzing m ixtures o f am ino ac ids because o fthe speed and
reproduc ibil
ity o fresu l
ts ob tained . A nu tr
ien t med ium wh ich con ta ins
allo fthe essen tial compounds neededforthe grow th o f a particu lar
microorganism excep tthe am ino ac idto be assayedis p repared. Add ition
ofth is amino ac id resu l
tsin g row th ofthe m icroorganismin p ropor tion
tothe amoun to fam ino acid added . Cu lturetubes a reset up andg raded
amoun ts ofthe unknown a re addedto a se r
ies oftubes . S tanda rds a re
set up a tthe samet ime w ith g raded amoun ts ofthe pu re am ino ac id.
The unknown can be compa red w ith standa rds by measur ingthera te o f
grow th ofthe m icroorganism . W i
th o rgan isms wh ichform ac id such as
Lactobacil
li,t i
tra t
ion o fthe ac idfo rmed can be used as a measure o fthe
numbe ro fcells p resent. Pu re cu ltu
res needto be usedinth is kind o f
evalua t
ion.

B
iolog
ica
leva
lua
tion

Afeedingexpe r
imen tisconductedtotes ttheeff
icacyofformu latedfeeds
.
Itis usua
lly doneintanks, ponds,o rcages. In alaboratoryexpe r
iment,
environmenta l condi
tions are eas
ily kept constan
t. In carrying outa
feed
ing experimen t
, thefol
lowingfac to
rs haveto be considered:

a
. The ob
jec
tive o
fthe s
tudy hasto be c
lea
rly de
fined
.

b
. Experimen ta
ltreatmen ts and statis
tical design approp r
ia tetothe
objective ofthe exper imen t have to be care fully selec ted. A
comp lete
lyrandomized des ign(CRD )o rrandom ized comp lete b
lock
design (RCBD )is usua lly app
liedin mos tfeeding exper imen ts
.
Howeve r
, whenthere are mo revariables
, afac tor
ial designis used .
Prior to afeeding exper iment the number o f replications per
 CHAPTER 6 Eva
lua
tion o
f Feeds
tuf
fs and Aqua
feeds 163
treatmen t andthe numbe ro ff ish pe rrepl
icate should be chosen
such tha t a mean ing
fu l statistical ana lysis of data can be
accomp lished a
t ase tlevelo fsignificance
.I tis bes
ttha tthe data
to be ga thered , samp l
ing f requency , number o f samp les per
replica
te, and s ta
tist
ica
l me thodsto be usedinthe da ta ana lys
is
are known be forehand. As tatist
ician should be consultedforthe
design ofthe expe riment
.

c
. Theexpe rimen talf ishspec ies havetobeiden ti
fied withi tssc ien t
ific
and common names , toge ther w ithits strain, source ,s ize, age ,
and prev ious nu tri tiona lh istory. Acc lima t
ion o fthetes t an ima l
is usua lly donefo ra tleast a weekto enab lethef ishto ad justto
its new env ironmen t. In it
ial body we ight andleng th haveto be
measured dur ing s tock ing
. I tis bes tto use an equa l numbe ro f
fish o f approx ima te ly the same s ize and age per rep licate. In
some s tud ies
, bodycompos it
ion( initia
l andf inal) hasto be known .
This is usua lly de term ined in three groups o ffish in three
replicates.F ish a re usua lly we ighed, sacrificed, chopped ,d r
ied
inaf reeze drier, and oven -dr
ied a 60 °Ctocons tan t weigh t
. Resu lts
of proxima te compos itionisthen re lated tothein itial we ights o f
thef ish to ob tain abso lute we ights ofthe nu trient componen ts.
The same procedure is fo llowed to de term ine the f ina l body
compos it
ion o fthef ish . The da tais usedto de termine nu t
rien t
retention .

d
. Theexpe rimen taltanksshou ldbela rgeenoughtoa llowsubs tant
ial
fish g row th(F igure 6 .11). Expe r
imen taltanks,d imensions, wate
r
sou rce, vo lume , and dep th should be c lear
ly de
fined. Wa ter may
be s tatic bu t rep laced regu larly
, rec i rculat
ing, orflow-through
depend ing on the ava ilable resources .
Fil
tra tion sys tem shou ld be adequa te and
eff
icien tin remov ing pa r
ticula te ma tte r and
metabo litestha t mayin fluencetheresponse
offishtot rea tmen ts
. Wa tertempe ra tureis
usua lly amb ien t and some t
imes con trolled.
Water qua lity ( temperature , d isso lved
oxygen , pH , am mon ia, e tc.) shou ld be
mon itored da ily and shou ld befavo rablefo r
maximum g row th.

e
. The durat
ion ofafeedingexperimentshou ld
belong enough,a tleast 8 weeks
, to al
low
fishto man i
fes
t defin
ite g
rowtht rends and
signif
icant d i
fferences in response
parame ters as affected by the d ie
tary F
igu
re6
.11
Alabora
toryse
t-upfo
rafeed
ing expe
rimen
t us
ing 250l
treatments.
tanks
.
164 CHAPTER 6 NUTR
ITIONIN TROP
ICAL AQUACULTURE

f
. Growth , the mos t wide ly accepted response parame ter for
evaluating treatments infeed ing
, is a sensit
ive and prac t
ica l
indica
to rofthe adequacy ofessent
ial nutr
ientsinthe diets
. O ther
parameters such as surviva
l,feed eff
iciency
,e tc. shou
ld also be
takenin to account
.

Pa
rame
terstobe mon
ito
redinafeed
ingexpe
rimen
t:

1
. G
row
th

a
) Abso
lute g
row
th

wt ga
in =wf- w
i,
where: w
f =f
ina
lw t
; w
i =in
itia
lwt

Abso
lute g
row
th ra
te

w
t ga
in/day = wf- wi
days o
f cu
ltu
re
b
) Re
lat
ive grow
th

w
f- wi
%w
t ga
in = x 100
w
i
Rela
tive grow
th ra
te

%w
t ga
in/day = wf- wi
x 100
(w
i)(days o
f cu
ltu
re)
Spec
ific Grow
th Ra
te(SGR
)
ln w2-ln w1
SGR =

where
: w1 and w2 are we
ights a
t per
iods 1(
t1 and 2 (
t2)
,
respec
tive
ly.

2
. E
ffic
iency o
ffeed u
til
iza
tion
d
ryfeed consumed
Feed conve
rsion ra
tio(FCR
)=
w
t gain

wt ga
in
Feed e
ffic
iency(%
)= x 100
d
ryfeed consumed
3
. D
iges
tib
ili
ty o
f nu
trien
ts

% D
iges
tib
ili
ty = nutrien
t absorbed
x 100
nu
trient consumed
4
. E
ffic
iency o
fpro
tein u
til
iza
tion
wt ga
in
Pro
tein E
ffic
iency Ra
tio(PER
)=
pro
teinin take

B
iolog
ica
l Va ) - N re
lue(BV ta
ined
N absorbed
 CHAPTER 6 Eva
lua
tion o
f Feeds
tuf
fs and Aqua
feeds 165
Ne
t Pro
tein Va
lue(NPV
)or Produc
tive P
rote
in Va
lue(PPV
)

= N re
tained o
r = BVx d
iges
tib
il
ity
N consumed

Net Pro
tein U
til
iza
tion(NPU
)

n
itrogeninc reaseinf ish n
itrogen dec reaseinf ish
= fed thetes
t p
r o
tein d
iet + fed the pro
t einfree d
iet
x 100
ni
trogenin takef romthetes t p ro
te in die
t

5
. Su
rviva
l ra
te

% Surv
iva
l= f
inal count
x 100
init
ial coun
t
6
. P
rox
ima
te compos
ition o
ffishsamp
les(
ini
tia
l andf
ina
l)

7
. B
iolog
ica
l pa
rame
ters
a
) am ino ac
id andfa tty ac
id compos it
ion
b
) stored nutr
ientlevelsint issues, serum,orplasma
c
) enzyme ac tiv
ity
d
) oxidat
ion ofrad ioact
ively-
labelled nutr
ien
ts

8
. H
isto
log
ica
l changesint
issues(e
.g.g
ill
s, sk
in,l
ive
r, musc
le)

Summa
ry

The cho ice ofh igh qua l

ityfeeds tuf
fsfo rincorporat
ionin to aquafeedsis
cruc ialto the success o f an aquacu l ture venture. Proper me thods of
feeds tu ffandfeed eva lua tion shou ld belea rned and app l
iedto a tta
in an
effectivefeed . The sys tema tic evaluation offeeds tu
ffs andfeedinc ludes
phys ica l
, chem ical, and b iological me thods.Inthe app l
ication ofthese
me thods ,s tandard samp ling procedures shou ld befo l
lowedin o rderto
effectively carry ou tthe eva luat
ion p rocess. Amongthese me thods, the
biologica l method wh ichinvo lves actualfeedingexpe riments gives a more
accura te est
ima te o
ffeed u til
ization.

Gu
ide Ques
tions

1
. Wha
tarethe d
iffe
ren
t me
thods o
ffeed eva
lua
tion? D
iffe
ren
tia
te one
f
romthe o
the
r.
2
. Wha
t does prox
ima
te ana
lys
is measureinfeed eva
lua
tion? Wha
t
a
rethel
imi
tat
ions o
fth
is procedure?
3
. Wha
tis the pr
inc
iple beh
ind the “K
jeldah
l me
thod” o
f pro
tein
de
term
ina
tion?
4
. Howisthe pro
tein qua
lity eva
lua
ted?
5
. Descr
ibe br
ief
ly how to ca
lcu
late the essen
tia
l am
ino ac
idindex
(EAA
I)o
fa p
rote
in.
166 CHAPTER 6 NUTR
ITIONIN TROP
ICAL AQUACULTURE

6
. Exp
lainthe p
rinc
iple beh
ind gas ch
roma
tog
raphy
.
7
. Wha
tarethetes
ts usedto de
tec
tlip
id ranc
idi
ty?
8
. Wha
t are perox
ides? D
ifferent
iate be
tween perox
ide va
lue and
th
iobarb
itur
ic ac
id numbe
r.
9
. Wha
tistheimp
lica
tion o
f hav
ing a h
ighf
reefa
tty ac
id va
lueinfeed
ma
ter
ials?
10
. Wha
tisthe s
ign
ificance o
ftheth
ioba
rbi
tur
ic ac
id numbe
ror TBA
numbe
r?
11
. Exp
lainthe p
rinc
iple beh
ind a
tom
ic em
iss
ion spec
troscopy?
12
. Wha
tarethe cha
rac
ter
ist
ics o
fa h
igh qua
lityfeed?
13
. Howis a
fla
tox
in p
roducedinfeed/
feeding
red
ien
t?
14
. Wha
tarethe an
tinu
tri
tiona
lfac
torstha
t may be presen
tinthefeed
ingred
ien
t and exp
lain how they are m
inim
ized or removedf
rom
feeds
tuf
fs.

Sugges
ted Read
ings

ADCP
. 1980
.Fish FeedTechno
logy
. Rome
, FAO
, ADCP/REP/80/11
, 395 p
.

AOAC(Assoc
iat
ion o
fO f
ficia
l Analy
tica
l Chemis
t). 1980. O
ffic
ial me
thods
of ana
lysis
.(13th edi
tion
). Washing
ton
. 1108 p .

Asakawa T
, Nomura Y
, Matsushita S. 1975
. A modif
ied TBAtes
tfo
rthe
determ
ina
tion ofl
ipid ox
idation. Yakagaku 24
:481-482
.

B
ligh EG
, Dye r WJ. 1959
. A rapid method ofto
tallipid ext
rac
tion and
pur
ifica
tion
. Can.J.B iochem. Physio
l. 37:911-917.

B
lock RJ
, Mitche
ll HH . 1946. The correlat
ion o f the am
ino acid
composit
ion of prote
ins w
ith the
ir nutr
itive va
lue. Nutr
. Abst
.
and Revs
. XVI
:249 .

Cheeke PR. 1991

. App l
ied anima l nutr
ition
: feeds and feeding.
Depa
rtmento f An
imal Sc
ience
, Oregon State Un
ive
rsi
ty. 504 p
.

Chow KW. 1980. Qua li

ty controlinfishfeed manu fac
tu r
ing. IN:F ish
FeedTechnology
, FAO/UNDPTra ining Course
, College o
fFisher
ies,
Unive
rsi
ty o
f Wash ington
, Seatt
le, 9 October
-15 Decembe r 1978.
ADCP/Rep/80/11 , p 369-385
.

Chu
rch DC, Pond WG. 1974
. Basic an
imal nutr
ition andfeed
ing
. O and
B Books, 1215 NW K l
ine P
lace, Co
rval
lis, Oregon 97330
, Un
ited
States o
f Amer
ica. 300 p
.

Cho CY, Cowey CB, Wa tanabe T

. 1985 . F
infish Nutr
ition in As
ia:
Methodo
log
ica
l Approachesto Resea
rch and Deve
lopment
. O t
tawa,
Ontar
io.IDRC. 154 p
.
 CHAPTER 6 Eva
lua
tion o
f Feeds
tuf
fs and Aqua
feeds 167
CRC Handbook o
f Chroma
tography
. CRC P
ress
, Inc
.,F
lor
ida
. USA
.2:
319
-355.

Cramp
ton EW , Harr
is LE
. 1969. App
lied animal nu t
rit
ion
; the use o
f
feeds
tuffsintheformula
tion o
flivestock ra
tions. W
.H.F reeman,
San Franc
isco
, USA
. 753 p.

Feed Deve
lopmen t Sec
tion
. 1994. Feeds and Feeding o
f M
ilkf
ish, Ni
le
Ti
lapia
, As
ian Sea Bass
, and Tiger Sh
rimp. SEAFDEC Aquacul
ture
Department
.T igbauan,Ilo
ilo
, Phil
ippines
. 97 p.

Fo
lch J
, Lees M ,Stanley GH. 1957
. Asimple me
thodfo rtheisola
tion
and pur
ifica
tion ofto
tall
ipidsfrom an
imalt
issues
.J .Bio
l. Chem.
226
:487-509.

F
ish Nutr
ition and Mar
icultu
re. 1988
. The Genera
l Aquacul
ture Course
,
AJ ICA Textbook. Wa tanabe T. (ed
). Kanagawa Internat
ional
Cooperat
ion Agency
. 233 p.

Has
tings WH . 1969
. Nutri
tiona
l Score
.In: Neuhaus
,O.W and J.E. Halve
r
(eds
).Fishin Research Academ
ic P
ress
,Inc. New Yo
rk. p 263
-292.

Has
tings WH , D
ickie LM. 1972
. Feedformu la
tion and eva
luat
ion. In
:
Halve
r,J.E.(ed).Fish Nutr
ition
. Academic P
ress
,Inc. New York
.
Chapter7, p 327
-370.

Jowaman, K, Duangsmorn,S , Ankana , H, Uthai

, K. 1987
. Manua lo f
feed microscopy and qua l
ity contro l
. Amer ican Soybean
Assoc
iation Na t
ional Renderers Assoc
iation and US Feed Gra
in
Council
. p 154
-160.

Love
ll RT
. 1984. Mic
rob
ialtox
insinf
ishfeeds
. Aquacu
lture Magaz
ine
10
(6A
):34
-36.

Mango
ld HK, Zweig G
, Sherma J. 1984
.L ip
ids. CRC Handbook o
f
Chroma
tog
raphy. CRC P
ress
,Inc.Flor
ida, USA
.

Mar
ine
tti
, GV. 1967.Lip
id chromatograph
ic ana
lys
is Vo
lume 1
. Ma
rce
l
Dekke
r,Inc.
, New Yo
rk. 537 p
.

Mar
ine
tti
, GV. 1967.Lip
id chromatograph
ic ana
lys
is Vo
lume 2
. Ma
rce
l
Dekke
r,Inc.
, New Yo
rk. 596 p
.

Maynard LA
, Loos
li JK
, Hin
tz HF
, Warner RG
. 1979
. An
ima
l Nu
tri
tion
,
7
th edn
. New York
, McGraw H
il
l.

Me
tca
lfe LD, Schmi
tz AA
, Pelka JR
. 1966 . Therapid prepara
tion offat
ty
ac
id me thy
l es
tersfroml ipidsfor gas chromatographic analys
is.
Anal
. Chem. 38:514
.
168 CHAPTER 6 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Mor
rison WR, Sm ith LM . 1964. Preparat
ion offat
ty acid me
thyl es
ters
and d
ime thylacetalsfroml ipids wi
th boron t
rif
lour
ide-methanol
.
J.Lipid Res.5 :600-608.

Ose
r BL
. 1951 . Methodforinteg
rat
ing essentia
l amino acid conten
tin
the nu
tri
tiona
levalua
tionofprote
in. J. Amer
.D ie
tetic Assn
. XXVI
I:
396
.

P
ierce JG. 1976 . Feed microscopy in qua l
ity control. In: Feed
Manu fac
tur
ing technology. Feed Product
ion Counc i
l, American
Feed Manufacturers Assoc
iat
ionInc .
, Ar
lington
,V irginia
, p 270-
273.

Roach AG, Sanderson P, Wi

ll
iams DR . 1967. Compar
ison o
f methodsfor
the determina t
ion ofavai
lablelysineva
luein anima
l andvegetab
le
pro
tein sources
.J . Sc
i. Food Agr
ic. 18
:274
-278.

S
impson RJ, Neube
rge
r MR,Liu TY
. 1976
. Comple
te amino ac
id analys
is
o
f prote
insfrom a s
ing
le hydro
lysate
.J. Bio
l. Chem. 251:1936-
1940
.

Tacon A
. 1987. The nutr
ition andfeeding o
ffa rmedfish and shr
imp.A
train
ing manua l
.2 . Nutrien
t sources and composit
ion
. FAO Fie
ld
Documen t
, Project GCP/RLA/075/ ITA,F ield Document, No
.5 ,
Brasi
lia
, Brazi
l. p 95-117.

Yu TC
,S innhuber RO. 1957. 2
-Th iobarb
itur
ic acid me
thodf rom the
measuremento franc
idi
tyinfishery p
roducts
. Food Tech
. 11:104
-
108.
 CHAPTER

7
ManagementofFeeding
Aquacu
ltu
reSp ecie
s
VERON
ICA R
. ALAVA

In
troduc
tion
Produc tion targe ts , env ironmenta l cond itions , and soc io-econom ic
cons iderat ions de term ine the leve lo f managemen tin aquacu l
ture
produc t ion sys tems . The con tribution o f na tura lfood as a source o f
nour ishmen tisimpor tan tin produc ingf ish w ithfewinpu ts. Howeve r
,
mo reinpu ts such as comp lete h igh qua lityfeeds a rerequ iredtoinc rease
fish y ield pe r un i
ta rea .
The use o f cos t-effec tive feeds , properfeed ing managemen t
, and
ma intenanceo fgood wa terqua l
itya rec r
itica lforasuccess fulaquacu ltu re
en terpr ise . As the s tock ing dens ityincreases , feed requ iremen t and
me tabo lic was tes a lsoincrease . Feed ing mus t be regular ly mon i
tored
and con tinuous ly ad jus ted so as no tto ove r
feed o r unde rfeed thef ish.
The success o f an a rtificiald ietis dependen t onthe managemen to fthe
feed onthefa rm . Feed ing prac t
ices mus tthe refore be d irec ted towa rds
op tim izingfeed u til
iza tionto p reven t pollu tion o fthe cultu re env ironmen t
andthe coas talv icinity .
Th is chap te rteachesthe reade rto :d if
fe rentiatethe d i
ffe rentfeed ing
stra teg iesin pond cu ltu re;lea rnfeed ing managemen t me thods such as
stocksamp ling andreco rd keep ing, calcu lating da i
lyfeedra t
ion , choos ing
appropr iatefeed s ize , and me thods o f app lyingfeeds ; unders tand the
impac to ffeed ing managemen t on wa ter qua l
ity and env ironmen t and on
the cu ltured an ima l’s g row th, surv ival, andfeed convers ion ra tio; and
descr ibe the d ifferen tfeed ing schemes usedto cu lturef ishes(m ilkfish,
tilapia , rabb itfish ,b ighead ca rp, native ca tf
ish , sea bass,o range-spo tted
groupe r, and mang rove red snappe r; and c rustaceans( tige r sh r
imp and
mud c rab )
. O the r spec iesfo r aquacu l
ture s tock enhancemen t(donkey ’s
ear aba lone , seaho rses , w indow -pane oys te r)a re also discussed .

Feed
ings
tra
teg
iesinpondcu
ltu
re

Fish production depends onthe s tocking density andfeeding strategy

used
.A tlowers tock
ing densi
ty asintheextensivecu l
turesystem, natural
pond produc tiv
ityisimpo rtant
. As stock
ing dens i
tyincreases asinthe
sem i
-intensive and intens ive sys
tems , art
ific
ial feed becomes more
essential(Figure 7.1)
.
170 CHAPTER 7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

F
igu
re 7
.1
Range o
faquacul
ture p
rac
ticesinre
lat
iontoinpu
ts.
Sou
rce
:Taco
n199
8

In pond aquacu
ltu
re,thefeed
ing s
tra
teg
ies used a
re:

A
. Product
ion o
fna
tura
laqua
ticfood

Dur ing the ear ly grow th s tages o fthe cu ltured an ima l and a tlow
stock ing density,thero le o f natu ral aqua ticfood , asthe on ly sou rce
of nu tr
itionis veryimpo rtant. The na tura lfood basesin ponds a re :
lab lab (a comp lex ofb lue -green and g reen a lgae ,d iatoms , ro tife rs,
crus taceans,insec ts, roundwo rms , detritus ,p lank ton(F igure 7 .2A ),
and lumu t( fibrousf ilamen tous green a lgae )(F igure 7 .2B ). Organ ic
or inorgan ic fert ilizers are
pe r
iod ica l
ly app l
ied as sou rces
of carbon , n itrogen , and
essent ial minera ls. Wi th
sun ligh t, fer ti
lizers enhance
the growth o f phy toplank ton
the rebyinc reasingthe na tura l
produc t
ivi ty and hence f ish
produc tion . An integrated
sys tem us ing var ious spec ies
in d if
feren tt roph icleve ls such
F
igur
e7 .
2 as seaweed , she ll
fish , and
Na
tural aqua
ticfoodin ponds
:lab
lab(A
) andlumut(B)
. herb ivorous and omn ivorous
fish spec ies such as m ilk fish
andt ilapia area lso usedtoe f
ficiently uti
lizeva riousfo rms o f nu trien ts
andto ob tain h igh produc t
ivity. In extens ive cu ltu re sys tems ,inpu ts
and produc tion cos ts are gene ra l
lylowe rthanin sem i-
in tens ive and
in tensive systems , bu ty ields a re alsolowe r
.

B
. Feed
ingasupp
lementaryd
iet

When stocking density and standing crop o

f cul
tured an ima
ls Is
increased
, the natural produc
tiv
ity can nolonger suppor
t adequate
 CHAPTER 7 Managemen
tof Feed
ing Aquacu
lture Spec
ies 1
71
growth. Supp lemen ta
lfeeds become necessa ry. The common ly used
supplemen ta
lfeeds a rerice bran, chickenfeed ,b readcrumbs , boiled
corn, cooked cassava , and chicken en trails
. These a re nutri
tiona lly
incomp lete and wou ld beinadequa teif used asthe on ly source o f
food. When supp lemen talfeedingis app lied,highers tocking densities
are possible result
ingin h igher production per unita rea. The mos t
cost
-effec t
ive way o f producing aqua t
ic an imalis to grow na tura l
food and to prov ide supp lemen ta lfeed when na turalfood becomes
inadequa te. As na turalfood dec reases w i
thincreasing standingc rop ,
the quality ofthefeed shou ld beimp rovedto sus tainfish growth.

C
. Feed
ingacomp
leted
iet

Ath igher s
tock ing densitiestypicalofin tensive culture
, a complete
die
tis necessaryto p rovidethe nu t
rients required bythe an imalfor
growth and su rviva
l.P roduction does notrely on na turalfood.Since
the aim ofh ighlevelo fp roductionisto max im ize y
ield per un
itarea
in a shorter period,h ighly e
ffect
ive ar
tif
icialfeeds a re used
.

Feed
ing managemen
t

Samp
lingandreco
rdkeep
ing

Regu lar samp ling of cultured s tocksis essen tialin o rderto assessthe
effectivity offeed ing management . Samp ling invo lves we ighing or
measur ing arepresen ta t
ive group(sub -samp le)o f an imals. These da ta
can then be used to de terminetheto talb iomass , andthe changesin
weigh t orleng th from prev ious samp ling da ta.A ll samp ling methods
shou ld be care fu
lly des igned to avoid bias, andto m inim ize any stress
onthe an imals. A work schedu le should be setp riortosamp ling
, and a l
l
equipmen t and ma terials should be on hand be fore anima ls are co
llected
and measured .Itisimpor tant thatthef ish hand l
ing and measur ing
procedures a re cons isten t and repeatable.
To es timate biomassinthe pond , tank o r cage,f ish are samp led by
using a cas t orlif
t ne t(F igure 7 .3 A,B). Whenf ishinla rge ponds a re

F
igu
re7
.3
Samp
ling bycas
tne
t(A
)andl
if
tne
t(B
).
172 CHAPTER 7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

samp led,the ne ts are castin severala reas o fthe pondin o rde rto ge ta
better samp le ofb iomass . For examp le,in a one -hec tare pond , a ne t
may be cas tinthe m iddle, and one ortwoineach s ide ofthe pond . The
samp les a rethen we ighedind iv
idua l
ly o rin bu lk. A good es t
ima te of
surv ival ra teis necessary to ca lculate to ta lfish b iomass bu t th isis
of
ten d i
fficultto ob taininthe ponds .In p rac tice, mos tfa rme rs assume
a cer tain va lue for surv ival based on da ta f rom prev ious cu lture
opera tionsin the same pond or o ther ponds w ith s im i
lar cond i
tions .
Thisva lueis mod i
fied byobse rvat
ions o factua l mo rtalit
ies, wa terqua l
ity,
and occur rence o fd iseases.
Data onfe rt
il
izers, anima lstock,feed , wa te r quality, and othe r usefu l
parame tersl istedin Tab le 7
.1 mus t be reco rded. Accura te reco rds a re
necessaryto enab lethefa rme rto assessthe e ff
iciency o ffeed ing and
farm managemen t duringthe pas t and cur ren t culture runs .

Tab .1 Some use

le7 ful pa
rame
terstha
t mus
t bereco
rdedinthefa
rm

Parame
ters

Pond Pond numbe

r,a
rea
, dep
th

Fer
ti
lize
r Sou
rce
, da
te o
ffe
rti
lize
r app
lica
tions
,type and quan
tity o
ffe
rti
lize
r used

An
ima
lstock Da
te o
fstock
ing
, sou
rce o
fstock
, spec
ies
, behav
ior
,stock
ing dens
ity
, ave
rage
in
itia
l we
igh
torleng
th, da
te o
f samp
ling
, da
te o
f ha
rves
t, ave
rage f
ina
l
we
igh
torleng
th, kg ha
rves
ted
, we
igh
t ga
in o
rgrow
th, su
rviva
lat ha
rves
t,
s
ize d
ist
ribu
tion a
t ha
rves
t

Feed Da
te o
f pu
rchase o
ffeeds
,feed
ing ra
te and amoun
t,type o
ffeed used
, da
te
eachfeedtypeis g
iven
,feed
ingf
requency
,feed
ingt
ime
,feed
ingloca
tionin
the pond
,presence o
fle
ft-ove
rfeeds
,feed conve
rsion ra
tio

Wa
ter qua
li
ty Secch
idisk read
ing (wa
tert
ransparency
), wa
ter exchange ra
te, sa
lin
ity
,
d
isso
lved oxygen leve
l, da
te o
f wa
ter change
, wa
tertempe
ratu
re, wa
ter
co
lor
,time and du
rat
ion o
f padd
lewhee
l ope
rat
ioni
f used

O
the
rs D
isease o
r abnorma
lit
ies o
fstock
,preda
tors in pond
, wea
ther cond
itions
du
ringthe g
row
ingcyc
le, unusua
leven
ts,cos
ts o
finpu
ts,inves
tmen
tre
turns

Feed
ing Ra
tion

The amoun to fthe da i

lyfeed ra tion , andthef requency andt iming o f
feeding areimpo rtantfacto rs affec t
ing g rowth andfeed conve rsion. Each
cultured spec ies has a dieta ryfeed ingra te opt
ima lforg row th andfeed
eff
iciency. The optima lfeedingleve lva ries wi
th age,fish biomass ,d ietary
nutrientrequ iremen t, water qua lity, and na turalfood ava i
lab il
ity. F ish
lose we igh t when the irfeedin take fa lls below the requ ired leve lfor
maintenance . As ra t
ion size andfeedin takeinc rease, the grow th rate
increases up to a cer tainl im it. Thelarva ls tages of aqua tic an ima ls
have very h igh me tabol
ic ac tiv
ity and rap id gastro-
intestinal evacua tion
rates. Thus,i tis crit
icaltha tthe i rh igh energyrequ iremen ts a re me t by
con t
inuousfeed ingto sa t
iation .F requen tfeedingisrecommended when
fish are sma ll
, when na tura lfoodisinadequa te
, and whenthefeeds a re
less wa ter-stable
.
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 173
Op t
ima l grow th andfeed e fficiency can on ly be a ttainedi f su fficien t
amoun to ffeedis prov ided , comp lete lyinges ted, and d iges ted . Pond -
rea red sh rimps mus t befedf requen tly on sma l
l amoun ts,w iththe bu lk
ofthefeed g iven dur ingthe n igh t when thefeed ing ac tivityis h ighes t.
Inc reas ingthef requency o ffeed ing reducesleach ing andfeedloss , and
improvemen to f grow th andfeed e f
ficiency . No rma lfeed ing pa tternsin
crus taceansa rein ter rup ted during mo lt
ing .Immed iate lybe foreand du ring
actua l mo lting,c rus taceans s topfeed ing . Th is usua llylas tsfo rtwo days
after wh ichfeed ingleve lsinc rease . Thus , an ad jus tmen tinfeed ingleve ls
is needed .
Various me thods are used to es tima te feed ra tion . These inc lude
calcu la tions based on the use o ffeed ing char ts, feed equa tions , and
grow th p redictions . Feed ingtab les a reca lcu latedf romes tima teso fg row th ,
su rvival, andfeed conve rsion . Tabu la tedfeed ingleve lsfo rva rious spec ies
are common ly p rovided byfeed manu fac turing compan ies. Da ilyfeed ing
rateis expressed as pe rcento f body we igh t pe r day . The pe rcen tage o f
feed consumedinre la t
ionto body we igh t dec reases asthef ish g row and
the ir me tabolicra tes dec l
ine. Feed ing cha rts provide on ly agene ral gu ide
tofeedin take. These do no ttakein to accoun t the sho rt-o rlong -te rm
fluctuat ions in appet ite in response to many phys io log ica l and
environmen ta l fac tors . However , they are use fu lin es tima ting feed
requ iremen ts. These base lineva lues ,la te r mod if
ied asfeed ing and g row th
da taf rom the var ious cu lture ba tches ,a re collec ted . C lea rly, op tima l
feed ing ra te andf requency mus t be de te rminedfo r eachfeed and pond
by care fu l mon i
tor ing o ffeed consump tion ,g row th, andfeed e fficiency
ove r seve ral grow ing cyc les.
In o rde rto ca lcu latethe da ilyfeedra tion o fthe cu ltu red an ima l
, da ta
on ave rage body we igh t andthe es tima ted surv ival ra te a t ag ivent ime
are needed , andthesetwo a re ob tained bysamp l
ingthes tock . Thefeed ing
rales recommendedfo rthe d i
fferen t spec ies a t va r
ious s tages o f cu lture
are g iveninthe sec tion on Feed ing Schemes . Thefo rmu lasto compu te
fo r ave rage body we igh t
, surv iva l rate, and da ilyfeed ra tion a re:

to
talw
tofsamp
led s
tock(g
)
Ave
rage body w
t(g
) =
numbe
rof samp
led s
tock(pcs
)

numbe
roff
ish a
ta g
ivent
ime(pcs
)
Surv
iva
lra
te (%) = x 100
o
rig
ina
l numbe
roff
ish s
tocked (pcs
)

or
igina
l numbe
r ave
rage body es
tima
ted feed
ing
s ) X
tocked (pcs w
t(g
) X su
rviva te(%) X
lra ra
te(%
)
Dai
lyfeedra
tion(kg
/day
)=
1
,000
174 CHAPTER 7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

FeedPa
rtic
leS
ize

Feed par tic

les mus t be ofas izetha t can be read ilyingested and th is
inc reases asfish grow. La rgefish caninges t smal
l part
icles butrequ ires
more ene rgyto cap ture an adequa te amoun t resu
ltingin reducedfeed
eff
iciency. Un likefishfeed ,the pa rt
icle size ofcrustacean pe l
letis no t
related tothe mou th size because crustaceans phys ically break down
pellets pr
iortoinges tion. Howeve r
,the pe l
le ts must be of an app rop
riate
size tha t crustaceans can carry and n ibb le on using the irfeed ing
appendages .

Feedapp
lica
tion me
thods

Feed ingf ishisthe mos timportan t da i

ly ac tivity a tfishfa rms requ i r
ing
acons ide rab le amoun
toftime
. The cho ice o ffeed ing me thods w il
l depend
onthefo llow ingfac tors:labo rcos ts, sca le
o ffa rm ope ra t
ion , spec ies be ingfa rmed ,
type o f ho lding sys tem(ponds , cages ,o r
tanks ) , ha tche ry and g row -ou t ope ration .
Manua lfeed inginc ludes broadcas t
ingthe
feedsin tothe pondas we l
lasp lac ingfeeds
in tof loa tingf rames( forf loa tingfeeds )o r
feed ingt rays( fo rs inkingfeeds ). Feed ing
t rays a re used e itherto mon itor ac tua l
feed ingo rtose rveasfeed inga reas(F igu re
7.4 ). Manua lly broadcas tingfeedsinthe
midd le ofa b ig pondrequ iresthe use o fa
sma ll banca . Mechan ica l means o f
par t
ia lly rep lac ing handfeed inginc lude :
au toma t ic feeders wh ich can be se t to
re lease con tro lled amoun ts o ffeed when
ac tivated , and demandfeede rs wh ich can
F
igure 7
.4
re lease a few pe l
lets each t ime a
Useoffeedingtrayto mon
ito
rfeeding
.
t rigger ing mechan ismis bumped bythe
fish (F igure 7 .5). These mechan ica l
feeders can save t ime , however , these
mus tbeca l
ib ra tedandad jus tedregu la r
ly.
Whe ther manua l or mechan ica l, the
feed ing me thod used shou ld be
econom ical and a llow some con trol and
re liabi l ity in d ispens ing feeds . The
me thod shou ld m in im ize feed was te,
prov ide a way to mon itorf ish appe tite
and feed consu mpt ion , and shou ld
distr ibute feeds wide ly in order to
promo te un iformfeed ing and g row th.I f
carr ied ou t care fully, manua l feed ing
can c lose ly mon itorthe appe tite,feed ing
behav ior , and abnorma lities i f any ,o f
F
igure 7
.5
t he a nim al. H ow eve r, it is la bo r
Ademandfeeder. intens ive and t ime consuming .
 CHAPTER7 Managemen
tof Feed
ing Aquacu
lture Spec
ies 175
Mechanizedfeed ing requ i
res h igh cap i
ta l and operating cos ts but can
be o
ffset bythereduc t
ion o flabo r cos
ts associated with manua lfeeding
.
Wi
tht imingcon trols, automa ted systems can be setto deliverfeeds at any
time,a t any givenf requencyin accura te amoun t
, and sp readfeed mo re
evenly and overla rge areas,thanis poss ib
le when broadcas ting by hand.
Manualfeeding maybecomb ined withthe useo f mechanicalfeed ingsys
tems
to e
ffect
ively controlfeedingrou t
ine ,the rebyreducing was te.

Feed
ing
, Wa
ter Qua
li
ty,andtheEnv
ironmen
t

Nutr
itional and econom ic success ofan aquafeedis based upon ava rie
ty
ofinterl
inkedfac tors andtha t no onefactor can be considered byitse
lf.
These are presen ted in F igure 7.6. Feeds tha t are uneaten w i
thin a
cer
ta in per
iod can de teriorate the water quali
ty o fthe cul
ture system .
Hence, properfeed ing st rateg
ies and good wa ter qual
ity managemen t
are veryimpor tantfac to
rs .

F
igure 7
.6
In
ter
linkedfac
torstha
tarec
ri
tica
lfo
rthesuccesso
fanaqua
feed
.
So
urce:T a
con1993

. Feed
A ing
,oxygenrequ
iremen
ts,and wa
terqua
lity

Oxygen ,togetherw ith ano rganicsubs trate,isrequi redfo ralloxidative

metabolic processes. Thera te of oxygen consump tionr isesto a peak
af
terfeed ing, andthen s lowlyfa l
ls backtothe p re-feedinglevel.
Low wa ter exchange ra tes, pho tosyn thes
isin p lants, and the
respiratoryac t
ivityofthepondb iomassc reatef
luctua tionsin dissolved
oxygen(DO )levels. Low DO depressesthefeed ing ac tivi
ty of aqua t
ic
anima ls. When DOleve lislow ,feed ing shou ld no t be done . Giving
lessfeed a t eachfeed ing whileincreas ingthef requency o ffeedings
helps preven tlow DO prob lems . Care ful mon i
tor ing o f DOleve ls,
comb ined w ith observations ofthe behav iorofthe aqua tic an
ima l
,is
176 CHAPTER7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

essen tial and should be rou t

inely done. Oxygen dep let
ion can be
alleviated byincreasing waterexchange(a tleast 30% o
fto ta
lvo lume)
andthe use o f mechanical aera
tors such as a paddlewheel aera
tor
.
Othe r wa ter qual
ity parame ters should be careful
ly mon i
tored so
tha t condit
ions tha t adverse
ly affec
tg row th and survival can be
min imized
. Some wa ter quali
ty problemsin brackishwater ponds and
me thodsto managethem a re g
ivenin Tab le 7
.2.

Tab
le 7
.2 Wa
ter qua
lity parame
ters
, me
thod o
f measuremen
t, wa
ter qua
lity prob
lems and poss
ible causes
, scheme
/ me
thod o
f
managemen
t, andtarge
t wa
ter cond
itionsin brack
ishwa
ter pond cu
lture

Pa
rame
ter Me
thod o
f Wa
terqua
li
ty Poss
iblecauses o
fprob
lems Scheme
/me
thod o
f Ta
rge
t wa
ter
measu
remen
t p
rob
lem managemen
t cond
it
ion

Dissolved DO me te
r, Low DO Increased DO demands dueto : Ae
rat
ioni
f DOis < 4 ppm > 4 ppm
oxygen t
itra
tion over-abundance o fp lankton and
(DO ) benthic organisms, and subsequen t
d
ie-offs o
fp lankton or an
ima l
, Wate
r change 30 -50%,
waste accumu lat
ionin pond bo ttom f
lushing of bo
ttom wastes
Slow O2d iffusionfrom a tmosphe re and det
ritus
Limited pho tosynthes
is du ring overcast
days o
rthe a
lga
l popu
lat
ion co
llapses

DO stra
tif
i­ L
imited l
ight penet
ration due A
lga
l con
tro
l No s
tra
ti
fica
tion
ca
tion to h
igh plankton densi
ty

pH pH mete
r, Low pH Production oforganic acids by Applicat
ion of do
lomi
tic
pH paper anae robic bac
teriafrom or agricul
tura
llime 7
.0 to 8
.5 pH
uneatenfeeds and me tabo l
ic wastes
Acidsleachf rom dikes of ac
id sul
fate so
il Wa
ter exchange
/flush
ing
Excessive CO2p roduct
ion

pH st
rati
fi­ L
im i
tedlight pene
trat
ion due A
lga
l con
tro
l No s
tra
ti
fica
tion
ca
tion to h
igh p
lankton dens
ity

Sa
lin
ity Ref
rac
tomete
r, Sa
lin
ity Ra
in andf
reshwa
terruno
ff Ae
rat
ion/ c
ircu
lat
ion 15to 30 pp
t
hydrome
ter s
trat
if
ica
tion

Tempe­ Thermome
ter Tempe
ratu
re L
im i
ted penetra
tion dueto A
lga
l con
tro
l Notemperature
ratu
re high p
lankton dens
ity s
tra
tif
ica
tion

Unstable In
fluenced by c
lima
tic cond
itions Deep wa
ter
, abou
t 150 cm S
tab
le,26to33
°C
wate
r
tempe ra
ture

Ammon
ia-N Ammonia H
igh NH3
-N Excessfeed, metabol
ic was
tes Wate
r change
/flush
ing <1
.0 ppm
tes
tki
t and decay
ing ma t
ter
; Aera
tion
phytop
lankton d
ie-of
f

N
itr
ite
-N N
itr
itetes
tki
t H
igh NO2
-N Excessfeed, me
tabol
ic was
tes Wa
ter change
/flush
ing <0
.5 ppm
and decay
ing mat
ter Ae
rat
ion

A
lka
lin
ity A
lka l
inity Low Excessfeed, me
tabol
ic was
tes Wa
ter change
/flush
ing > 20 ppm
testkit, alka
lin
ity and decay
ing mat
ter Ae
rat
ion
ti
trat
ion

Hydrogen Sul
fide H
igh H2S Excessfeed, metabo
lic was
te and Wate
r change
/flush
ing <0
.003 ppm
Sul
fide testk
it decaying mat
ter Ae
ration

Transpa­ Secchidisk Tu
rbid Increased suspended sol
ids,erosion Coagulan
ts 30to 45 cm
rency vis
ibi
li
ty ofdikes,turbid wa
tersource
, excessive Wate
r change
and rapid p
lankton b
loom

C
lea
r Absence o
fgood plank
ton b
loom
, Fe
rt
il
izers
:urea
,ch
icken Golden brown,
plank
ton d
ie-o
ff manure l
ightto brownish
g
reencolor

Sou
rce
: Mod
if
iedf
rom Ch
iu 1988
 CHAPTER 7 Managemen
tof Feed
ing Aquacu
lture Spec
ies 177
B
.Fishfa
rm was
tes

Theimpac to faquacu lture onthe env ironmen tis pr ima rilyre latedto
feed managemen t. The compos ition o ffeeds andfeed conve rsion affect
both the phys ica l and chem ica l na ture o f was te ma teria ls andthe
amoun ts produced . Was tes produced dur ingthe cu lture o f aqua tic
anima lsfa llin to two groups : so lid was tes resu ltingf rom unea ten
feed, dus t and feces , and so lub le excre tory produc ts , pr imar ily
ammon ia and u rine,d issolved o rgan ic ma terials, and ca rbon d ioxide .
These ma teria ls are po tentially ha rm fulto bo ththeimmed iatefa rm
environmen t and wa ter bod ies rece ivingthefa rm e ff
luen ts.
Ifo rganic ma terials accumu la teto h ighleve ls onthe bo ttom o fthe
ponds o rinthesed imen t belowf loa tingcages ,thesed imen tw ill become
anaerobic,re leas ingtox ic hyd rogensu lf
ideand me thanegas . Suspended
sol
id ma terials have a ha rmfu le ffect onthe g ills of aqua tic an ima ls,
af
fectresp irato ryfunc tion , and p rov ide po ten t
ials i
tesfo r bac terial and
funga lin fec tion . Long -term e ffec ts may resu ltfrom the re lease o f
phospha tesand n itratesf romf ishfa rme f
fluen tslead ingtoeu troph ication
oren r
ichmen to ftherece iving wa ter. Inex treme cases ,these maylead
tothe occu rrence o f po tentia
lly ha rm fulp lank ton b looms .

Pe
rfo
rmance Measu
res

The success offeed

ing ope ra
tionsisreflectedin overal
l production and
pro
fit
. Growth, surv
ival, andfeed conversion ra
tio
, the ma jo
rind ica
tors
common ly used byfarme rsto measure performance are computed atthe
end ofeach cu l
ture cyc
le(See Chapter6 ).

B
iomass
. Th
is re
fers to the to
tal we
igh
tof spec
ies be
ing cu
ltured
,
expressedinterms o
fag
iven area o
r vo
lume o
fthe hab
ita
t,e
.g.
, pe
r
hectare
.

o
rig
ina
l no o
fstock(p
cs)x ave body w
t(g
)x su
rviva
l(%
)
B
iomass(kg/ha
)=
1,000
Feed conve
rsionra
tio(FCR
). Manyfarmers use FCR astheind
ica
toro
f
the effectiveness of bothfeed ing prac tices and ove ral
l husband ry. FCR
can on ly be accura tely determ inedi fthe amoun to ffeed consumedis
accura tely mon i
tored and recorded . To compare the FCRs o ffeeds,
moisture values mus t be co rrected when ca lculat
ingthe we igh
ts offeed
fed. Poor or var iable FCRs may re flect problems w ithfeed o rfeed ing
methods ,o r may beind icators o f prob lems such asthe occur rence of
diseasein s tocks or worsen ing wa ter qua l
ity
. Low FCR o f1.1 means
thatit onlytakes 1 .1 kgo ffeedto p roduce 1kgo ffish.H igh FCRo f2.5is
considered as poo rs inceittakes 2 .5 kg o ffeedsto p roduce 1kg o ffish.
Thelowe rthe FCR va lue, the be t
te risthefeed u t
il
ization.

amountin d
rywto ffeedg
iven(kg)
Feed conve
rsion ra
tio(FCR
)=
we
tw tga
ined wi
thin afeed
ing pe
riod(kg
)
178 CHAPTER 7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Feed
ing Schemes

A
. M
ilk
fish
Milkfish (Chanos chanos )(Figure 7.7
)is a ma jor pro te
in sourceinthe
Ph i
lipp ines
.I tis omn ivo
rous and g rowsin a w ide range ofsa linity
. The
trad itionallow -dens
ity culturein brackishwa te
r ponds has expandedto
high-dens ity cu l
ture in
freshwa terlakes, brack ishwa ter
ponds , mar ine pens , and
offshore cages . High qua lity
feeds and good feed ing
management become more
critical in these cu l
ture
systems. M ilkf
ish are ha rvested
and ma rketed asfresh , chil
led,
or processedin to value -added
produc ts, such as deboned ,
F
igure 7
.7 ma r
inated,‘cho ice cuts’ packs ,
M
ilk
f
ish Chano
schanos
.
smoked , and canned .

1
. Feed
ing m
ilk
fishb
roods
tockinf
loa
ting cages andtanks
The m i
lkfish broodstockrea r
ingfac i
lit
ies used a t SEAFDEC AQD
arefloat
ing ma rine net cages and concretetanks(F igu re 7
.8A,B ).A
formu la
ted broods tock d
iet(Table 7.3)is broadcasttw ice dai
ly(0800
to 0900 h and 1400 to 1500 h )a t 2to 4% o fb iomass . M ilk
fish
antic
ipatesfeedingt ime and can swallow who le pellets measur ing 1
cm d iameter x 2 cmlong . Mature milkf
ish can spon taneously spawn
duringtheseason w i
th nutri
tional
lyadequa tefeeds and prope rfeeding
managemen t
.

F
igu
re 7
.8
F
loa
t
ing ma
rinene
tcages(A
),andcon
cre
tetank
s(B
),fo
rbrood
sto
ck.
 CHAPTER 7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 179
Tab .3 P
le 7 rac
tica
ldie
tfo
rmu
las(g
/kgd
ryd
iet
)fo
rmi
lkf
isha
tva
riouss
tageso
fcu
ltu
re
Ingred
ien
t Broods
tock La
rva
l F
ryfreshwa
ter F
ryseawa
ter Grow
-ou
t
d
iet3 d
iet4
d
iet1 diet2 d
iet5

F
ish mea
l 200 330
.0 566
.0 300 110
Soybean mea
l 430 180
.0 114
.0 200 308
Squ
id mea
l - 100
.0 - - -
Sh
rimp mea
l(Ace
tes sp
.) - 120
.0 90
.0 - -
Sh
rimp head mea
l - - - 160 -
R
ice b
ran 255 - 87
.0 115 492
Breadf
lour 40 66
.9 - 150 50
Sago pa
lm s
tarch - - 50
.0 - -
K-car
rageenan - 50
.0 - - -
Codl
ive
roi
l 20 80
.0 25
.0 30 20
Soybean o
il - - 25
.0 - 20
Lec
ith
in - 10
.0 - - -
V
itam
in m
ix 15 30
.0 6
.9 10 -
β
-caro
tene - 2
.5 - - -
DL-α-
tocophero
l ace
tate - 0
.1 - - -
M
ine
ral m
ix - 30
.0 36
.0 35 -
D
ica
lcium phospha
te 40 - - - -
Bu
tyla
ted hyd
roxy
toluene - 0
.5 0
.4 - -

Prox
ima
te compos
ition (% DM)
:
Crude p
rote
in 37
.6 46
.3 43
.9* 37
.6 26
.7
Crudefa
t 8
.7 11
.4 10
.3* 8
.7 10
.9
Crude f
iber 3
.9 5
.6 3
.6* 3
.9 8
.4
N
itrogen
-free ex
trac
t 36
.4 27
.3 16
.8* 36
.4 45
.1
Ash 13
.4 9
.4 16
.2* 13
.4 8
.9

Sou
rces
:1 Ma
rte & Bor
longaninFDS Manua
l 1994
;2Bo
rlongane
tal
.2000
;3San
tiagoe
tal
.1983
;4A
lavaandL
im 1988
;5S
umagaysay 1998
*as-
fed basis

2
. Feed
ing m
ilk
fishla
rvaein ha
tche
ries
Inin tensive produc tion, m ilkfish larvae a re rearedin concre te
tanks(3to 10 m3capac i
ty,F igu re 7 .9)a thigh s tockingrates(30 ,000
to 50,000larvae pe r m3 ). M ilk f
ishlarvae a re pe lagicfeede rs, thus
theirfeed shou ld be buoyan t and rema in suspendedin the wa ter
column un ti
l ea ten. Afeed ing schemeis deve loped
wi
ththe use o fa rtif
icia ld ietsto reducethe needfo r
naturalfood produc tionin ha tcher ies(F igure 7 .10).
A nutrit
iona l
ly -ba lanced ,f laked m ic robound d ie to r
MBD (Tab le 7.3) may be fed to m ilkfish larvaein
comb ination with B rach ionus s tart
ing day 2 o r day 7,
and used so lely asfeedf rom day 15 onwa rds. The
naturalfoods are g iven once da ily(0900 h ), wh ile
formu latedlarva ld iets a re d ispensed three t imes a
day at 0800, 1100 , and 1300 h .
For a sem i-in tens ive ha tchery sys tem, m ilk f
ish
larvae are fed in itia lly w ith Brach ionus. At la ter
stages, copepod Acar tia and Pseudod iap tomus
nauplii and MBD are prov ided . MBD is g iven a t FIGURE7.9
2 g/m3 /day, four t imes da ily (0800 , 1100 , 1400 , Intens
ivelar
valrea
ringtan
ks.
and 1700 h ) un ti
l ha rvest.
180 CHAPTER7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

F
igure 7
.10
Feed
ing managemen
tschemefo
rla
rva
lrea
ringo
fmi
lk
fi
sh.
Sou
rce
:B or
longa
neta
l.200
0

3
. Feed
ing m
ilk
fishf
ryin nu
rse
rytanks and ponds
Trad it
iona l
ly, m ilkf
ishf ry obta ined f rom the w i
ld a re rearedin
fe rt
ile nurse ry ponds w ith abundan t na turalfood. Ha tchery-bredf ry
are a lso stocked in to nursery ponds . The na turalfood bases for
milkfishin ponds a re lablab
,p lank ton , andlumu t
. Lab labiscu ltivated
dur ingthe d ry season and p lank ton dur ingthe we t seasonin deep ­
wa ter ponds(70to 100cm ).
Wild-sourcedf ry me tamorphose and has good g rowth and su rvival
inf reshwa ter whenfed on an a rt
ificiald iet(Table 7.3)a t 15to 10% o f
biomassfourt imes da ily(0900, 1200 , 1500 ,and 1800 h )fo r 35 days
intanks .L ikew ise,w i
ld-sourcedf ryrea redin seawa tertanks andfed
artif
ic ialdie t(Tab le 7.3)threet imes da ily(0900 , 1300 , and 1700 h )
at 20 to 15% o fthe body we ight per day a t
tain good grow th and
exce llent surv ivalin one mon th.

4
. Feed
ing m
ilk
fishing
row
-ou
t ponds
Milkf
ishg row-outcu l
tu res tartedfromthet radi
tionalsha l
low wa ter
straight culture of uni
form-s izedfish s tockin ponds to mu l
ti-size
stocking and harves ting us ing modu lar, and deep -wa ter plank ton
methods.
Milkf
ish depends on na tura lfood growninthe ponds dur ingthe
firs
t and second mon ths ofcu l
ture(F igu re 7
.2). Ar
tif
icialfeeds(Tab le
7.3)can befed a t 3to 4% o fbody we igh t per day when na turalfoodis
low. Thereis no commonfeed ing tab lefor supp lemen talfeed ing;
feeding depends onto talf ish biomass , the amoun to f na turalfood ,
and on wa ter qual
ity
. Feed can be b roadcas ta lwaysinthe same a rea
ofthe cage o r pond (Figure 7 .11A, B) . Beforefeeding , thef ish a re
cond i
tionedtosoundto makefeed ingeas ierand mo reeffectivebecause
thef ish gatherin one a rea.
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 1
81

F
igure 7
.11
Feedcanbeb
roadca
stinthesamea
reao
fthecageo
rponds
incef
ishga
the
rinsamea
reafo
rfeed
ing(A
,B)
.

Milkf
ish a re daytimefeede rs. Feedingis donetw ice a day,at m id­
morn ing(when DOis no tless than 3 ppm ) andla te af
ternoon.A t
mid-day , milkfish prefertofeed on na turalfood. Thus ,high plankton
dens ity shou ld be ma inta ined through per iodic app licat
ion o f
fert
i l
izers. Fo r examp le, 25 kg o f 16
-20-0 monoammon ium phospha te
may be app lied pe r hectare every 15 days.
Whenfeedisthe ma jor source of nutr
ients,feed ingisinc reased
toth reet imes a day ; 0800to 0900 , 1200, and 1600to 1700 h . The
feed ra tion cou ld be highe ra t noon andinthe a fternoon(3/4 o fthe
dai
lyra tion)thaninthe mo rn ing(1/4 ofthe da i
lyra tion)
.
Milkfish feed ing act ivity is a f
fected by abrupt changes in
envi ronmen talfac tors such astempera ture, salinity
, and DO . The
dai
lyfeedra t
ionisreduced by 25% whenthetempe rature dropsfrom
32°C to 25 -28 °C, and by 50% when thetempe raturefa llsfrom 26 -
33°Cto 21 -24°C , when sa l
in ityis mo rethan 36 pp t,o r when DOis
lessthan 3 ppm .

B
.Ti
lap
ias
Inthe Ph ilippines,t i
lap ias(T i
lap ia nilot
ica or
Oreoch rom is n i
loticus,T . mossamb ica or O .
mossamb icus,T .au rea o rO .aureus , redt i
lapia
Oreoch rom is spp .(Figu re 7 .12A ,B ,C ,D )rank
second to m ilk
fishin econom icimpo r
tance .
Tilapias are omn ivorous ; they feed on
phytop lankton , zoop lankton , bottom
organ isms , and de tri
tus .T ilap ias areidea l
for cu lture because o fthe irflex ib
lefeed ing
hab i
ts . They can read i ly take a var iety o f
feeds , in mea l form , and in s inking and
floating pe l
letform .T i
lap iasfeedcon t
inuous ly Figure 7
.12
andtheyrespond we lltof requen tfeeding.They T
ilap
ia nilo
tica (A
),mossamb
ica (B
),au
rea (C
),redt
ilap
ia(D
).

prefer sma ll pellets

, and chew ra ther than
swa l
lowthem who le
. Fo rh igh su rv
ival and e f
fic
ientfeed conve rsion,d ry
pelle
tc rumb les are bet ter u t
ilized than the non -pe l
le ted m ixture o fthe
same d iet
.
182 CHAPTER 7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

1
.Broods
tockintanks
, hapas o
r ponds
Sexua lly ma ture ti
lap ias stockedinfer til
ized ponds a re usua l
ly
given asupp lemen tofrice bran o ra 3:1 comb ina t
ion o frice bran and
fish mea l.B roodstockintanks andin hapas a refed comp lete dietsto
obtain h ighf ry product
ion .Ad ietfort ilapia broods tockis shownin
Table 7.4. Th is d
ietisg iven a tafeed ingra te of 1%o fthe body we ight
twice da ily at 0900 h and 1500 h andis su f
ficien ttoinc rease g rowth
andf ry p roduction.
Ti
lap ias spawn asynchronous ly. The fema les
depositthe ireggs a tthe bo ttom o fthe pond ,tank o r
hapa where the ma lesfer ti
lize them . Thefema les
incuba te their eggs and she lterthelarvaeinthe ir
mouths un t
ilthef rya refree-sw imm ing(F igu re 7.13).
Duringth istimethey ea tveryl i
t t
le or none a ta ll
.
Res tricted feedingis su itab le for broods tock .
Ti
lapiatha t have beenfedtosa tiety attempe ratures
28 to 30 °C can be s timu lated to reproduce by
reducingthefeedra tionby 25to 50% .Itisimpo r
tan t
notto ove r
feedthe b roods tock.F requen tc leaning o f
F
igure 7
.13 tanks and hapas d isturbs the brooders , caus ing
Lar
vaere leasedf
rom mou
th-b
rood
ingfema
le. themto sp it outo r swallowthe eggs andla rvae.

2
. La
rvae o
rfryin nu
rse
ry ponds
,tanks
, and hapas
Nursery ponds arefer ti
lized to grow p lankton.T i
lapia f ry are
stockedin ponds o rin hapasins idethe pond. Na turalfoods d if
fe rin
nutritional va lue for t i
lap ia f ry
. The d iatom Nav icu la and
cyanobac terium Ch roococcus are more accep table and are be tter
assim i
lated than the green a lga Ch lo
rella and the phy toflage l
la te
Euglena.
When na turalfood startsto dec l
inein ponds ,the nursery ope rato r
introducesfeed wh ich may bef ine r
ice bran ora 7 :3 comb ina t
ion o f
fine rice bran andf ish mea l
. In nursery tanks and hapas where
naturalfoodis sca rce, aformu la
ted diet(Table 7.4
)is giventof ry as
soon asthey s tartfeeding.F ry arefedto sa t
iety
,o ra t abou t 30% o f
the biomass da i
ly,fou rtimes a day . Astheyg row,thefeed ingra teis
gradua l
ly decreasedto 15% . Thefeedis broadcas t overa reasinthe
nursery sotha tthef ry do no t haveto swimfa rto searchforfood .

3
.Ti
lap
iasin g
row
-ou
ttanks
, ponds
, andcages
Polycul
turesys tems oftilapias wi
thf ish ofd i
fferentfood prefe
rence
are usedto maxim izethe u ti
lizat
ion of natu ralfoods. The manures o f
chicken, ducks,o rp igs are used tofer t
ilize the ponds . Loca l
ly
avai
lable crude andinexpens ivefeedstuffs such asr ice bran, po
lla
rd,
and cop ra mea
l,le f
toverb read and b iscuits, and comme rcia
l poul
try
feeds are given as supp lemen talfeeds tw ice a day at 2to 3% o f
biomass.
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 183
Tab
le7
.4 P
rac
tica
ldie
tfo
rmu
las(g
/kgd
ryd
iet
)fo
rti
lap
iaa
tva
riouss
tageso
fcu
ltu
re
Ing
red
ien
t B
roods
tockd
iet1 La
rvae
/f
ryd
iet2 G
row
-ou
tdie
t3
F
ish mea
l 362 301
.7 182
.5
Co
rn g
luten mea
l 204 - -
Soybean mea
l 177 259
.5 250
.0
Cop
ra mea
l 118 114
.8 100
.0
Ip
il-
ipi
l mea
l - 81
.0 -
Cassavaf
lour - - 364
.2
R
ice b
ran 75 149
.7 60
.0
S
tarch 32 30
.0 -
Codl
ive
roi
l 5 10
.0 -
Vege
tab
le o
il 5 10
.0 -
V
itam
in-m
inera
l m
ix 22 43
.3 43
.3

Prox
ima
te compos
ition (% DM)
:
Crude p
rote
in 44
.0 38
.1 28
.1
Crudefa
t 5
.5 8
.7 3
.8
Crude f
ibe
r 9
.1 5
.6 3
.6
N
itrogen
-free ex
trac
t 29
.6 30
.8 54
.6
Ash 11
.8 16
.8 9
.9

Sou
rces
:1,
2,3San
tiago e
tal
.1985
,1986
,1987
,respec
tive
ly

Broadcas ting by us ing acup o ralad leisthe mos tcommonfeed ing

method . Thefeedis d is tribu ted even ly o rin seve rals ites sotha ta ll
fish have easy accesstofeed . Wa tertempe rature,fish s i ze
,the qua l
ity
of the feed , and the presence o f na tura l food a ffect the feed
consump t
ion o ftilap ias. Thus , the da ilyfeed ing ra tionis ad justed,
depend ing onf ish appe tite.
Intensive cu lture o ft ilapia has ga ined popu lar
i tyin recen t years.
This me thod o ffarm ing requ ires the use o fh igh qua lityfeeds and
modern managemen t too ls.F ish are s tocked a th igh dens itiesin
conc retetanks , ea r
then ponds o rfloa ting ne t
cages (50to 74f inge rlings pe r m3). Fo rg row -
ou tcagesinlakes(F igu re 7 .14),feed ingt rays
or a secondf ine-mesh ne ta tthe bo ttom o f
the cage a reins talled to he lp m inim ize the
loss o ffeeds .Ad ietw ith 28% pro tein(Tab le
7.4)is giventhreet imes da ily a t 5to 6% o f
biomass . Da ilyfeed ingra te can be dec reased
to abou t 2to 3% asthef ish s izeinc reases
from 30gto mo rethan 100g .The da ilyra tion
is giveninthreefeed ings 3to 4 h apa r
t,a t
0900 , 1200, and 1600 h . Dur ingcoo l mon ths,
when me tabo l
icra teislow ,feed ingf requency
can be reduced totw ice a day a t 0900 and
1500 h . The appe titeis gene ra l
ly poo r when Figure 7
.14
watertempe ratureislow , and h ighes tinthe Grow-outfloa
ting ne
tcagesin alake
.

afternoon , whenthe wa teris wa rme r.

184 CHAPTER 7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

C
. Rabb
itf
ish
Rabbi
tfishes are widely d is
tributedintheIndo-Pacif
ic region.
The loca l species S iganus gu tta
tus (Figure 7.15 ) is an
econom ical
lyimpor tan tfoodf ishinthe Phil
ippines
.I t breeds
in captivi
ty, spawns regu larly
, and growsin brack ishwa ter
pondsfeed ing on
ly on a lgae.

1
. B
roods
tockintanks
Rabbit
fish spawns na turally every mon th the whole year
round. Broodstock arefed with an a r
tif
icialdiet(Table 7
.5)at
F
igure 7
.15 3% o fthe body weight per day, morn ing and afte
rnoon. This
Rabb
it
fish S
iganus gu
t tus ju
ta ven
iles
. die
t enhances reproduc t
ive performance , and egg andlarval
quali
ties o
frabb it
fish.

Tab
le7
.5 P
rac
ticaldie
tfo
rmulas(g
/kgd
ryd
iet
)fo
rrabb
it
fish 2
. La
rva
lrea
ring and nu
rse
ry
b
roodstockandfry Inthe ha tche ry, rabb i
f
tishlarvae
Ingred
ien
ts Broods
tock diet1
Nu
rse
ryd
iet2 (day 1to 21 ) can grow and surv ive
well w ith feed ing e ither h ighly
F
ish mea
l 100
.0 145
.0
unsaturated fat ty ac ids (HUFA )-
Sh
rimp mea
l(Ace
tes sp
.) - 145
.0
Squ
id mea
l
enr iched rot ifer a t 15 -20 ind/m l
,
100
.0 134
.0
Sh
rimp mea
l 100
.0 - HUFA -enr iched ro t
ifer supp lemen ted
Soybean mea
l 80
.0 - with a r
tificiald iet(0 .5 g/m3 /day),o r
Co
rn g
luten mea
l 80
.0 - Ch lore l
la -fed ro tifers supp lemen ted
Whea
t ge
rm 80
.0 - with ar tificiald iet. Good g row th and
Brewe
r’s yeas
t 60
.0 - surv iva l are a lso obta ined when
Ce
llu
lose 187
.3 28
.5 larvae (day 25 to 45 ) are fed an
Breadf
lour 22
.5 376
.0 artif
ic iald iet(Tab le 7.5)
. Feed ingra te
K-car
rageenan - 50
.0
is a t 23% o fin i t
ialb iomass da ily,
Codl
ive
roi
l 100
.0 71
.0
givena t 0900 , 1030 ,1200 ,1300 , 1500,
Soybean o
il 15
.0 -
Soybean lec
ith
in 45
.0 -
and 1700 h . Afeed ing schemef rom
V
itam
in m
ix 20
.0 30
.0
day 1to 45 ou t
linedin F igure 7.16
M
ine
ral m
ix 10
.0 20
.0 yields cons is
ten tlyfa ir surv ival
.
Bu
tyla
ted hyd
roxy
toluene 0
.2 0
.5
3
. Cu
lturein ponds andf
loa
ting
Prox
ima
te compos
ition (% DM)
: ne
tcages
Crude p
rote
in 38
.4 39
.2 The na tura
lfood recommendedin
Crudefa
t 17
.4 9
.9 the rear ing pond are f i
lamentous
Crude f
iber 13
.1 5.5
algae such as C ladophora l inum ,
N
itrogen
-free ex
trac
t 23.6 37.8
Chae tomo rpha spp.
,andEn te romo rpha
Ash 7.5 7.6
tubu losa. These a lgae are p lan ted
Sources:1
Duray e
tal
.1994
;2Pa
razo 1991
.
un i
form ly in sma l
lc lumps in the
enti re pond , done dur ing the la te
af
ternoonto preven tthemf romf loatingthe nex t day. Rabb i
tfishes
rearedinf loa
ting cages andin ponds a re a
lsofed supp lemen tary
feeds or commercialti
lap
iafeeds.
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 185

F
igure 7
.16
Feed
ingschemefo
rrabb
i
tf
ishla
rvae
.
Sou
rce
: Ha
raeta
l.198
6

D
.Bighead Ca
rp
Carps are amongthe mos tcomme rc
ial
ly desirablef reshwaterfishspecies
in Asia and theIndo -Paci
fic reg
ion. Bighead ca rp(Ar
ist
ichthys nobilis)
(Figure 7.17A) has a high reproduct
ive capac ity andfas t growth ra te
,
feedslow onthefood cha in, and can be grownin monocu l
turein ponds ,
cages andf ish pens o
rin po lycu
lture wi
th m ilkfish andt i
lapiain pens .1

1
.Broods
tockinf
ish pens and cages
Bighead carp ma turein cagesinthelake w ithout supplemen tal
feeding, butfry produc tion and surviva
lareenhanced whenthe b rood-
stock arefed supp lemen taldie
ts. Broodstock d
ie t(Tab
le 7.6
)isfed a t
2to 0 .8% of body we igh t pe
r day at 0930 and 1500 h. Ar
tif
icia
ld iets
posit
ivelyin f
luencethe onse to f gonad maturation and enhancethe
hatchab i
lity of eggs.

F
igure 7
.17
B
igheadca
rp A
ris
tich
thysnob
i
lis (A
),indu
cedtospawnu
singho
rmona
lin
jec
t
ion(B
),andindoo
rla
rva
lrea
ringfa
ci
li
tie
s(C
).
186 CHAPTER7 NUTR
ITIONIN TROP
ICAL AQUACULTURE
Tab
le7
.6 Prac
tica
ldie
tformu
las(g
/kg d
ryd
iet
)fo
rbigheadca
rpa
tva
rious
s
tages ofcu
ltu
re

Ingred
ien
ts Broods
tock La
rva
l La
rva
l
d
iet 1 d
iet A2 d
iet B3
Cop
ra mea
l 200
.0 - -
F
ish mea
l 150
.0 406
.0 566
.0
Sh
rimp mea
l(Ace
tes sp
.) - - 90
.0
R
ice b
ran 25
.0 150
.0 127
.0
Soybean mea
l 320
.0 361
.3 114
.0
Co
rn g
luten mea
l - - -
Mea
t and bone mea
l 224
.0 - -
L-
lys
ine 1
.0 - -
DL-me
thion
ine 5
.0 - -
S
tarch 26
.9 - 10
.0
Codl
ive
roi
l 5
.0 32
.7 25
.0
Co
rn o
il 5
.0 - 25
.0
V
itam
in-m
ine
ralp
rem
ix 38
.1 50
.0 43
.0

Prox
ima
te compos
ition (% as
-fed
):
Crude p
rote
in 41
.5 40
.2 41
.5
Crudefa
t 5
.1 6
.8 11
.9
Crude f
ibe
r 3
.9 no da
ta no da
ta
N
itrogen
-free ex
trac
t 30
.4 26
.5 24
.5
Ash 14
.6 no da
ta 12
.5

Sou
rces:1San
tiago and Gonza
l 2000
;2Fe
rmin and Recome
ta 1988
; 3San
tiago and Reyes 1989

2
. La
rva
l and nu
rse
ryrea
ring
Induced spawn ing ofthe b roods tock us ing hormones(F igure 7 .17B )
produces b ighead carplarvae . In the nursery(F igure 7 .17C ), the
comb inat
ion o fl
ivefoodo rgan ism and a rt
ific
iald ie
tenhancestheg row th
ofb igheadca rplarvae . Thecomb ina tiono fMoina andla rvald ietA(Tab le
7.6)fed dailytosa tiationa t 0900 , 1100 , 1300, and 1600 hg ivesthe bes t
grow th o fb ighead carplarvae (day 5to 12 weeks ). Sim ilarly, the
comb inat
ion o fB rach ionus andla rva ld iet B(Tab le 7.6) has a pos it
ive
effect on grow th. B ighead ca rpla rvae can g row well on a rtif
iciald ie t
givenin it
ially at 100%o fthe body we igh t per day, andfinally a t 30% o f
thef ish biomass da ily. Thesela rvae a rerea redinindoo r nu rserytanks
fo r 30-45 days be forethey a re harves tedfo rs tockinging row -ou tcages .

3
. G
row
-ou
t cu
ltu
re
Culturedinf ish pens
,b igheadca rps do no
t haveto befed ar
tif
icial
feeds,sincetheyth r
ive wel
l on abundan t naturalfoodin Laguna de
Bay. Ca rpfa rm ing has a
lready expandedto up land andland locked
areasinthe Ph i
lippines.Bigheadca rps arerearedtogetherwithtilapia
inferti
lized ponds whe retheyre ly mainly on natura
lfood organisms,
wi
th occas ional supplemen talfeeds
.

E
. Na
tive Ca
tfish
The native catf
ish Clar
ias mac rocepha lus (Figure 7
.18A)is animpo rtan
t
indigenousf reshwaterfoodf ishin Ma lays
ia,Tha i
land, andthe Phi
lippines
.
At SEAFDEC AQD , catfishisinducedto breedin cap tiv
ityin orderto
arrest the decline ofthe na tural popu lation, and ensure a suff
icient
supply offry andf ingerlingsfors tocking.
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 187

F
igure 7.18
Na
t
iveca
tf
ish C
lar
ias mac
rocepha
lus (A
),rea
redinha
tche
ry(B
),andinne
t-cages inpond(C
).

1
.Broods
tock
Traditionally
, rawf ish alone orin comb inat
ion with art
ific
iald iet
is used asfeed to cond i
tion the broods tockforinduced spawn ing.
Livefoodsl ike Tubifex worms a re also used to supplemen tthe d ry
broods tock d iet
. W ithout supp lemen tat
ion o f natura
lfood , two
arti
ficia
ld ietformu lations enhancethe reproduc t
ive per
formance o f
catf
ish and can be usedin p roduc tion and maintenance o
fb roods tock
(Tab le 7
.7). Broods tock arefedtwot imes a dayto sa t
iat
ion o r abou t
8to 10% o fthef ish biomass daily.

Tab
le7
.7 P
rac
tica
ldie
tfo
rmu
las(g
/kgd
ryd
iet
)fo
r As
ianca
tf
isha
tva
riouss
tageso
fcu
ltu
re
Ingred
ien
t Broods
tock Broods
tock Larva
l Grow-out
d
iet A1 die
t B1 diet2 diet3

F
ish mea
l 250
.0 150
.0 73
.8 200
Soybean mea
l 350
.0 350
.0 380
.0 300
Sh
rimp mea
l(Ace
tes sp
.) - - 77
.5 -
Mea
t and bone mea
l 53
.3 224
.0 - -
Squ
id mea
l - - 71
.3 -
R
ice b
ran - 156
.1 310
Cop
ra mea
l 261
.2 34
.4 -
Ip
il-
ipi
llea
f mea
l - 35
.5 -
Breadf
lour - - 191
.4 90
S
tarch 20
.0 - -
κ-car
rageenan - - 10
.5 -
Ce
llu
lose - - 14
.7 -
Codl
ive
roi
l - - 80
.8 -
Soybean o
il 30
.0 20
.0 50
Vi
tam
in m
ix 35
.5 30
.0 50
.0 -
V
itam
in-m
inera
l m
ix - - 10
Mine
ral m
ix - - 10
Dica
lcium phospha
te - - 50
.0 30

Prox
ima
te compos
ition (% DM)
:
Crude p
rote
in 42
.5 43
.1 39
.9 34
.2
Crudefa
t 7
.2 7
.9 10
.6 9
.5
Crude f
ibe
r 4
.2 3
.4 1
.5 5
.8
NFE 33
.4 30
.2 40
.2 36
.3
Ash 12
.7 15
.4 7
.5 14
.2

Sou
rces
:1San
tiago and Gonza
l 1997
;2Fe
rmin and Bo
liva
r1996
;3Con
iza e
tal
.2001
188 CHAPTER7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

2
.La
rva
landnu
rse
ryrea
ring
Livezooplank tonfeed(A r
tem ia o rMoina) isrequ iredfo r successful
larva lrear
ing(F igu re 7.19B )ofca t
fish. Dur ingthef irs t weeko frearing,
new ly hatched A rtem ia a refedtof ish a t 10ind ividua ls per day and
arti
ficialdieta t 50% o f body we ight.A tthe second week o f cul
tu re
,
Artem ia is g
iven a t 20ind ividua ls pe r day and a rt
ificiald ietat 25% o f
body we ight per day . The da i ly rationis d iv
idedin tofourfeed ings
given be tween 0900 and 1500 h . Wean ing o f catfishlarvae to dry
arti
ficialdiet can be s tartedfour days a fterfeed ingl ive zooplankton.
Acomb ined dieto fA r
tem ia andad rya rti
ficialfeed(Tab le 7.7)improves
the g rowth and surv ivalo f catfishla rvae .
Hatchery-bred ca tfishf ry stockedin ne t cagesins tal
ledintanks
andin ponds(F igure 7 .18C ) grow we ll on ar tif
icia ld iet. In ponds,
natura lfood(copepods and c ladoce rans)a lso serves asfood source
for ca t
fishjuven iles.

3
.Grow
-ou
tcu
ltu
re
Traditionally, cat
fishfeeds on decaying organic ma t
terinthe pond
with kitchenre fuse assupp lemen t. Feedingca tf
ish b lanched chicken
en t
ra i
ls andr ice bran(80:20 )at 10% ofthef ish body we ight per day
is also p ract
ised.
Culturedin pensins ide a pond(Figure 7.18C), catf
ishfry arefed an
arti
ficiald iet(Table 7.7)at5 ,4.5,4 , and 3.5% off ish body weight per
dayfo rthe 1 s
t, 2nd
,3 rdand 4th month ofculture
,respec t
ive
ly. Feeds are
given a t 0800 and 1600 h w iththec rumble dietforthef irs
ttwo mon ths
and abou t2 .5 mm d iameter pe
lletsthereaf
ter until harvest
.

F
.As
ianseabass

Sea bass (Lates calcar

ife
r) (Figure 7.19)is an
impor tant aquacu lture speciesin Tha iland,
Hong Kong,S ingapo re, and Ma laysia
. Ca l
led
“barramund i”in Aus tralia and Papua New
Guinea, sea bassis animpor tant resource,
forming the bas is ofthe ir commerc ial and
F
igure 7
.19 recreational f isheries. Sea bass is an
As
ianseaba
ss La
tesca
lca
ri
fer
. opportunis
tic predatorthroughou titsli
fecycle.

1
. B
roods
tockinseacages andtanks

Sea bassrea
redto matur
ityinfloa
ting netcages andtanks spawn
natura
lly. Sea bass a
refed rawfish at 3to 5% body weight once
dai
ly and reduced to 1to 2% body weight dur
ingthe peak o fthe
spawning season.

2
. La
rva
l and nu
rse
ryrea
ring

The phy top

lankton Ch
lore
lla, Te
traselmis orIsoch
rys
is a
re addedto
larval rearingtanks as water condit
ioners and asfoodfor roti
fers
.
Sea bass begintofeed 50 h af
ter hatching bu
tfeedis g
iven ear
lierat
36 h. Larvae a
re weaned g
radua l
lytoeach newfeedtype byincreasing
 CHAPTER 7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 189

F
igure 7.20

Feed
ing managemen t schemeforthela
rva
lrea
ring o
f sea bass
.
Sou
rce
:Parazoe
tal
.1998

the da ily propor tion o fthe newfeedtype wh ile gradua lly dec reasing
tha to fthe preceed ingfeed . (Figu re 7 .20)
As a par tial or comp le
te rep lacemen to fA rtem ia, thef reshwa ter
cladoce ran Mo ina may befedtosea bassla rvae s ta rt
ing w i
th 17to 20
day o ldla rvae a t afeed ingra te o f notlessthan 1ind/m l
. Mo ina a re
firsts ievedto ob tain sma ll adu lts and neona tes be forefeed ingto 17 -
dayo ldla rvae . Uns ieved Moina can befed d i rectlyto 20 -dayo ldlarvae.
Whenfeed ing w ith Mo ina,the sa l
in ity ofrear ing wa teris gradua l
ly
lowe redto abou t 10 pp twith in a 24 -h period . La rvae a refed a tleas t
fourt imes da i
ly. The brack ish wa ter cladoce ran ,D iaphanosoma , is
also a po ten tiallivefoodfo r sea bassla rvae.I t can befed once a day
to 15 -day o ld and o lde rlarvae a t 2ind/m la t 32to 35 pp t.
Sea basslarvae andjuven i
les can be rearedin nursery cages
insta l
ledin open wa ters using n igh t
-
ligh t to a ttract zoop lankton (mos tly Table7.8 P ract
icaldietformu
la(g/kgdryd ie
t)
copepods ) asfood . As tock
ing dens ity forjuvenileseabass
of600/m2can be used whenf ishfeed Ingred ients i Amount1I
onzoop lank ton a lone bu t mincedf ish Fish meal 420.0
flesh g ivenadl ibitum dur ing day time Soybean mea l 90.0
is needed when s tock ing dens ityis Shrimp mea l (Acetes sp
.) 100.0
inc reased to 1200/m2 . Squid mea l 50.0
B
readf
lou
r 77
.5

3
.Grow
-ou
t cu
ltu
re Ce
llu
fil 145
.0
Codl
ive
roi
l/ soybean o
il(1
:1) 57
.5
Sea bassjuven i
les respond we l
l
V
itam
in m
ix 40
.0
to dry arti
ficia
ld iet aslong asthey
M
ine
ral m
ix 20
.0
aretra ined tofeed an artif
iciald iet
are earlyin the nursery . In tanks, Prox
ima
te compos
ition (% DM)
:
Crude p
rote
in 43
.2
juvenilesfed an ar t
ific
iald ie
t(Tab le
Crudefa
t 9
.3
7.8
)a t 17to 14% , 12to 8%, and 8to
5% ofthe body we ightforthef irs
t 20 Sou :'C
rce a
tacu
tanand Co
loso1995
.
190 CHAPTER 7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

days, next 20 days

, andlas t 14 days, respective
ly. Da
ily ra
tionis
giveninthree equa l port
ions at 0930
, 1300 , and 1600 h.In ponds
and in sea cages , sea bass grow we ll on raw f ish alone orin
comb inat
ion with art
ific
ialfeeds.

G
.Orange
-spo
ttedg
roupe
r
Groupers a
re amongthe mos tva luable spec
iesfo r expor
t ma rke
ts. Thei
r
demandis mos t
lyin countries with high seafood consump tion orw i
th
high econom ic grow
th. Groupe rs spawn read ilyin cap t
ivi
ty bu tlarval
surv
ivalis stil
llow.A tpresent,the grow -outindust ryrel
ies onfrytaken
fromthe w ild.

1
.Broods
tockinseacagesandtank
s

Orange -spo
tted grouper(Ep inephelus co
ioides)(Figure 7.21)first
mature asfema les at around 3to 4 kg body we ight, some o fthe
fastest growingfema les changein to males when they reach more
than 6 kg. Broodstock heldintanks o rcages spawn mon th
ly, usually
wi
thin a week before orafterthelast quar
ter moon phase
. Broods tock
maintainedin sea cages andintanks a refed a var
iety ofspecies of
rawf ish every othe
r mo rning at sa
tiat
ionlevel.

F
igure 7
.21
O
range
-spo
tted g
roupe
r Ep
inephe
lusco
io
ide
s.

2
. La
rva
lrea
ring
Inin tensivelarva l rearing, the rearing pro tocolinvo lvesfeeding
younglarvae (day 2to day 15 ) w i
th screened ro tifers previously
enriched w ith high n-3 HUFA boos ters.O lderlarvae(day 20to 50 ),are
feden richedA r
tem ia naup l
iiorArtemia me ta -naupli
i unt
il metamo rphosis
tothejuven ile phase(F igure 7.22)
.
In sem i-intensivelarva l rearing, grouperlarvae s tocked atlow
densit
ies(10la rvae/l
)w i
th m inima l water exchangefeed on copepod
naup l
ii duringthe ea rlyrear ing phase. Copepods a re collectedfrom
brackish wa ter ponds andinocu latedinthela rvaltanks 2to 3 days
before stocking o flarvae.F irst-
feedingla rvae prefer copepod naup l
ii
than ro tife
r.
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 1
91

F
igure 7
.22
Feeding and wa
ter managemen
tschemefo
rin
tens
iverea
ring o
fgroupe
rla
rvae
.
Sou
rce: Durayetal
.1997

3
. G
row
-ou
t cu
ltu
re
Gene rally,rawf ishis used asfeed Tab
le7
.9 P
ract
ica
ldietfo
rmu
las(g
/kgd
ryd
iet
)fo
rgrow
-ou
tcu
ltu
re
for groupers farmed in sea cages , o
fgrouper
ponds , and net -cages in ponds . Ingred
ien
ts Grow-ou
tdie
t Grow-ou
tdie
t
Ano therfeed ings trategyistoin it
ia lly
Ch
ilean f
ish mea
l 80 200
stock adu ltt i
lap ias(5 ,000to 10 ,000 Mea
t and bone mea
l,loca
l - 200
per hec tare )in ponds and a llowthem Mea
t and bone mea
l,impo
rted 320 -
to reproduce .T i
lap iaf inger l
ings a re B
lood mea
l 80 80
Sh
rimp mea
l 100 100
then used as food for grouper
Soybean mea
l 60 60
juven iles. In add i t
ion , chopped raw
Squ
id mea
l 10 10
fishis g iven every o ther day a t 5% Whea
tflou
r 150 150
ofgroupe rb iomass . Ha l
fo fthera t
ion R
ice b
ran 70 70
is g iveninthe morn ing and ha l
fin DHA-Se
lco
Codl
ive
roi
l 60 60
the a fternoon , some are p laced in
V
itam
in m
ix 40 40
feed ing trays for mon itoring
M
ine
ral m
ix 30 30
purposes and the res t are V
itam
in-m
ine
ral m
ix - -
broadcas ted . I frawf ishisthe so le Bu
tyla
ted hyd
roxy
toluene - -
feed , groupers may be g iven up to Prox
ima
te compos
ition (% DM
):
10% o fb iomass da ily. Crude p
rote
in 44
.0 44
.0
Ha tchery -bred grouperjuven iles Crudefa
t 11
.5 11
.5
accep t dry ar ti
ficial pe l
lets aslong Crudef
ibe
r 1
.8 1
.8
as they are t rained tofeed on the NFE 25
.8 25
.8
pellets ea rlyinthe nu rsery. Art
ificia l Ash 16
.9 16
.9

feed(Tab le 7 .9)is g ivento g roupe rs Sou

rces
: M
il
lamenae
tal
. 2001
cul tured in o ffshore cages and in
cagesins talledins idethe pond a t4
to 6% o fb iomass da ily.
192 CHAPTER7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

H
. Mang
roveredsnappe
r
Mang rovered snapper Lutjanus a
rgentimacu
latus (Figure 7
.23)is anim­
portantfoodf ishin Southeast As
ia andis popularly cul
turedin brack­
ish wa ter ponds and mar
ine net cages
. Snappers are opportun
ist
ic car
­
nivores andfeed ma in
ly onfish and crus
taceans.

F
igure 7
. 23
Mang
roveredsnappe
r Lu
t
janu
sargen
t
imacu
latu
s.

1
.Breed
ing
Wild
-caugh tor hatchery p
roducedf ry are sexua
lly mature a
fter5
(ma les) and 6 years (females) in f
loa t
ing ne t-cages orin tanks .
Broodstock arefedrawf ishevery othe
r mo rning at 5% o
fbody we igh
t.
They are sexua lly mature f
rom Apr il to Oc tober and they spawn
natural
lyfo r uptofour consecutive days.

2
. La
rva
lrea
ring
Thefeed ing reg imefo r
the red snapper larvae
cons is ts o f Ch lorella,
rotifers , Ar temia, and
minced f ish f lesh (F igure
7.24). Grow th and su rv
ival
ofred snapperlarvae a re
bes t when fed screened
ro t
ifers (<90 µm ) dur ing
the f irst 14 days and
Artem ia naup l
ii(2 pe r ml
)
given four t imes a day
starting on day 22 . Day 21
to 35larvae have s im i
lar
surv iva l w ith Ar temia
alone, ar t
ificiald ieta lone,
orcomb inationA r
tem ia and
artif
icia ldie t
.

F
igur
e7.2
4
Feed
ingand wa
ter managemen
tdu
ringla
rva
lrea
ringo
fthe mang
roveredsnappe
r.
Sou
rce
: Dura
yeta
l.1994
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 193
3
.Grow
-ou
tcu
ltu
re Tab
le7
.10 P
rac
tica
ldietfo
rmula(g
/kg
Snappe
rs are usua
llyfed rawf ishinf loat
ing ne t
-cages d
rydiet
)forredsnappe
r
butthey a
lso respond we
llto a rt
ific
iald iet.Intes ts done Ingred
ien
ts Amoun
t1
in 250ltanks thefeed(Tab le 7.10)g iventw ice a day a
t
Pe
ruv
ianf
ish mea
l 430
.0
0930 and 1600 h at3.5to 4 .5% o f body we ight per day Squ
id mea
l 80
.0
enhances growth and su
rvival. Sh
rimp mea
l (Ace
tes sp
.) 54
.0
De
fat
ted soybean mea
l 80
.0
I
. T
ige
rsh
rimp B
readf
lou
r 200
.0
The tiger shr
imp Penaeus monodon (F igure 7 .25A ,B)isthe R
ice b
ran 58
.0
mostw idely cultured shr imp spec ies and accoun tsfor over Codl
ive
roi
l 38
.0

oneth ird ofgloba l

lyfa rmed sh r
imp . H
igh -density product
ion Soybean lec
ith
in 5
.0
V
itam
in and M
ine
ral m
ix 50
.0
is suppor ted by greater use o fformu lated feeds , where
Asco
rby
l monophospha
te 0
.2
produc t
ion may y ield up to 10tons/ha/year .T iger shrimp
D
ica
lphos 5
.0
culture has evo lved f rom trad i
t ional ex tensive to sem i-
intensive andin tens ive sys
tems us ing mode rn equipmen t and Prox
ima
te compos
ition (% DM)
:
Crude p
rote
in 44
.0
techniques. The a imisto p roduce s igni
fican t
ly high yie
ld per
Crudefa
t 10
.0
unit area wh i
le m in imizing farm was te d ischarges to the
Carbohydra
te 26
.5
environmen t.
Ene
rgy(MJ
/kg d
iet
) 18
.9

S
our
ce: 1Ca
tacu
tane
tal
.2001

F
igure 7
. 25
T
ige
rsh
rimp Penaeus monodon (A
),ands
tageso
fova
rian ma
tur
i
ty(B
).

1
.Broods
tockintank
s
Shrimp broods tock a refed on a var ie ty ofl ive,f resh ,o rf resh -
frozen na tura lfood , such as mar ine anne l
ids, musse l mea t, squ id,
clam mea t, and o the r meato f mo llusks and can be g iven a lterna tely.
These na tura lfoodi tems con tain essen tial nu trients pa rt
icu larly n -3
HUFAtha ta re neededfo r shrimp ma tura tion(F igure 7 .25B ). B rown
musse l mea t and squ id are choppedin to sma llp ieces. Ma rine wo rms
areg iventosh rimps preferably alive. Fo rconven ience ,theya re bough t
in bu lk, we ighed , packed,f rozen, andthawed as neededfo rfeed ing.
An a rt
ific
iald iet has beenfo rmu la ted us ingthefa tty ac id and am ino
acid pro fi
les o f mature shr imp ovar ies as gu ide . The d iet is
supp lemen ted withf resh wetfeedsin an a ttemp ttoensu re an op tima l
nutrient ba lance and afu llrange o fessen tial nu tr
ients . Rep roduc t
ive
performance o f pond-reared sh r
imp broods tockis ma rked lyimp roved
whenfed a comb ination dieto f natu ralfood and pe llets.A m ix ture o f
variousfeeds tu f
fsis be t
te rthan a s inglei teminthe d iet. Mos to fthe
fema les given na turalfood a lone unde rgo ova rian reg ression and do
not spawn .
194 CHAPTER7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

The standa rdfeed ing reg

imefo r shrimp broods tock at SEAFDEC
AQD cons is
ts o ffreshfood and pe l
lets(Tab le 7.11
)g iven alternately.
Fresh orfrozen na turalfoods(squ id, musse l meat,or ma rine wo rms )
are g
iven daily at 0800to 0900 h a t 5to 10% o fbiomass, and pe llets
at 1600to 1700 ha t2to 3%o fb
iomass . Naturalfoodsa re broadcas ted
intothe ma tura t
iontank wh ile pellets a
re placedinfeed ingt rays .

Tab
le7
.11 P
rac
tica
ldie
tfo
rmu
las(g
/kgd
ryd
iet
)fo
rtige
rsh
rimpa
tva
riouss
tageso
fcu
ltu
re
Broods
tock La
rva
l Grow-ou
t
Ingred
ien
t
diet1 diet2 diet3

Squ
id mea
l 300
.0 300
.0 –
Sh
rimp mea
l - - 150
.0
Sh
rimp head mea
l 200
.0 350
.0 -
F
ish mea
l 200
.0 - 250
.0
Soybean mea
l - - 250
.0
B
readf
lou
r 55
.0 110
.0 130
.0
R
ice b
ran 22
.5 - 69
.5
Seaweed (Grac
ilar
ia) 40
.0 - 50
.0
κ-car
rageenan - 5
.0 -
Ce
luf
il - 22
.0 -
Codl
ive
roi
l 60
.0 80
.0 25
.0
Soybean o
il - - 25
.0
Soybean lec
ith
in 30
.0 25
.0 -
Cho
les
tero
l 5
.0 10
.0 -
V
itam
in m
ix 27
.0 60
.0 20
.0
β
-caro
tene - 2
.5 -
M
ine
ral m
ix 60
.0 40
.0 10
.0
D
ica
lcium phospha
te - - 20
.0
Bu
tyla
ted hyd
roxy
toluene / E
thoxyqu
in 0
.5 0
.5 0
.5

Prox
ima
te compos
ition (% DM)
:
Crude p
rote
in 52
.8 50
.3 41
.7
Crudefa
t 12
.1 14
.2 8
.8
Crude f
ibe
r 3
.8 20
.4 5
.9
N
itrogen
-free ex
trac
t 13
.4 10
.1 29
.2
Ash 17
.9 5
.0 14
.4

Sou
rces:1 M
il
lamenae
tal
.1986
;2Bau
tis
tae
tal
.1989
;3M
il
lamena and T
riño 1994

2
. La
rvaein ha
tche
ryand nurserytanks
In the ha tcheries , the food o f shr imp pro tozoea llarvae up to
postlarvae PL5cons ists mos tly ofplank tonic microa lgae (Ske letonema ,
Tetrase lm is, Chae toce ros) and Ar tem ia naup li
i. At PL6onwards ,
postlarvae a re gradua llyintroducedto m inced musse l mea t, rawf ish,
and sh rimp mea l Ace tes sp.o r pel
letc rumb les.
Ar
ti f
ic iald iets tha t part
ially replacel ivefood he lp a l
leviate the
needfo rla rgetanks necessa ryfo r phy toplank ton p roduc t
ion. Sh rimp
larvald iets areread ily avai
lable and a reeasyto use . Manycomme rcia
l
micropar ticulate d ie ts can par tially rep lace the natura l food
requ iremen ts of shr impla rvae. A t SEAFDEC AQD .a m icrobound
diet(Tab le 7.11,F igu re 7.26)is usedto pa rt
iallyrep lace diatoms and
Artemia asfoodfo rsh rimpla rvae. Howeve r
,i tneedstobe mo rebuoyan t
and shou ldrema in suspendedin wa te rlonge rto be mo re ava i
lableto
thela rvae . Seawa te risrep laced a tthera te of 30% pe r dayf rom Z2to
Z3s tage and 50% pe r day during mys is and pos t
la rvals tages.
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 195

F
igure 7
.26
Feed
ingschemefortige
rshr
impla
rvaeZ
-zoea
,M-m
ysis
, PL
-pos
t
lar
vaea
tda
ys1to20
.
Sou
rce
:B au
ti
stae
tal.1991

3
. Sh
rimpsing
row
-ou
tponds
Shr imp pos tlarvae are re leased d irec tly in to nursery ponds .
Farme rs propaga te na tura lfood o rgan isms such as copepodsinthe
ponds by app ly ing organ ic (ch icken manure ) and/or inorgan ic
(monoammon ium phospha te 16 -20-0 ,d iammon ium phospha te 18 -46 -
0)fertilizers. In ex tens ively managed ponds , shr impfeed mos tly on
detri
tus , an ima l rema ins ,d iatoms , cyanobac teria, and g reen a lgae .
The mos t cos t-e f
fec tive way o f produc ing shr impisto grow na tura l
food in the ponds and prov ide ar tif
ic ialfeed when necessary . A
sugges tedfeed ingformu lafor shr imp grow -ou t cu l
tureis shownin
Table 7 .11 .A t 50 ,000/ha ,th isfo rmu lated d iet may be g iven s ta rting
1-1.5 mon ths a fte rs tock ing depend ing onthe amoun to f natu ralfood
inthe pond .A t 100 ,000/ha ,th is d ietis g iven soon a fterthe sh rimps
are stocked .
Ear lyjuven ile s tages pre fertofeedinthe sha llowe r wa ter a long
the wa lls ofthe d ike . As deve lopmen t proceedsthroughla tejuven ile
into the adu lts tages , the shr impfeedf ree ly onthe ben thos o fthe
whole pond . Shr imp a re s low bo t
tomfeede rs andfeed con tinuous ly
and mo re active ly a tn igh t. Theinges tedfeedtakes abou tfou r hou rs
toleavethe m idgu t
. Da i
lyfeed ingis usua lly done 4to 5t imes(0600 ,
1000, 1400 , 1800 ,and 2200 h )w i
th abou t 30% o ftheto talfeed g iven
inthe mo rning and 70%inthela te afte rnoon and even ing.
Me tabo lic ra tes o f an ima ls dec rease w ith s ize
. Thefeed ing ra teis
notfixed , butislowe red asthe sh r
imp g row . Ded icated managemen t
and appropr iatefeed ing prac tices represen t a ma jorlabor compo ­
nent bu tis cruc ialtothe success o fthe cu lture. Feed ing based on
biomass a lone w ithou t ad jus tmen t overt ime may resu l
tin under ­
feeding when sh rimp a re sma l
l, and ove r
feed ing whenthey a rela rge r
.
The mos te f
f icient waytofeed sh rimpis“by demand ”ra therthan
byfo l
low ing afeed ingtab le. Da ilyra te can be ad jus ted acco rdingto
how much o fthe prev ious day ’sra t
ionis consumed . Feed ingt rays
are usedto obse rvethe s ize and hea l
th cond ition o fsh rimp as we ll as
to mon itorfeed ing ac tivity to avo id over feed ing and under feed ing .
196 CHAPTER7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Trays measur ing 1x 1 mcan be madef rom nylon screen and wood o r
bamboo s tr
ips and s tone s inke rs(F igure 7.27)
. Usua l
ly 1to 3% o fthe
daily rat
ion a llotted tothe pondis spread be tween severa lfeeding
trays .Fivetotwen tyfeed ingt rays a re usua l
ly distr
ibu ted over one
hectare, posit
ioned no ttoo nearthe d ikes orthe pond bo ttom . Feedis
placed ont raysto mon itorfeed ing ac t
ivi
ty. Feedingt rays shou ld be
ins tal
led immed iate ly a fter s tock ing to perm i
t observa t
ion o fthe
cond it
ion and g row th o fthe sh rimp , andto de tec
tthe p resence o fany
pests orp reda tors. Thefeed a llowance can be ad justed based onthe
F
igu
re 7
. 27 amoun to ffeedle ft onthe t rays a fter oneto two hours . The nex t
Afeed
ingt
rayfo
rsh
rimp
. feed ing rationis then ad justed accord ingly based on the average
amoun to ffeed rema ining onthet rays.
Atthe end o fthe g row -ou t cycle
, sh rimps avoidvery sha llow areas
dur ing the day to avo id l ight and those areas where anaerob ic
sedimen ts and s ludge accumu la te
. Thus ,feedis d istributed on lyin
deepe rareas dur ing day time and nonein a reas where sludge depos i
ts
are high because sh rimp w ill no t consumei t
.

J
. Mud C
rabs
Various species of mud c rabs occurthroughou ttrop icalto warm and
tempera te zones. They are h ighly va
lued and a reimpor tant sources of
incomeforf ishers throughou tthe As ia-Paci
fic region. Ex tensive mud
crab(Scy l
la serrata, S.tranqueba rica,S.o l
ivacea)(F igure 7.28)farm ing
has been an on -goingindus tryfo r many decades bu tthetechno logyfo r
broodstock, hatchery, nu
rse ry, and semi-
intensive grow -out cul
tu re have
been developed on lyrecently.

1
. B
roods
tockintanks
Mud crabs a re provided w i
thf low-through aera ted seawa terin
tanks. They arefed acomb ina t
ion o fna turalfoodcons ist
ing o f mussel
meat
, rawf ish
, andfo rmu lated diet(Tab le 7.12
). Feed ingra teis 3to
5% o fbiomassfor na turalfood and 1to 1 .5%forformu lated diet
.
Feedis giventw ice da
ily wi
thfo rmu la
ted d ietgiveninthe mo rning a
t
F
igure 7
.28
Mang
rovec
rabs Scy
l
la spp
. 0800 h, and na turalfoodinthe a fternoon a t 1700 h . Reproduc t
ive
per
formance andla rval quality are betterw ithth is comb ination die
t
than on either naturalfood o rfo rmulated d ie
ta lone.2

2
. La
rva
lrea
ring and nu
rse
ry
Spawned eggs ha tch and a reraisedtozoeathento mega lopa,the
highestlarva lstage. Init
ial
ly, zoea a refed Brachionus a t 10to 15
ind/m la refedthenA rtemia naup l
iia reintroduced at zoea3and a re
graduallyincreased f rom 1to 5ind/m l as thelarvae deve lop to
megalopa(F igure 7.29). As a supp lemen t
, a commerc iallarva ld ie
t
for shr
imp may be g iven at1 .5to 2.0 g/m3wa ter pe
r day.
Megalopas are nursed un ti
lc rab stage ei
therintanks o rin ne t
cagesins talledin ponds . In tanks , food consis
to f new ly ha tched
Artemia at 3to 5ind/m lo r adul
tA r
tem ia
,thento m incedf ishf lesh,
green musse l meat
,o r smal
l sh r
imps g iven adlibi
tum tw ice a day as
soon as the mega lopa me tamorphose to crab s tage. In ne t cages
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 197
Tab
le7
.12 P
rac
tica
ldie
tfo
rmu
las(g
/kg d
ryd
iet
)fo
r mudc
rab b
roods
tockand g
row
-ou
t

Ingred
ien
ts Broods
tock Grow-ou
tdie
t Grow-ou
tdie
t
diet1 in pond 2 intank 3

Ch
ilean f
ish mea
l 200 250 335
Sh
rimp mea
l(Ace
tes sp
.) - - 75
Soybean mea
l, de
fat
ted - - 110
Sh
rimp head mea
l 200 - -
Brown musse
l mea
t - 250 -
Squ
id mea
l 200 - 65
Whea
tflou
r 170 170 118
Seaweed (Grac
ila
ria sp
.) 40 50 50
R
ice b
ran - 125 -
Co
rn b
ran - 100 -
Ce
llu
fil - - - 162
Ca
rboxy me
thy
l ce
llu
lose - - 25
Codl
ive
roi
l 50 25 14
Soybean o
il - 25 -
Lec
ith
in 30 5
Cho
les
tero
l 10 - 1
V
itam
in m
ix 30 - 15
M
ine
ral m
ix 40 - 5
D
ica
lcium phospha
te 30 - 20
E
thoxyqu
in - 5 -

Prox
ima
te compos
ition (% DM)
;
Crude p
rote
in 46
.0 40
.1 43
.3
Crudefa
t 11
.6 11
.9 4
.7
Crudef
iber 4
.2 1
.4 12
.0
N
itrogen
-free ex
trac
t 23
.2 38
.0 27
.1
Ash 15
.0 8
.6 12
.9

Sou
rces
: 1M
il
lamenaand Qu
ini
tio2000
;2T
riñoand M
il
lamena2001
;3Ca
tacu
tan and Tesh
ima2001

F
igure 7
.29
Feedingscheme usedinthecu
ltu
re o
f mudc
rabla
rvae
.
Sou
rce: Qu
ini
tioe
tal
. 2001
198 CHAPTER7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

installedin brackish water nursery ponds,c rab mega lopas arefed

macerated b rown musse
l meato rfishflesh at 30to 20% o fb
iomass
per day. Feedrat
ionisgiven a
t0800 , 1300, and 1700 h dai
ly.Stocking
30 mega lopas/m2in net cagesisfeasible
.

3
. Pond cu
ltu
re
Cultureo fe i
ther ma le,fema le
,o rbo thsexeso f mudc rabinea r
then
ponds(F igu re 7 .30)o rin pensins tal
ledint idalf lats wi
th re forested
mangrove a re econom ically v iable. S tocking dens ity of 1/m2g ives
high surv iva l. Rawf ish andf resh brown musse l mea ta re used as
feed . Crabs a refed a t 10% o fthe irb iomass da i
ly when carapace
leng th(CL )is <6 cm and 5% when CLis >6 cm . The dai
lyfeedra tion
is broadcas ted over areasin be tween mangrove t rees o f each pen ,
40% a t 0700 h and 60% a t 1700 h . Feed ra t
ionis de term ined by
placing 1% o fthe schedu ledfeed ing ra t
ion onfeed ingt rays. When
feed onthet rayis consumed a fterfour hou rs,thera t
ionfo rthe nex t
feed ingisincreased by 5% . Feed ing rateis reduced by 5% when
0.5% o r mo re ofthefeedisle ft unea ten onthe t ray wh ile nofeed
adjus tmen tis madei flessthan 0 .5%feed rema ins onthet rays. D ry
art
ificiald iet can a lso be used as g row -outfeedfo r mud c rabs(Tab le
7.12). This d ietis given a t 5% o fb iomass da ily when CLis <6 cm and
2% when CLis >6cm .The da ilyfeedra t
ionisg ivena t0730and 1700 h .
Mud c rab can a lso befed Ace tes , greenf i
lamen tous a lgae, anima l
hides and en t
rails
, and sna i
ls.C rabs a refed at 10%then 6% o fbody
weigh t as cu lture progresses . Da ilyfeed a l
lowanceis p laced on a
feedingt ray o r broadcas ttw ice a day , halfofthefeedinthe mo rning,
halfinthe a fternoon .
A mud c rab a rtif
iciald iet(Tab le 7 .12) has been deve lopedinthe
labora tory .I tis presen tly used as ma intenancefeedfor mud c rab
unde rindoo r conditions .

F
igu
re7.
30
G
row-ou
tcul
tureinpond(A
)andinpen
sin
sta
l
ledinre
fore
sted mang
rove
s(B
).
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 199
O
the
rspec
iesfo
raquacu
ltu
res
tockenhancemen
t

A
. Donkey
’s ea
r aba
lone

Donkey ’s ear abalone (Halio

tis as inina)(F igure 7.31A)is a gas tropod
mol
lusktha tl
ive around algals tandsin rocky and sha l
low sea wa ters.
Abalones are popu larin spec ialty restauran tsin the Ph i
lippines and
other Sou theast Asian countries bu ttheir dec l
ining y
ieldf rom the w i
ld
necessi
ta tesintensive aquacultu re. Research a t SEAFDEC AQD a imsto
develop seed produc t
ion and cu l
tu retechn iquesin orderto p roduce seed
for stock enhancemen tinthe w i
ld. Abalones a re herbivores andfeed
mainlyonseaweeds . Amongtheth ree naturallyoccurring abalonespec ies
inthe Ph i
lipp
ines, donkey’sea r aba lone g
rowsto a max imum s ize of 100
to 110 mmin she l
lleng th(SL ).

F
igu
re7
.31
Donkey
’sea
raba
lone Ha
l
iot
isas
in
ina (A
),feed
ingonreda
lga(B
). Tab .13 P
le7 rac
tica
ldie
tformu
la(g
/kg d
ry
d
iet
)fo
raba
lone broods
tock

1
.Breed
ing Ingred
ien
ts Amoun
t

Aba lone breeders are kep tin per fora ted F

ish mea
l 100
.0
plastic baskets and fed the red a lga , Sh
rimp mea
l 150
.0
Gracilar iops is (F igure 7.31B ), to satiation De
fat
ted soybean mea
l 200
.0
replen ished a t week ly interva ls. Cap tive R
ice b
ran 149
.5
abalone a t
tains sexua l ma turity with
in 6to 8 Whea
tflou
r 200
.0
months o fcu lture at shells izerange o f 35to Seaweed 70
.0

40 mm . Artif
icial d iet (Tab le 7 .13 ) in Codl
ive
roi
l 15
.0
Soybean o
il 15
.0
comb inat ion w ith Grac i
lariops is improves
V
itam
in m
ix 30
.0
ferti
liza tion ra te and spon taneousfecund i
ty
M
ine
ral m
ix 40
.0
in aba lones . Aba lones a refed 2to 5% o fthe ir
D
ica
lcium phospha
te 30
.0
body we igh t per day usingthe d ry d
iet wh
ilethe BHT 0
.5
red algais g iven adl ibitum . The d ryra tionis
Prox
ima
te compos
ition (% DM)
:
given once a t 1600 h .
Crude p
rote
in 27
.9
Crudefa
t 5
.8
Carbohydra
te 40
.4

Sou
rce
:1Bau
tis
ta-Te
rue
land M
il
lamena 1999
200 CHAPTER 7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

2
. Pos
tla
rva
lse
ttlemen
tandnu
rse
ryrea
ring
Set
tlementtanks a re p repared one week before stockingto al
lowthe
growth ofepiphytic diatoms onthe p late substrates(Figure 7.32A,B )
.
Vel
igerlarvae and earlyjuven ilesfeed on ep iphy t
ic diatoms such as
Navicula andNitzschia, then g radual
ly shifttored a lgaG rac
ilar
iopsis
whentheyreach >5 mm SL .
Feedinglarvae a commerc ia
lfeed orthe SEAFDEC formu lated
die
t(Tab le 7
.13)isfeas ible. Thesefeeds a re given at 3to 5% o fthe
body weighttw ice daily.

F
igure 7
.32
P
la
tesub
stra
tes(A
)fo
r ep
iph
yt
icd
iatom
s,l
ivefoodfo
rla
rvaein
sta
l
ledintanks(B
)
.

3
.Grow
-ou
t
Culture ofaba lonefrom 30 mm SLjuven i
lesto ma rketable si
ze o f
55to 60 mmisca rr
ied outinflow-throughtanks andinseacagesfo r
8to 10 mon ths. Abalones arefedfresh seaweedsto satiat
ion given a t
weeklyin tervals.
In cages ,feedingra tes as we
ll as g
row th and survivala re higher
atlowe rs tocking densit
ies(17to 35 pe r cage). The rapidly growing
smalljuven iles(16 -20 mm )feed at 35to 40% o f body weight wh ile
the bigger abalones(> 50 mm )feedless, only 5to 10% ofbody we igh t
.

B
. Seaho
rses

Live seahorses(F igure 7 .33A)a ret raded onthe aquar iumf ish marke t
whilethe d r
ied ones a re high ly valuedint radi
tional Ch inese med icine.
Seahorses came nearto ex t
inc t
ion dueto des truct
ion o ftheir na tural
hab itat and overexp loita t
ion .A t SEAFDEC AQD , breed ing and seed
produc tiontechn iques a re being unde rtaken on H ippocampus kuda and
H. ba rbouri u l
timatelyfo rs tock enhancemen tinthe w ild.
Livefood organ isms are used asfood for seahorses to breed and
produce seedsintanks(F igu re 7.33B). The comb inat
ion d ie
to f HUFA -
enr iched Ar tem ia adu lts , mys ids, or tilapia fry promo tes frequent
partur it
ion even ts and grea t brood s ize perfema lein tanks . Artem ia,
mysids , andtilapiafry are given a t 15
,6 , and 5% o fthe body we igh t per
day, respec t
ively. A comb ina t
ion d ietofro ti
fers and copepods enhances
surv ivalo fH. kuda la rvae.
 CHAPTER 7 Managemen
tof Feed
ing Aquacu
lture Spec
ies 2
01

F
igu
re7
.33
Seaho
rses(A
)andarea
ringtank(B
).

C
. W
indow
-paneoys
ter

Window-pane oys tero r kap is she l

l(Placuna p lacen
ta)(F igure
7.34A) is a b ivalve mo llusk that is com merc ially and
econom ical
lyimpor tantinthe Ph i
lippines
. The irt
rans lucen t
shel
ls are used asraw ma terialsforthefab ricationofbeau t
iful
handicraftslike window s il
ls, wa l
l panels,lamp shades , and
chande l
iers(F igure 7.34B ,C )
. W ith very h igh world marke t
demandforthese produc ts, they become overexp loited and
their natura l popu lation is near ly dec ima ted. Ar t
ificial
propagationisca rr
ied outa t SEAFDEC AQDino rdertoreseed
and enhance the dec lining na tural popu la t
ionin the w ild
,
thus rehab i
litat
ingthe kap is indus t
ry.

1
. B
rood
stockandseedp
roduc
tionintank
s
Broods tockfed da ily w
ith Isoch rysis andTe t
raselmis a t
3:1 rat io at 200 ,000 ce lls/m l promo tes rap id gonad
developmen t
. Sexua l maturityisreached a f
ter 4 mon ths,
and theninduced spawn ingis done to produce v iable
game tes . Thelarvae a re rearedinf iberglass tanks a ta
dens i
ty o f 900larvae perl iter
. Larvae grow and surv ive
bestw ithIsoch rysisasfooda tinit
ialconcen t
rationof 10,000
cel
ls/m l, and a re progressivelyincreasedto 30 ,000 cells/
ml asthela rvae grow. Se ttl
ingisreached a fter 14 days.

2
. S
tock enhancemen
t F
igure 7
.34
W
indow-paneoys
ter(A
)
,and hand
ic
raf
tsu
sin
g
Kapis she l
ls are f i
lter-feeders and the irfood cons istof kap
isshel
ls(B
,C)
.
plankton , detritus and other suspended mater ia ls in the
estuary. The natura l beds of kap is shells are the b luish-
soft mud or s l
ight ly sandy-muddy subs tratesin areas w ith
high primary produc tivity
. To rehab il
itate the natura l beds
of kapis shella tT igbauan ,I lo
ilo coastl
ine(F igure 7 .35A ,B)
,
im mature broods tock(72 m m SL , 14
.5gbody wt ) ares tocked
at 75ind/m2 . The broods tockfeed onthe ava ilab le na tural
food in the estuary . Two months a fter reseed ing, they
spawn natura l
ly.
2
02 CHAPTER 7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

F
igu
re 7
.35
Rehab
i
li
tat
iono
fIlo
i
locoa
st
line(A
,B)
.

Summa
ry

The ob jective o ffeed managemen tisto make ava ilab letothe an ima ls
the mos t cos t-effec tive,fo rmu latedfeedinthe prope r amoun ts and a t
the r ightt imes andloca tions . Feed ing prac t
ices mus t be con tinua l
ly
mod ified and adap ted to accoun tfor changesinfeed ing ac tivi ty and
prefe rences asthean ima lsg row and asenv i
ronmen talcond itionschange .
Feed managemen t prac tices a re spec i f
ically adap tedto spec ies, cu l
ture
sys tem , and a reain o rde rtoinc rease p roduc tion e fficiency and m inim ize
env ironmen talimpac t.
Thefarmer wan ts the cu ltured aqua tic an ima lto grow max ima lly
with a m in imum amoun to ffeed was te. Thus , one shou ld have a good
know ledge o f how muchthef ish shou ld befed andthen mon itorthe ir
feed ing ac t
ivity ca refu l
ly . Feed consump tion can vary by spec ies ,s ize/
age o f anima ls,in tens i
tyleve lo f cu l
tu re, wa tertempe rature ,d issolved
oxygen ,feed a t
trac tab i
li ty and wa ter s tability,f requency and t imes o f
feed ing, and o the rfac to rs. Feed ingra tions a rebased onfeedconsump t
ion
fromfeed ingt rays ,o rsu rvivales tima tes , ave rage s i ze, andfeed ingtab les
deve loped bythefa rme rh imse lf using da taf rom seve ra l cultu re cyc les .
Many b iotic and ab ioticfac tors a re used to ad jus tfeed quan t
ity , and
these cr iteriainc ludelow DO , ex treme wa tertempera tures , excess ive
plank ton b looms ,feed consump tion , mo lting/ lunar cyc le
, an ima ls ize
and wea ther.
How o f
tento app lyfeedis animpo r
tan t dec isionthe cu ltur ist hasto
eva luate and de termine based on exper ience , season , chang ing
env ironmen ta l cond it
ions , spec ies , age and s ize,s tock ing dens ities ,
produc tion sys tem used ,s tress , ava ilab le resources , and other
cons ide rations . Cons tan tfeed ing o f new ly ha tchedla rvaeisimpo rtan t
fo r good su rvival. Pa rt
icle s ize shou ld beinc reasedin c losere la tion w ith
grow th.
Hand feed ingis common ly used and has the advan tage o fc lose
obse rva t
iono fthef ish . The reisthen anassu ranceo ffeed ingto max imum
effectiveness and m inimum was te sincethef ish cu l turistis mo re awa re
ofthe cond itions and p rob lemsi fthey deve lop . Au toma tic and demand
feede rs have advan tagesintha tthey can belabor -sav ing and can a llow
fishto befed manyt imesth roughou tthe day . Au toma t
icfeede rs can be
used as a use fula l terna t
ive to manua l broadcas ting , bu t shou ld no t
 CHAPTER7 Managemen
tof Feed
ing Aquacu
ltu
re Spec
ies 203
replaceregular observat
ion andv isual assessmen
t byt ra
ined personne
l.
Consider
ing the d ivers
ity of spec
ies and produc t
ion systems, bes
t
managemen tp ractices
, particu
larlythose onfeed ing managemen t,are
impor tant steps towards assur ing that aquaculture wi
ll grow in a
responsib
le manner and can be sus tained.

Gu
ideQues
tions

1
. Compa
rethe d
iffe
ren
tfeed
ing s
tra
teg
iesin pond cu
ltu
re.
2
. Whyis supp
lemen
taryfeed
ing cons
ide
red asthe mos
t cos
t-e
ffec
tive
feed
ing s
tra
tegy?
3
. D
iscusstheimpo
rtance o
fsamp
ling andreco
rd keep
ing
, ad
jus
tmen
ts
o
ffeed
ingra
tions and use o
fapp
rop
ria
tefeed s
izeto op
tim
izethe use
o
ffeeds
.
4
. How a
refeed ra
tions andto
talfeed requ
iremen
t de
term
ined?
5
. Wha
tarethe s
ign
ifican
t con
tribu
tingfac
torsinthe cho
ice o
ffeed
ing
me
thods?
6
. Wha
tarethe advan
tages and d
isadvan
tagesinvo
lvedin manua
l vs
.
mechan
ica
lfeed app
lica
tion me
thods?
7
. Why does the h
igh va
lue for FCRind
ica
te poorfeed convers
ion
e
ffic
iency?
8
. Wha
tarethe ma
in was
te p
roduc
ts assoc
iated w
ithfeed
ing and how
a
rethey m
inim
ized?
9
. Wha
tfeed nu
trien
ts a
re agen
ts o
f“eu
troph
ica
tion
”inrece
iving wa
ters
forf
ishfa
rm e
ffluen
ts? D
iscussthe
ir ha
rmfu
l consequencesinthe
env
ironmen
ts.
10
. How do we de
term
inethe app
rop
ria
tet
ime andf
requency o
ffeed
ing
forf
ish and c
rus
taceans?
11
.Discussthefac
torstha
tin
fluencefeedconsump
tion o
faqua
tic an
ima
ls.
12
. Wha
tarethe do
’s and don
t’sinfeed
ing managemen
t?

Sugges
ted Read
ings

A
lava VR,Lim C. 1988 . Ar
tif
icia
ld ie
tsfor m i
lkfish Chanos chanos
(Fo
rsska
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Bau
tis
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lamena OM, Kanazawa A
. 1989
. Use o
fk-carrageenan
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iet(c-MBD) asfeedfor Penaeus monodon larvae
.
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.

Baut
ista-Terue l MN, Mil
lamena OM . 1999. Diet deve
lopment and
evaluationforjuvenile aba
lone, Hal
iot
is asin
ina: pro
tein/energy
levels
. Aquaculture 178:117-126.
204 CHAPTER 7 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Bombeo RF, Fermin AC, Tan -Ferm in JD. 2002

. Nurseryrearing o
fthe
Asiancatf
ish
,C lar
ias mac rocephalus (Gunthe
r),a
tdif
ferentstocking
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tiesin cages suspendedintanks and ponds .(Aquaculturein
press
).

Bo
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Ca
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ios
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l, and body composi
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, Lates calcari
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Ca
tacu
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r GE , Teshima SI
. 2001.E f
fecto fdietary prote
in
andlipidleve
ls and proteinto energy ra
tios on growth, survival
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tion o fthe mangrove red snapper Lu tjanus
argen
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). Aquac. Res. 32:811-818.

Cap
inp
in ECJr
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iM . 1999. Dens
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liot
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.
Aquacul
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Ch
iu YN. 1988. Water quali
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ilo C
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Con
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cat
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rias mac rocepha lus juveni
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ferentd ietsin
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ian Fisheries Soc
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Duray MN. 1998

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,A lpasan LG
. 1997
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Duray MN,Alpasan LG, Estud i

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tcheryrear
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.J. Aquacult.
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Fe
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-
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Fe
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Ferm
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iotis as
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Ferm
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ist
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206 CHAPTER 7 NUTR
ITIONIN TROP
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Go
lez NV. 2000.E f
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Ha
ra S
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itf
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.

H
ilomen-Garcia G
. 1999. AQD’s mar ine ornamenta
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t, In
:
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Mad
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indow -paneshel
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l Publica
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Mad
rones-Ladja JA
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. The ef
fecto
fm icro
a
lgald ie
t andrearing condi
tion on gonad maturity
,fecundity and
emb ryon
ic deve
lopmen tofthe window-paneshe l
l,Placunaplacen ta
L
innaeus . Aquacu
lture, 206:313-321.

M
illamena OM, Primavera JH , Pudadera RA , Caballero RV. 1986. The
ef
fectofdiet onrep roduct
ive pe r
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los
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M
illamena OM . 2002. Rep lacemento ffish mea
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t for grow-out cu l
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,
Epinephelus co
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.

M
illamena GM , Trino AT. 1994 . Eva
luat
ion offish prote
in concen trate
andlactic yeast as potent
ialprote
in sourcesfor shr
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M
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ffec
ts ofd
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t mudcrab Scyllaser
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Mo
toh H
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razo MM . 1991. An arti
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iald ie
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ini
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illamena OM, Rodr
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les
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.

Qu
ini
tio ET , Estepa FP,A lava VR . 1999. Developmen to f ha tchery
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ian
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,A ldaba MB, Abuan E
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ts ofart
if
icia
l
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romis n
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ietaryc rude
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ilo
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lo fthe Phil
ipp ines
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ietform onthe g rowth and surv
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ilo
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r
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icia
l
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208 CHAPTER7 NUTR
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. 2000.Effec
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fec ts o
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ture 132:273
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Tr
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lture 174:109-118
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Tr
iño AT, Mi
llamena OM, Keenan C. 2001. Pond cu l
ture of mud c
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Scyl
laserra
ta (Fo
rska
l)fedformulated die
tw ith orwithoutv
itamin
and minera
l supp
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ian Fish
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Tr
iño AT, Rodr
iguez, EM. 2002
. Pencu
lture o
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rrata in
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. Aquacul
turein p
ress.
 CHAPTER

In
troduc
tion
Econom
icso
fFeed
ing
RENATO F
. AGBAYAN
I
8
The numbe rofaquacu lturefa rme rs who adop th ighe rs tock ing dens ities
and p rovide supp lemen ta ryfeedstof ish stocksisinc reas ing because o f
increased produc t
ion andimproved pro fits
. For examp le, in shr imp
cultu re, the na turalp roduc tivity o fthe wa ter may gene ra tef rom 100to
300 kg/ha/y r andfe r
tilization mayfur therinc rease produc tionto 600 -
1,000 kg/ha/y rinthe Ph i
lipp ines. The use o ffeeds canra ise p roduc t
ion
upto 20 ,000 kg/ha/y rin Ta iwan and 30 ,000 kg/ha/y rin Japan .
In shr impfarm inginthe Ph ilippines ,a lthoughthe reis aninc rease
in p roduc tion,the cos to ffeedstakes up 54 -63% o fope ratingcos ts when
stock ing dens ityisinc reased such asinthe sem i
-intens ive andin tensive
farm ing me thods . In mudcrab cu lture , cos ts o ffeedsis 50 -57% o f
ope rating expenses wh ilein m i
lkfish cu lture,the cos ts o ffeeds comp rise
10-77% o ftotalcos ts. Hence , aneva luation o ffeed qua lity and econom ic
eff
iciencyis a veryimpor tan t unde rtakingto de term inethe pro f
itabil
i ty
of an aquacu l
ture ven ture .
Th is chap ter a ims to in troduce concep ts and me thods in do ing
econom ic analysis app licab lein aquacu l
turein genera lw ith emphas is
in feed produc tion and feed ingin aquacu lture farms . Th is chap ter
discusses thefo llow ingtop ics : cos to f produc ingfeeds ;s imp le single ­
inpu t( feeds) and s ing le ou tpu t( f
ish ) produc tionfunc tion; ind icesfo r
measur ing econom ic efficiency o ffeeds ; theleas t-cost comb ina t
ion ; and
linear programm ing as usedinthe a l
loca t
ion o flim ited resources such
asfeedingred ien ts tha tw ill mee tthe nu tritiona l requ iremen ts ofthe
fish.

Cos
tofp
roduc
ingfeeds

The cos titemsin produc ing aquacu lturefeeds can be c lassif

iedin to
direct costs andind irect costs. Direct costsinc ludethe raw ma terials
such asf ishmeal, m ineral and v itam in mixes, and o theringred ients,
and directlabor usedinthe manu factureo fthefeeds. The direct materials
become pa r
to fthef inalp roduc tsinthefo rm offeeds. D irectlabo r cost
itemsinc lude wages and sa lar
ies ofthe worke rstha t manu facture the
feeds.
Indirect cos
tsinc lude (1 ) supervisory and managemen t overhead
expenses ;(2)electr
icity;(3) supp l
ies otherthanfeeding redients such as
gaso l
ine and mach ine o il
,i f any ; (4) repair and ma in tenance ; (5)
210 CHAPTER8 NUTR
ITIONIN TROP
ICAL AQUACULTURE
depreciat
ion o ffixed asse ts such as equ ipmen t, bu ildings; and(6 )o the r
incidental expenses requ iredin manu fac turing thefeed . The cos to f
packaging and s to rage o fthefeedsis no tinc ludedinthe compu tation .
Deprecia t
ionisthe a llocation o fthe or iginal cos to fa llfixed asse ts
over their econom ic or use fu lli
fe . For examp le,i f afeed m il
l cos ts
P1,000,000and w i
llbeeconom ica
lly usefulfo rten years ,the dep reciation
is compu ted by d iv iding P1 ,000,000 over 10 years , wh ichis equa l to
P100,000.I tis a non -cash expense becausethe moneyfo r cons t
ruc ting
or purchas ing a f ixed asse tsuch as afeed m i
ll has been used up bu tis
being al
loca ted ove rthe pe riod ofi ts usefull i
fe.
Table 8.1 shows al isto fcos titemsin p roduc ingsh r
imp d iet. The d irect
costofing red ients comp rises 92% o ftheto tal costo fp roduc ingfeeds .
The cos to fd i rect labor used in gr inding , pu lverizing, m ixing ,
pel
let
izing, and oven d ryingis compu ted by mu ltiplyingthe numbe r(2 )
of workers w iththe wagera te(P200 pe r day )fo r examp le whichis equa l
to P400. The cos to fe lect r
icityiscompu ted by mu ltiplyingthe numbe rof
hours used(200 -kilowa tt hou r)w i
ththe cos t(P2 .50 pe r kwh) wh ichis
equalto P500 .
Theto ta lcos to fp roduc ing oneton o fsh r
impfeeds wh ich amoun ted
to P25,293 .83isthe sum o fthe d irec
t cos t(ing redien ts and direc tlabo r
)
andtheind irect cos to r P25 .29 pe r kg.

Tab .1 P
le8 roduc
tioncos
tofsh
rimpd
iet

I
tem Quan
tity Un
itcos
t pe
rkg Cos
t/kg Cos
t/ton

A Ingred
ien
ts Compos
it
ion (%) (
in Ph
il
ipp
ine Peso
)

Fish mea
l 25
.00 25
.00 6
.2500
Soybean mea
l 25
.00 11
.25 2
.8125
Sh
rimp head mea
l 15
.00 21
.00 3
.1500
Breadf
lou
r 16
.00 8
.60 1
.3760
Rice b
ran 3
.95 4
.96 0
.1959
Seaweeds 5
.00 12
.00 0
.6000
Codl
ive
roi
l 2
.50 84
.79 2
.1198
Soybean o
il 2
.50 45
.00 1
.1250
Vi
tam
in m
ix 2
.00 200
.00 4
.0000
Mine
ral m
ix 1
.00 130
.00 1
.3000
Dica
lphos 2
.00 14
.00 0
.2800
Ethoxyqu
in 0
.05 8
.00 0
.0040
Sub
tota
l 100
.00 232
.132 23
.2132

B
. D
irec
tlabo
r 2 a
ides 200
.00 400
.00
(g
rind
ing
, pu
lve
riz
ing
,
m
ixing
, pe
lle
tiz
ing
,
oven d
rying
)

C. Ove
rhead
Elec
tric
ity (200 kwh
) 2.5 500
.00
M
isce
llaneous 1,180
.66
(5% o
fItems A & B
)
Sub
tota
l 1,680
.66

D
. Cos
t pe
rton 25
,293
.83

E Cos
t pe
r kg 25
.29

Sou
rce
: M
il
lamena 1994
 CHAPTER 8 Econom
ics o
f Feed
ing 2
11
S
ing
le-
inpu
tands
ing
le-ou
tpu
tproduc
tionfunc
tion

The prev ious sec tion d iscussed s imp le me thods o f comput ing the
produc tion cos tof feeds us ing severalinpu ts. Feeds,intu rn,a re one o f
theinpu tsin produc ing f ish. Th ischap ter describes aneconom ic me thod
of eva lua ting the re lationsh ip be tween theinpu ts used to produce a
particu lar outpu t. Th is me thodis ca l
led p roductionfunc tion.P roduc tion
func t
ionis de fined asthetechn icalre lat
ionship be tweenthefa rminpu ts
andthe ou tpu ta tag iven t ime us ing atechno logy.S imp ly stated, the
totaly ield o rou tpu tva r
ies w iththe quan ti
ties and comb ina t
ion o finpu ts
usedinthe produc tion p rocess . Thef ish growe ro rthe managemen to f
corpora tefarms dec ides wha t, how , when , and how muchto produce
from thel imited resources o fthe company .
In aquacu lture , farm inputs are genera lly compr ised o ff ry or
fingerlings ,feed ,fer ti
lizers, chem icals,labo r
, as we ll astechn ical and
managemen t services . The des ired ou tpu tisthe ma rketable-sizef ish.
For examp le, the re lationsh ip between the var iousinpu ts, deno ted
as X1,...Xn , and an ou tpu t
, deno ted as Y,is expressedinthefo llowing
equa tion :

Y =f(X1
,X2
,X3
,X4
,X5
,…Xn
)

where
:

Y =tota
lf ish yield(ou tput)
X1= amoun toffeed
X2= s
tocking size off inger
lings
X3= amoun toffe rt
ilizer
X4= s
tocking dens i
ty
X5= amoun toflabo r
Xn= o
ther var
iab les relatedto grow
th o
ffish andto
taly
ield

The ma thema tical express ion showstha tthef ish y ield(Y )is re lated
oris a“ func tion”o fthe va r
iables o rproduc tioninpu ts(X1 ,…Xn)in some
particular way . The purpose o fthe produc t
ionfunc t
ion ana lysisisto
estimate the phys ica l and marg inal re lationsh ips be tween ou tpu t
(dependen t variable) and a number o finpu ts( independen t variables).
Theinpu ts o rindependen tva riablestha ts ignif
ican tlyinfluencethey ield
areinc ludedinthe produc tionfunc t
ion. The re may be o the rinpu tstha t
are used bu ttheir ma rg
ina linfluenceonf ish produc tionis no ts ignif
ican t.
As imp leinpu t( feed) and s ing le ou tput( f
ish ) produc t
ionfunc tion
(Shang 1990 )is shownin Tab le 8 .2. Theleve lo ffeed ingg iventothef ish
is represen ted by Y and the ou tpu t ortheleve lo fthe to tal phys ical
produc t(TPP )isrepresen ted by Y . The Y o rTPPinc reases upto ace r
tain
level astheleve lo f X1( feed)inc reases . The h ighes tleve lo f outpu to f
produc tionis 69 un its. This ou tpu tleve lis a t
ta ined whenfeed ingleve l
is 11 un its. This produc t
ionleve lis ca l
led the max imum sus tainab le
yie
ld(MSY ). They ield o r TPP sta r
tsto dec rease beyondth isfeed ingleve l.
There fore, thereis no reason orlog icinincreas ingtheleve lo finpu t
when MSY has been a ttained. Thereis no add itiona l bene fi
tin g iving
additiona lfeedstothef ish beyondthe MSY .
212 CHAPTER 8 NUTR
ITIONIN TROP
ICAL AQUACULTURE

The average physical product(APP)isthe amoun toffish produced

per unitofinpu t(
feed)g iventothef ish.I
tis computed by d
ividingthe
tota
l phys ica
l produc t(TPP ) overthelevelofX1(feed)
. The marg ina
l
physical product(MPP )is de f
ined as theincremen
t orthe changein
outpu t(f
ish) resu
ltingf rom one add it
ional un
itofinpu t(feed)given
thef ish
.

Tab
le8
.2. Re
lat
ionshipamongtota
lphys
ica
lproduc
t,ave
ragephys
ica
lproduc
t,
and ma
rginalp
roduct
Leve
lof X1 Averagephys
ica
l
Leve
lofto
tal Ma
rgina
l phys
ica
l
product
comb
ined w
ith phys
ica
lproduc
t(Y
) (APP ) produc
t
m
ixed resou
rces (TPP
) (MPP
)

0 0
0.5
1 0
.5 0
.5
1
.5
2 2 1
3
3 5 1
.67
7
4 12 3
11
5 23 4
.6
16
6 39 6
.5
13
7 52 7
.43
8
8 60 7
.5
5
9 65 7
.22
3
10 68 6
.8
1
11 69 6
.27
-1
12 68 5
.67
-3
13 65 5
-5
14 60 4
.29

Sou
rce
: Shang 1990

Graph ica l
ly, the re la
tionsh ip among TPP , APP and MPPtothe y ield
orleve lo f outpu tY , andlevelo finpu t(X1, …Xn)is shownin F igure 8 .1.
The g raph can be d ividedin tothree stages.In S tageI , TPP, APP and MPP
exh ibit increas ing re turns . Fur thermore , the TPP increases a t an
increas ing ra te. MPPinc reases un t
il marg inalincremen t per unitis a t
its peak . APP cont inues to increase . Stage I I is the phase where
dimin ish ing re turns occu rs
. MPP dec reasesfo r every additional un i
to f
inpu t( feed )giventothef ish.A tthiss tage, TPPiss ti
l lincreasing al
though
at a dec l
in ing ra te. The peak o f the TPPisthe max imum sus tainab le
yie
ld(MSY )o rthe po int whe rethereis nolonge raninc rease ofp roduction
in sp ite o fthein t
roduc t
ion o f an add i
tional un itofinpu t(feed)in tothe
fish cu ltu re system .A tthis pointthe MPPisze ro. APPis s ti
llincreas ing
but beg ins to dec line. Beyond th is po intis S tageI I
I orthe dec lining
 CHAPTER8 Econom
ics o
f Feed
ing 213
phase. Thereis nolonge r an advantage o
fp rov
iding an addi
tionalinpu t
becausethe re w
ill be a decreaseinthe ou tputo rproduction y
ield. This
meanstha tfeedingthef ish a tthis s
tage w i
llresultto decreaseinf ish
production
.

F
igu
re8
.1
Re
lat
ionsh
ipbe
tweenTPP
,APP
,and MPP
.

Thep
roduc
tionfunc
tionandthecos
tofp
roduc
tion

Thein t
roduc tion o fcostin produc tion analysisis an econom ictoo ltha t
can gu ide thef ishfarmerin h is dec isions related to produc t
ion. The
main ob ject
ivein produc tion analysisisto max imize profi
t. Wha tlevel
of produc tion w il
l resultto max imum pro fi
tg iven aleve lo fresources
such asfeed? I f afishfarme rw i
ll base h is decision onthe p roduct
ion
func tion discussed ea r
lier
, he may betemp tedto max im ize production
uptothe MSY(Tab le 8
. 3and Figu re 8.2)or whe rethe p roduc t
ion outpu t
is atthe max imum . However
,th is section will showtha tthef ishfarme r
shou ld cons iderthe econom ic pro fi
ttha t cou ld be attained a t var
ious
levels o finpu t
.

Le
t us de
fine some newte
rms a
tth
is po
int
.

□ Value o
fTPP(VTP )isthe mone ta ryvalue o fthe output(fish) based
onthefa rm ga te pr
ices orprices o fthe p roduce so
ldinthefa rm.
As an examp le, Table 8
.3 assumedtha tthe price of one un itof
outputis equ iva
lentto one peso(P1 .00).
□ Value of APP (VAP)is APP mu ltiplied bythefa rm gate price
.
□ Value of MPP(VMP )isthe add itional revenue or sales resu l
ting
from aninc reasein add it
ional un itofinpu t
.
214 CHAPTER 8 NUTR
ITIONIN TROP
ICAL AQUACULTURE

□ Totalvar
iab
leinpu tcos t(TVIC
)isthesumo fthecostoftheva r
iab
le
inpu t(
feed
)a tthe d i
fferen
tlevels o
finput
.
□ Prof
itisthe d
ifference betweenthe VTP o
rg rossrevenue andthe
TVIC orinput cost
.

Thelas tco lumn o fTab le 8

.3 showsthe h ighes tp rof
itlevelis 20 un its
whichis a ttained a tthefeed inglevelo f 8 un i
ts andthe co rrespond ing
yieldis 60 un its. This pointis the max imum econom ic yie
ld (MEY ).
Figure 8 .2i l
lus tra
testhe re lat
ionship between MEY and o the r cost and
revenue va riables. VTP o rthe value oftheto tal produc tisthe revenue
curve . The T ICisthe cos t curve. The shaded areais the pro fi
t area
whe rethe d ifference be tweentherevenue andthe cos tare positive. The
bigges td i
fference be tween thetwo curvesis a tthe MEY . The MEYis
reached be forethe po in t where maximum y ield o rthe MSYis a ttained.
There fore, w ith profi
t asthe ma in objective, the des iredfeed ingleve l
shou ld be atthe MEY whe reitis mos t cos tefficient.A tfeed ingleve ls
beyondthe MEY ,thereis no add it
ionalo r ma rgina l econom ic bene f
itin
inpu tt
ing add itional unitoffeed sincethe add it
iona l costo ffeed w il
l be
grea terthanthe add itionalincome de r
ivedf romthe add it
ional ou tput.

There aretwoconclus
ionsthatcanbe de rivedfromTable 8
.3 and Figure 8
.2;
1. max imiz
ingfa rm production does no talways maximize p
ro f
its
;
2. decision on maximizing profi
tis based on marg inal analysis or on
whetherthe reis addi
tional benef
itf rom addit
ionalinput( feed
).

Econom
ic E
ff
iciencyo
fFeeds

The usua lp ract

icein measur ingthe e f
ficiency offormu latedfeedsisto
measure thefeed convers ion rat
io (FCR ). FCRis de fined as the to ta
l
amoun toffeeds by we ightgiven over a cer ta
in period d iv
ided bythe
yie
ld in k i
lograms . Thelower the ra t
io , the more e ff
icient the feed,
regardless ofthe cost.
FCR , howeve r
, does nottakein to accoun tthe cos to ffeeds andthe
econom ic e
ffic
iency of usingfeeds. A method o f measuringcos teff
iciency
offeeds calledtheinc idence cost
,indica testhe costo fp roducing a unit
(by weight)o ffish.

Inc t = to
idence cos tal cos
to ffeed used
wto fthefish produced

In compar ingtwofeedformu lat

ions ava i
lable to afishfa rmer, the
feed w ith alowerinc idence cos tis more econom ically ef
fic
ient and,
therefore, more benef
icia
ltothefa rmer.
Anothe r method of measuring economic effic
iency offeedsistolook
atthe va lue ofthefish produced andthe cos to ffeeding. Thisis ca
lled
the profitindex, whichind icatesthe profi
tfor every un it cos
to ffeeds
incurred . Th
isis compu ted usingthefo l
low ingformu la
;

to
talva
lue o
fthef
ish p
roduced-to
tal cos
toffeeds
Pro
fitindex =
to
tal cos
toffeeds
Tab
le8
.3Re
lat
ionsh
ipamongto
talphys
ica
lproduc
t,ave
ragephys
ica
lproduc
t, marg
inalphys
icalproduc
t,va
lueo
fto
talphys
ica
lproduc
t,va
lueo
fave
rage
phys
ica
lproduc
t,va
lueo
f ma
rgina
lphys
ica
lproduc
t,to
talva
riab
leinpu
tcost,andprof
it
.

To
tal Average Marg
ina
l
Leve
lof X1 Leve
lofto
tal Ave
rage phys
ica
l Ma
rgina
l Phys
ica
l Va
lue o
f Va
lue o
f Va
lue o
f var
iab
le var
iab
le var
iab
le P
rof
it
comb
ined w
ith phys
ica
lproduc
t(Y
) produc
t Produc
t TPP APP MPP inpu
t cos
t inpu
t cos
t inpu
t cos
t
m
ixedresou
rces (TPP
) (APP
) (MPP
) (TPPx1
) (APPx1
) (MPPx1
) (X1
x5) (APPx5
) (MPPx5
) (
TVP-TV
IC)

0 0 0 0 0
0
.5 0
.5 0
1 0
.5 0
.5 0
.5 0
.5 5 2
.5 -4
.5
1
.5 1
.5 7
.5
2 2 1 2 1 10 5 -8
3 3 15
3 5 1
.67 5 1
.67 15 8
.35 -10
7 7 35
4 12 3 12 3 20 15 -8
11 11 55
5 23 4
.6 23 4
.6 25 23 -2
16 16 80
6 39 6
.5 39 6
.5 30 32
.5 9
13 13 65
7 52 7
.43 52 7
.43 35 37
.15 17
8 8 40
8 60 7
.5 60 7
.5 40 37
.5 20
CHAPTER 8

5 5 25
9 65 7
.22 65 7
.22 45 36
.1 20
3 3 15
10 68 6
.8 68 6
.8 50 34 18
1 1 5
11 69 6
.27 69 6
.27 55 31
.35 14
Econom

-1 -1 -5
12 68 5
.67 68 5
.67 60 28
.35 8
ics o

-3 -3 -15
13 65 5 65 5 65 25 0
-5 -5 -25
14 60 4
.29 60 4
.29 60
f Feed
ing

Sou
rce
: Shang 1990
215
216 CHAPTER8 NUTR
ITIONIN TROP
ICAL AQUACULTURE

F
igure 8
.2
Re
lat
ionsh
ip be
tween TVP
, TV
IC and p
rof
it
.

Theto ta
lva lue o
fthef ish does not only considerthe weigh t buta lso
the overal
l quali
tyincludingf reshness ofthef ish. The va lue a t
t r
ibuted
to qual
ityis accountedinthe ma rketp r
ice ofthef ish
. Thesefac tors are
includedinthe pro f
itindex. Thefarmer can then compare the pro fit
index ofva r
iousfeeds when choos ingfeedsfo rh is s
tock.
A thi
rdind icatorof determiningthe econom ic eff
iciency offeedsis
the returns onfeeds . Th is ind icator shows the ra te of return on
investmen t onfeeds. Thisis measu red bythefo l
lowingfo rmu la;

ne
tp ro
fit
Re
turns onfeeds =
costoffeeds

Tab le 8.4 is a cos t and re turns computa t

ions tha t shows the
compa rativeeconom ice ff
icienc
ies based onthethreeindicato
rs(incidence
cost, profitindex, and re turn onfeeds )in the pen cu l
ture of As ian
catf
ish(C larias mac rocephalus)fed three di
ffe
rent types ofdiets
. Us ing
the aforemen tionedfo rmulas, the summary ofthe econom icind ica
tors
usingtheth ree die
tsis :

Die
t 3 and D
iet 2 havelowerinc
idence cos
ts,higherpro
fitindexes
,
and returns onfeeds comparedto D
iet1. We conc
ludetha tthe use of
Die
ts 3 and 2 are economical
ly v
iab
le. The use ofD ie
t 1is alosing
proposi
tion.
 CHAPTER8 Econom
ics o
f Feed
ing 217
T
able8
.4Costandre
turnsofthepencul
tureo
fClarias macrocephalus a
tas tock
ingdens
ityo
f 10f
ish/m2and
fedthreed
if
feren
tdietsfo
r120days
.Va
luesa reonape rhape rcropbasisinPh
il
ipp
inePeso(PhP)*
Quant
ity Un
itP
rice
I
tems (kg
) D
iet1 D
iet2 D
iet3
Revenue

D
iet 1(30
.2 g MBW) 2208 120 264960
Die
t 2(58
.3 g MBW) 4696 120 563520
Die
t 3(67
.35 g MBW) 4808 120 576960

Less
:
Var
iab
le cos
ts
F
inge
rlings (pcs
) 100000 1
.5 150000 150000 150000
Feeds(kg
) D
iet1 6308 14
.69 92665
Die
t2 9979 15
.4 153677
Die
t3 21964 7 153748

L
ime (
ton
) 1 1000 1000 1000 1000
ch
icken manu
re (
ton
) 1 800 800 800 800
45-0
-0 (bag
) 0
.5 415 207
.5 207
.5 207
.5
16
-20
-0 (bag
) 1 410 410 410 410
D
iese
lfue
l(l
) 100 15
.15 1515 1515 1515
M
isce
llaneous expenses 5299
.2 11270
.4 11539
.2
Repa
ir and ma
intenance 598
.8 598
.8 598
.8
Care
taker
’s sa
lary (mo
) 5 1000 5000 5000 5000
Pondren
t(y
r) 0
.5 8000 4000 4000 4000
Sub
tota
l 261495 328478 328819

F
ixed cos
t
Deprec
iat
ion 2649 12849 12849
In
teres
t expenses 12849 15595 15598
Sub
tota
l 15498 28444 28447
To
tal cos
ts 276993 356922 357266

Ne
t prof
it be
foretax
/crop -12033 206598 219694

to
tal cos
toffeed used
Inc
idencecos
t = = 41
.97 32
.73 31
.98
w
tofthef
ish p
roduced

to
talva
lue o
ffish-to
talcos
toffeeds
Prof
it index = = 1
.86 2
.67 2
.75
to
tal cos
toffeeds

ne
tpro
fit -12
.99% 134
.44% 142
.89%
Re
turns on feeds = =
cos
toffeeds

Sou
rce
: Con
iza e
tal
.2001

Leas
t-cos
t Comb
ina
tiono
fFeeds

Cons
ideringthe sensitiv
ity ofthe costo ffeedsinthe profitab
ili
ty offish
farming
, keepingthecos t downisc rucialtoensu retheeconom icviabil
ity
ofthefishfarm ing enterprise
. Least-cost combina t
ion techniques are
usedin determ iningthelowes t costf rom di
fferentfeed comb inations
that would resultinthe same p roduction.
218 CHAPTER 8 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Forexamp le,twotypes o ffeeds(F 1and F2 )are so

ld a
td i
fferentp r
ices
and are available tothefishfarme r .Ifthe two types offeeds can be
subst
ituted,thera te ofsubs t
itutionind icatesthe amoun t by which one
feed must be changedin o rderto offsetthe changeinthe amoun tofthe
otherfeed. Thisis called margina l rate o
f substi
tu t
ion(MRS ) between
feeds
, the value of whichis nega t
ive.

amountoffeed rep
laced(FJ
MRS =
amountoffeed added(F2
)

The MRS betweeninpu ts(

feeds)is a phys
ica
lre lat
ionsh ip and canno
t
de
terminetheleas t-cos
t combination o
ffeeds. Feed prices(prices o
f PF1
and PF2
)a re needed andthera t
iois compa red with MRS .

pr
ice offeed added(PF2
)
P
rice ra
tio =
p
rice o
ffeed replaced(PF
)1

Theleas
t-cos
t comb
ina t
ion o
ffeeds F1and F2occu
rs whenthe MRS
is equa
ltotheinverse o
fthe price ra
tio
.

F1= pf2 o
r F1
(PF1
)= F2
(PF2
)
F2 PF1

The costofchangein onefeedis equa ltothe costo fchangeinthe

otherfeed.In Table 8
.5,theleast cos
t combina
tion offeedsis a
t P180,
whichcorresponds w
iththe useo
f 10pa r
ts o
fF2and 15pa rtsof F1
.Inthis
connec
tion,thecos tofchangein F1isequaltothe cos
to fchangein F2
.

Tab
le8
.5 Hypo
the
tica
lre
lat
ionsh
ipfo
rcomb
iningfeedsto p
roduceag
ivenleve
lofou
tpu
t
(P
1=P
9,P
2=P
6)

X2 X
1 F1
/F2 P2
/P1 Cos
t
(MRS*
) (F1xP1+F2xP2
)

0 40 240
4 24 0
.25 0
.67 180
10 15 0
.67 0
.67 180
15 8 0
.71 0
.67 183
20 3 1
.00 0
.67 198
25 0 1
.67 0
.67 225

*Ma
rgina
lra
te o
f subs
titu
tion
X1
isthefeed be
ing rep
laced
X2
isthefeed be
ing added
F1
isthe changeinthe quan
tity o
f X1
F2isthe changeinthe quan
tity o
f X2

Sou
rce
: Shang1990
 CHAPTER8 Econom
ics o
f Feed
ing 219
M
inimum Cos
tofFeedFo
rmu
lat
ionus
ingL
inea
rProg
ramm
ing

Linear programm ingis a compu tat

iona l method usedto a l
loca te scarce
resourcesto max im ize p
rofito r m inimize cost
.In aquacu ltu re,itis often
used in the m inimization mode lo f comput ing the leas t-cost feed
comb ina t
ions o rra tion and s ti
ll mee ttherequ ired nutr
ien tsfo rthef ish.
The concep tsinvo lvedinl inea r programm inginclude:
□ Ob jective- usua llyto max imize p rof
its or minimize cost;
□ Cons tra ints- resources such as rawfeeding redients a re res
tricted
orl imi ted;
□ A lterna tive ways o fa t
ta iningthe ob jec
tive are dete
rm ined ;
□ Re la t
ionsh ip betweeninpu t and ou tputis assumedto bel inear;
□ P rices pa id o rreceived a re assumed cons tant; and
□ Quan t
i ty ofinpu ts used shou ld be equalto o rlessthanthe quan ti
ty
ava i
lab le.

Table 8.6 presents a hypothetical example tha

tillustra
testhe da ta
on nutrient avai
labi
lity
, cost
, requ irements, ob
ject
ives, and cons
traints
to cons ider in comput ing for leas t cos
t comb ination using linear
programm ing as adoptedfrom Shang 1990 .

Tab
le8
.6 Da
taonnu t
rien
tava
ilabi
li
tyandrequ
iremen
ts,feedcos
t, ob
jec
tive
,and
cons
tra
intsinl
inea
rp rogramm
ing
M
inimum da
ily
Nu
trien
t Feed A Feed B requ
iremen
t

A Nu
trien
t ava
ilab
il
ity and requ
iremen
ts
(Un
ito
f nu
trien
t pe
r un
ito
ffeed
)

Ca
lcium 2 1 18
Pro
tein 2 2 20
Ca
lor
ies 1 5 25

B
. Cos
t pe
r un
ito
ffeed 1
.00 2
.00

C. Ob
jec
tive
: M
inim
ize cos
t= p1
a +p2
b

D. Sub
jec
tto cons
tra
ints such as
: 2A + 1B => 18
2A + 2B => 20
1A + 5B => 25
A=>0
, B=>0

whe
re:A
,isthe amoun
tof Feed A
, and
B
,isthe amoun
tof Feed B
.

Sou
rce
: Shang 1990

Theconstrain
tsintheequa t
ionsinTab le 8.5insuretha tthe minimum
requiremen tforingredientis me ta ttheleas t cost
. For examp le, the
minimum un i
ts of calc
ium , protein and ca lor
ies are 18, 20, and 25
respect
ively
.S implelinearp rogramm ing can be solved graphica
lly
. Bu t
as the problem becomes more comp licated, systema t
ic compu tat
iona l
techniques using compu te
rs are required.
220 CHAPTER 8 NUTR
ITIONIN TROP
ICAL AQUACULTURE

Summa
ry

The concep ts and approaches on the eva lua t

ion and compu tation o f
selectedind icato rs ofthe econom ic e ff
iciency o f feed ingin aquacu lture
opera tions a re discussed . The d iscuss ion s ta r
ted w iththeiden ti
fica tion
ofcos ti temsinvo lvedin p roduc ing aquacu lturefeeds . These cos ti tems
are class if
iedin to d i
rec t cos ts andind irect cos ts. To tal costs and pe r
unit cos ts are use ful econom icind ica tors to gu ide the aquacu lture
farme rsinthe ir opera t
ions .
Th is chapter introduced the produc tion func tion approach for
evalua tingthere lationsh ip betweenfa rminpu tssuchasfeeds , andou tpu t
(f
ish ). Quant i tative examp les demons trated marg inal ana lysis tha t
providedbas isfo runde rstand ingtheva r
iouss tagesinproduc t
ionsys tems .
These a rethe s tages whe re pro fi
tisinc reasing , max imum ,o r decreas ing.
The concep ts o f max imum sus tainab le y ield (MSY ) and max imum
econom ic yield(MEY ) we re discussedin o rde rto h ighlightthe op timum
feedingleve ltha tw il
l resu lttothe h ighes t pro fitlevel.
Th is chap ter a lso presen ted ind ices for measur ing the econom ic
ef
ficiencyin p roduc ingfeeds . These ind icesinc ludeinc idence cos t and
profi
tindex . Theleas t-cost app roach was d iscussedto demons tratethe
method o f de term ining comb ina tions o fingred ien ts ofd i
fferentfeed
formu la tions g iventhe preva iling p r
ices .
Fina lly, the chap terin troduced the concep to fl inear programm ing
in de te rm iningthe m in imumfeed cos ttha t wou ldy ieldthe des iredleve ls
of nu trien ts requ ired bythef ish .

Gu
ideQues
tions

1
. Wha
titems comp
risethe d
irec
t cos
ts o
fproduc
ing sh
rimpfeeds?
2
. Wha
titems comp
risetheind
irec
t cos
ts o
fproduc
ing sh
rimpfeeds?
3
. Inte
rms o
fyie
ldleve
ls,a
t wha
t po
intis add
itiona
lfeed
ing nolonge
r
bene
fic
ialto thef
ishfarmer? Wha
tis the mos
t pro
fitab
leleve
lof
feed
ingthef
ish? Exp
lain us
ing concep
ts o
f ma
rgina
l ana
lys
is.
4
. How do you measure the econom
ic e
ffic
ienc
ies o
fdi
fferen
t feeds
ava
ilab
letothef
ishfa
rme
r?
5
. Wha
tisl
inear p
rog
ramm
ing?
6
. Wha
tarethe concep
ts and requ
iremen
tsinl
inea
rprog
ramm
ing and
ana
lys
is o
ffeedfo
rmu
lat
ions?

Sugges
ted Read
ings

Agbayani RF, Bal

iao DD , Samonte GPB, Tumal
iuan RE
, Caturao RD.
1991
. Econom ic analys
is o
fthe monocul
ture o
f mudc
rab (Scy
lla
se
rrata) Forsskal
. Aquaculture 91
:223
-231.

Con
iza E, Tan-Fe
rm in J
, Catacutan M. 2001. Pencultu
re o
fAsian cat
fish
Clarias macrocephalus fedfour dif
ferentd ie
tsinthe Phi
lippines
.
Paper presen ted in the As
ian F isheries Forum. Kaohsh iung,
Taiwan. Novembe r 26-29
, 2001.
 CHAPTER 8 Econom
ics o
f Feed
ing 2
21
Ha
tch U, AgbayaniR
, Be
lleza E
. 1996. Econom
ic analysis o
fprawn(P.
monodon)inthe Phil
ipp
ines
.I I
: Grow-ou
t operat
ions. As
ian F
ish
.
Sc
i.9 :127
-141
.

M
illamena OM , Trino AT. 1994 . Evaluation off
ish protein concentrate
andlac tic yeas
t as p ro
tein sourcesfor shrimp Penaeus monodon .
In:P roceedings of the Th i
rd Asian Fisher
ies Fo rum, Oct. 26-30,
1992
.S ingapore . p 675-678.

New MB. 1986. Formulated aquacu lturefeedsin Asia: somethough ts on

comparat ive econom ics, indus tr
ial po tent
ial, prob lems , and
research needsinre lationtothe sma l
l-scalefarme r
. Repo rtofthe
Workshopon Shr imp and Fin f
ish Feed Developmen t
.Joho re Bahru,
Malays
ia. 25-29 Oct. 1986. ASEAN/UNDP/FAO/ . Reg ional Sma l
l-
scale Coasta
lF isher
ies Deve lopmen t
. RAS/84/016 . p 19 -30.

Shang YC
. 1990 . Aquacul
ture Economic Ana
lys
is: AnIntroduct
ion
. The
World Aquacu l
ture Soc
iety. Louis
iana State Un
iversity Baton
Rouge, LA 70803.

Sumagaysay NS, BorlonganIG. 1994. Growth and product

ion ofmilkf
ish
(Chanos chanos)in brackishwater ponds:effec
ts o
fd ie
tary pro
tein
andfeedingleve ls
. Aquaculture 132:273-283.

Sumagaysay NS
. 1998. M i
lkf
ish(Chanos chanos) production and water
qual
ityin brack
ishwater ponds a
tdi
ffe
rentlevels andfrequencies
.
J
. Appl.Ichthyo
l. 14
:81-85.

V
incke MM. 1969. Compte-rendu d ’activ
ite annee. Division des
Recherches Pisc
icoles
, Centre Technique Fores
tier Tropica
l,
Tananar
ive, Madagascar
, 30 p
.
 Append
ix A AP
-1

Samp
le Wo
rkshee
tfo
r
Ca
lcu
lat
ingtheNu
tr
ien
tComposi
tiono
fFeed
Feedformu
lat
ioncode :________
Datecomputed:______________
Formula
tedby: ______________

Ing
red
ien
ts g
/100g P
rote
in% Fa
t% F
ibe
r% Ash% NFE%
(1
) (2) (3) (4
) (5) (6) (7
)
2-AP Append
ix B

1
. Me
thodo
fpro
teinana
lys
is(K
jeldah
l me
thod
)

P
roc
edu
re:
1
. P
re-hea
tdiges
torto400
°Cin
sideahood
.

2
. We
igh60
-70 mgd
rysamp
leinata
redf
i
lte
rpape
randp
laceind
ige
st
iontube
.

3
. Re
cordthe we
igh
tofthesamp
le. Ma
ke2rep
l
ica
tes
/samp
le.

4
. Addca
talys
tand5 m
lconcen
tra
tedsu
lfu
rica
cidtoea
cho
fthed
ige
st
iontube
s.

5
. D
ige
stthesamp
leun
t
iltheso
lut
ioni
sclea
r(1h
).

6
. Coo
lthed
iges
tandadd25 m
lammon
ia-
freed
is
ti
l
led wa
ter
.

7
. D
is
ti
l
landt
i
tra
te.

8
. Compu
tefo
r %c
rudep
rote
in.

Com
puta
tion
:
%c
rude p
rote
in = % n
itrogen x fac
tor(6
.25
)

Spec
if
icconve
rsionfac
torfo
r pe
rcen
tcrude p
rote
in va
ries acco
rdingtotypes o
fproduc
ts asfo
llows
:
Mea
t(an
ima
lorf
ish
) 6
.25
M
ilk and da
iry p
roduc
ts 6
.38
Ce
rea
lpro
teins 5
.90
F
lou
r 5
.70
O
il seed p
rote
ins 5
.40

2.C
rudefa
tana
lys
is(Soxh
tec me
thod
)

P
rocedu
re:
1
. We
igh ou
ttwo(2
)rep
lica
tes(abou
t 1g
)fo
r each d
ry samp
lein c
rudef
il
ter pape
r.

2
. P
lacein
to c
ruc
ibles
. P
lace c
ruc
iblesins
idethe p
re-we
ighed ex
trac
tion cups w
ith 5 p
ieces o
f
beads
.

3
. Pos
it
ionthe c
ruc
iblesinthe ex
trac
tion un
it.

4
. P
lace abou
t 50 m
lofe
the
rins
ide each cup
. Inse
rt each cupin
tothe un
it andlockf
irm
ly.

5
. Ex
trac
tfo
r 30 m
inu
tes(ex
trac
tion knobsin“bo
il
ing
” pos
it
ion
).

6
. R
insefo
r 30 m
inu
tes(ex
trac
tion knobsin“
rins
ing
” pos
it
ion
).

7
. Recove
r so
lven
t(15
-30 m
inu
tes
). Condenser va
lves a
re c
losed
.

8
. Remove ex
trac
tion cupsf
rom ex
trac
tion un
it. Rese
rve de
fat
ted samp
lesfo
rcrudef
iber
ana
lys
is.

9
. D
rythe cups(w
ith c
rudefa
t)in an ovenfor 10 m
inu
tes
.

10
. P
lacethe cupsin a dess
ica
tor and coo
lfor 30 m
inu
tes
.

11
. We
ighto cons
tan
t we
igh
t.
 Append
ix B AP
-3
12
. Compu
tefor % c
rudefa
t

Compu
tat
ion
:
%c t= w
rudefa toffa
t(cor
rected
) x 100
wtofsample

3.L
ipidex
trac
tion(B
lighand Dye
r me
thod
)

P
rocedu
re:
1
. We
igh 1 g we
tsamp
les o
r0.5 g d
ry samp
le.

2
. Homogen
izefor 2 m
inin a so
lven
tmix
:(40 m
l CH3OH
, 20 m
l CHC
I3, 16 m
l H2O)
.

3
. Add 20 m
l CHC
I3and homogen
izefo
r2 m
in.

4
.Fi
lte
rth
rough ce
lite
-Rf
il
ter and co
llec
tthef
il
tra
te, sc
rape o
ffthe uppe
rlaye
rofce
lite pape
r and
t
rans
ferin
tothe homogen
izingtube
.

5
. M
ix w
ith new quan
tity o
fso
lven
tmix and homogen
izefo
r2 m
in andf
il
terth
rough ce
li
tef
il
ter
.

6
.Rinsethe ce
litef
il
ter w
ith 40 m
l CHC
I3andt
rans
ferthe comb
inedf
il
tered ex
trac
tin asepa
rato
ry
funne
l.

7
. To b
reakthe emu
lsion
, add 1 m
lof a 40 m
l H2Oin wh
ich a p
inch o
f NaC
I was p
rev
ious
ly
d
isso
lved
.

8
. Shakethe sepa
ratoryfunne
lfor abou
t2 m
in andle
tstandfor 10 m
into a
llow sepa
rat
ion
.
Upperlaye
r = CH3OH & H2O
Lowerlaye
r = CHC
I3 &l
ipid

9
. Dra
in o
ffthelowe
rlaye
rin
to af
lask pass
ingth
roughthe wa
terf
ree Na2SO4f
il
ter
.

10
. R
insethe Na2SO4
fil
ter w
ith CHC
l3and evapo
ratethe co
llec
tedl
ipidf
rac
tion us
ing aro
tary
evapora
tor un
ti
l nea
rly d
ry andf
lush ou
tthe rema
ining so
lven
ts w
ith n
itrogen (
tempe
ratu
re o
f
evapo
rat
ion shou
ld no
t exceed 40°C
).

11
. De
term
inethe quan
tity o
ftheto
tall
ipidsin g
/100g d
ry w
t.
4-AP Append
ix B

4
.Sapon
if
ica
tionandt
ranses
ter
if
ica
tion

Sapon
if
ica
tion

P
rocedu
re:
1
. T
rans
ferthel
ipids(4 m
l benzene
)in a py
rexcen
tri
fuge g
lass
-tube w
ithte
flonl
ined cap
;rinsethe
pea
r-shapedf
lask w
ith 1 m
l mo
re benzene
.

2
. Add 1 m
ldis
til
led wa
ter and 10 m
l0.5 N KOH
/CH3OHtothel
ipid ex
trac
tinthe cen
tri
fuge g
lass
tube
.

3
. F
lushthetube w
ith n
itrogen
.

4
. C
lose we
ll and shake
.

5
. Sapon
ifyin a bo
il
ing wa
ter ba
th (100°C) and shake we
ll eve
ry 10 m
in (
tota
lof 40 m
in)
. Coo
l
downthe cen
tri
fuge g
lasstube
.

6
. Add 5 m
l 2M HC
I and 5 m
l pe
tro
leum benzene
. Shake we
ll
.

7
. Le
tstandfor 5 m
inu
tes o
r un
ti
ltwo d
ist
inc
tlaye
rs a
refo
rmed
.

8
. T
rans
ferthe benzene phaseto a pea
r-shapedf
lask w
ith a p
ipe
tte(ge
tthe uppe
r pa
rt)
.

9
. D
ist
il
lof
fthe so
lven
t us
ing vacuum ro
tary evapo
rato
r.

Es
ter
if
ica
tion
1
. Add 2 x 1 m
l CH3OH andt
rans
ferthefa
tty ac
idsin
to a sc
rew-capped g
lasstube
.

2
. Add 2 x 1 m
l benzene andt
rans
fertherema
iningfa
tty ac
idstothe g
lasstube
.

3
. Add 3 m
lof 14% BF3
-CH3OH
,flush w
ith n
itrogen
,close we
ll
, and shake
.

3
. P
lacethe sc
rew-capped g
lasstubein a bo
il
ing wa
ter ba
thfor 8 m
in and shake a
fte
r4
m
inu
tes
.

5
. Coo
l downthetube
, add 5 m
l H2O(
to s
topthereac
tion
) and 5 m
l pe
tro
leum benzene
.

6
. Le
tstandfor 5 m
in andt
rans
ferthe pe
tro
leum benzene phase(uppe
rlaye
r)to a pea
r-
shapedf
lask
.

7
. D
ist
il
lof
fthe so
lven
t us
ing ro
tary evapo
rato
r.

8
. D
isso
lvethefa
tty ac
id mehy
l es
ter(FAME
)in 1 m
lisooc
tane andt
rans
ferto an ambe
rvia
lwi
th a
sc
rew cap andte
flon
-faced s
il
icone sep
tal
ine
r.

9
. Samp
leisreadyfo
r GLC (gasl
iqu
id ch
roma
tog
raphy
)in
jec
tion
.
 Append
ix B AP
-5
5
. Me
thodo
fpe
rox
ideva
luede
term
ina
tion

P
rocedu
re:

1
. P
lace 0
.3 gfa
t samp
lein 250 m
lflask w
ith s
toppe
r; add 10 m
lof CHC
I3-CH3COOH
shake
.

2
. Add 1 m
lofsa
tura
ted K
l so
lut
ion
.

3
. S
toppe
r and a
llowto s
tandinthe da
rkfo
r5 m
in.

4
. Add 20 m
ldis
til
led wa
ter
, shake
.

5
. T
itra
te w
ith 0
.01 N Na2S2O3so
lut
ion (un
ti
lligh
t ye
llow co
lor appea
rs)
.

6
. Add 1 m
lof1
.5% s
tarch so
lut
ion (un
ti
l co
lor
less
).

Ca
lcu
lat
ion
:
PV = (Vs- Vb
) xfx 10
/w
whe
re;Vs = t
itra
tion vo
lume o
f samp
le
Vb = t
itra
tion vo
lume o
fblank
F = fac
tor o
f0.01N Na2S2O3so
ln.
W = we
igh
toffa
tin vo
lume o
f ex
trac
t used(g
)
N = no
rma
lity o
f Na2S2O3so
lut
ion (
inth
is case N
/100
)

6. P
rocedu
reo
ffa
ttyac
idva
luede
term
ina
tion

F
reefa
ttyac
idva
lue(FFA
)
Ca
lcu
lat
ion
:

FFA(%) = ac
id va
lue x mo
lwto
fole
ic ac
id/mo
lwto
f KOH x 100
/1000
= ac
id va
lue x 282
.27
/56
.11 x 1
/10
= ac
id va
lue x ½

7
.Th
ioba
rbi
tur
icac
id(TBA
)va
luede
term
ina
tion

P
rocedu
re
1
.Place 0
.2-0
.4 gfa
t samp
lein ates
ttube w
ith a sc
rew cap
.

2
. Add 3 d
rops o
f an
tiox
idan
t so
lut
ion
.

3
. Removethe so
lven
t us
ingthe ro
tary evapora
tor unde
rreduced p
ressu
re a
t 35
-40°C
.

4
. Add 3 m
l TBA so
lut
ion
. And 17 m
loft
rich
loroace
tic ac
id so
lut
ion(TCA
).

5
.Flush N2gasin
tothetes
ttube andimmed
iate
ly s
toppe
r.

6
. Hea
tat 100°Cin a bo
il
ing wa
ter ba
thfor 30 m
in un
ti
lthe co
lou
r appea
rs.
6-AP Append
ix B

7
. Coo
ltoroomtempe
ratu
re w
ithtap wa
ter
.

8
. Add abou
t5 m
l ch
loro
form and m
ixfo
r afew seconds w
ith a Vo
rtex m
ixe
r.

9
.Trans
fer abou
t 15 m
lofthe co
lor so
lut
ionto a g
lass cen
tri
fug
ingtube
.

10
. Cen
tri
fugefo
r 10 m
in a
t3,000rpm
.

11
.Ifthe aqueous so
lut
ionis no
tclea
r, cen
tri
fuge aga
in a
t 10
,000rpmfo
r 10 m
in.

12
.Trans
fer a pa
rto
fthe c
lea
r aqueous so
lut
ion andread abso
rbance a
t 532 nm
.

13
.Blanktes
tshou
ld be ca
rried ou
tinthe same manne
rwi
thou
tfa
ts.

Ca
lcu
lat
ion
:
TBA(mg ma
lona
ldehyde
/kgfa
t) = abs xfx 0
.2/w

whe
re: abs = abso
rbance a
t 532 nm
w = we
igh
toffa
tinvo
lume o
f ex
trac
t(g
)
f =fac
tor = 46
Glossa
ry

A
Acc
limatiza
tion
: adap
tat
iono
rincrea
sed A
lga
ld ie
-of
f: anab
rup
t mass
ive mo
rta
l
ity Anox
ic: devo
ido
fox
ygen
to
leranceofaspec
iestoachanged o
funicel
lu
lara
lgaeresu
lt
ingf
rom natu
ral
env
ironment o
r man-madecause Ant
inut
rit
iona
lfac
tors
: subs
tancesinthe
feeds
tuf
f wh
ichcanredu
cenutr
it
iona
l
Ac
id:acompoundtha t whendisso
lvedin Algebra
icequation: a ma
thema
tica
l va
lue
wa
ter
,dissoc
iatestoy
ieldhydrogenions methodusedinformulat
ingfeed
su s
ing
(H+
) onlyfewingred
ients Ant
iox
idant
: astrongreduc
ingagent
,
wh
ichiseas
ilyox
idizedandthuspreven
ts
Activet ransport: movemen tofa Al
imen ta
rycanal
: atubula
rpa ssagetha
t theox
ida
tionofothersubs
tances
moleculeac rossa memb raneorothe
r ex
tendsfromthe mouthtoanusand
barrierdrivenbyene rgyo the
rthanthat func
tionindiges
t
ionandab so
rpt
ionof Appe
ti
te: na
tura
lde
siretosa
tis
fyhunge
r
storedintheconcen tra
tiongrad
ientor foodande l
imina
tionofres
idual was
te
e
lec trochemicalgrad ientofthe Aquacu lture: thefa rm
ingo faquatic
transpo rted molecule A
lkal
ini
ty:(
tota
l)thetota
lconcentra
tionof organism s
,inc ludingfish,crus
taceans ,
base
sin wa
terexpressedasmil
ligrams mol
lu sk
s,andaqua ticplants
. Farming
Adl
ibi
tumfeed
ing
:p rov
iding un
l
imi
ted pe
rli
terofequ
ivalen
tcalc
iumca rbonate imp l
iessomefo rmo finterven
tioninthe
amoun
toffeedun
t
ilsat
iat
ion (CaCO3) rearingp ro ce
sstoenhancep rodu ct
ion
,
sucha s
,regu lars tock
ing,feeding
,and
Addit
ive: aningrediento
rcomb inationof Aminoac id: acarboxyl
ica c
idtha
t protect
ionf romp reda to
rs
ingred
ients
,o therthanpremix,addedto includesanam inogroupaspa r
tofi
ts
thebasicfeed mixo rpar
tsthereoftofulf
i
ll st
ructure;anyoneclassoforgan
ic Aqua
farm: natura
land/o
rrecon
struc
ted
aspecif
icneed ;usuallyusedin micro­ compounds whichcontainboththeamino bodyof wa
terin
tendedfo
raquacul
ture
quant
it
ies andrequiresca re
ful handl
ing (NH2 )groupandthecarboxy
l(COOH )
and mix
ing group Aquafeeds: feedstha
tarein
tendedfo
r
aquacul
turespecies
Adiposet
issue: connect
ivet
issue Aminoacidan tagon
ism : occurs when
specia
l
izedforthestorageofla
rge someam inoacidsarefedine xcessof Arachidonicac
id: a20
-carbon
amountsoftr
iglyce
rides requi
redleve
lscau s
inganin c
rea sein unsatura
tedfat
tyac
idhavingfou
rdoub
le
requi
rementforanotheram inoacidof bonds
Aeration: a mechanicalp rocessusua l
ly s
imilars
truc
ture,e.g
.a rg
inine-
lys
ine
bybubb lingai
rthrough wa te
ro rsp ray
ing an
tagonism Adenosinetr
iphospha te(ATP):a
wa
te rintoai
rb y whichthele ve
lo f un
iversa
lene rgycarr
ierincel
ls;the
d
issolvedoxygeninthe wa terisraised Ammon ia(NH3
): aproduc
to ff
ishpro
tein pr
inc
ipaldriv
ingforceinene rgyrequ
ir
ing
and/orthelevelofd issolvedno xious me
tabol
ism anddecomposi
tionoforgan
ic b
iochemicalprocesseso fl
i
fe
vo
latilegasesi sredu ced ma
tte
rb ybacte
ria
ATPase: oneo falargecla
ssofenzymes
A
ldose: asimplesuga
rtha
tconta
insan Anabol
ism: me tabo
licreac
tionsresu
lt
ing tha
tca talyzeap rocesstha
tin
volvesthe
a
ldehydegroupatoneendofthecha
inas inthesynthesisof morecomplex hyd
rolysiso f ATP
pa
rtofi
tss
tructu
re compoundsfroms impleprecurso
rs;
commonlylinkedtotheh yd
rolys
isofATP At
trac
tant
: subs
tancesaddedtofeedsfor
Aldehyde: acompound w
ithan HC=0 to ADPandpho sphate fas
tconsumpt
ionespecia
l
lyforcrus
tacean
groupattachedtoanal
iphat
icca
rbon spec
ies
chain Anaerob ic
: acondit
iono rchem ical
rea c
tion wheregaseouso xygeni sno t Automat
icfeeder: adevice
,u sual
ly
Alga
lbloom : anexponent
ialinc
reasein presento rnotrequ
ired
,e.g. e
lect
ron
ical
ly ope
rated
,thatdispenses
the popu
lat
ion ofunice
llu
laralgaecaused decompos it
ionoforganic wastesb y feedatpre
-se
lectedtime
s
byhighnutr
ientloadinthe wate
r m
i c
roo rgan
ism s
,releas
ingtoxic h ydrogen
sul
fideand me thanega s
2-G G
lossary

Autotrophic:o rgan
ismstha
trequire Carbohydrate: alargeg roupoforganic Concen t
rate
: afeedu sed w
ithanotherto
carbond iox
ideo rcarbohyd
ratesorsimple compoundscommoninp lants wh
ich imp rovethenut
ri
tivebalanceand
inorganic ni
trogencompoundsfor includesimplesuga
rs,starches
, intendedtobefurthe
rd i
lutedand mixedto
metabol
icsyn thes
is;theseo
rganismscan ce
lluloses
, gumsandrelatedsubstances produceasupplemen torcomp le
tefeed
manufacturetheirownfood(e.g
.plan
ts)
Carbond iox
ide: (CO2 )aco lor
les s
, Cr
iticals tand
ingc rop: thes tand
ingcrop
Averagephysicalproduct(APP): the odorlessga s
,re sul
tingfromtheo xidat
ion orbioma ssby whichtheinc reasein
amountoff
ishp roducedorproductper ofca rbon-conta
iningsubstances,h igh
ly nutr
ien trequi
remen tcould nolonge rbe
un
itofinpu
t(feed)giventothef
ish solublein water
,toxictofishatlevels suppl
iedb ythena tura
lfood,thusg row
th
h
ighe rthan20ppm ;tox
ici
tyincreasesw i
th ratei slessthanthepo tent
ial ma x
imum.
lowle ve
lsofo xygen Th
isi sthepo int wherefeedingcan
ef
fectivelyincrea
sey ield

B Card
sho
i
r
t
fo
,f
rmteeth
: tee
ineandpoin
ththa
ted
ta renume
rous
,
Cy
ou
toso
ts
l: thepo
idethenu c
rt
ionofthece
leusando the
l
ltha
tl
r memb
ie
s
rane
-
Benth
ic: per
tain
ingtobottomter
rainof Carn
ivores
: animalsthatfeedexc
lus ly bounded o
ive rgane
lles
aquat
icreg
ions;thepar
to ftheaquat
ic onanimal ma
tte
r
env
ironmentinhabi
ted b
yo rgan
ismsthat
l
iveono

Ben
thos
rinthesed

: o
rgan
imen

ismstha
tli
t

veonorinthe
Car
ry
tha
ing capaci
ty: ma
tapondcansuppo r
x
imumb
tand ma
ioma
in
ta
s
in
s
D
g
ivenace rta
inleve
lof management,e.g
. Denovob iosyn
thes
is: syn
thesiso
f
sed
imen
tofaquat
icenvi
ronmen
ts nofer
ti
l
izat
ion,withfer
ti
l
izat
ion
, b
iomolecu
lesthatoccu
rs wi
thinthe
fer
ti
l
izat
ion plu
sfeeding an
imal body
Binder
: substancesaddedtofeedsto
makeitstab
leinthe wa
ter
,usual
lya Case
in: theco
l
loida
lpro
teinin m
i
lk
carbohydra
te Deprec
iation: thedec
lineinva
lueof
f
ixedas setsinaprodu
ctionpro
cess w
i
th
Ca
tabolism: metabo
l
icreactionsre su
l
ting thepassageo ft
ime
Biomass: inaquacu lture
,thisnormal
ly inbreakdowno fcomplex molecu
lesto
refersonlytothetota
l weigh
tofspe c
ies s
implep roduc
ts,commonlyox ida
tionto
beingcul
tured,expressedintermso fa Deprec
iat
ioncos t
: thealloca
tionofthe
ca
rbond iox
ideand water or
ig
inalcostofal
lfixedas
setsoverthei
r
g
ivena reao rvolumeo fthehabi
tat
economicl
ifespan
Ce
llu
lose: apo lymero
fgluco
se,an
Biosynthesis
: thefo
rmationofchem
ica
l impor
tantst
ructura
l ma
ter
ia
linplant
s;
substancesfromothe
rchem ica
l Desmosome : specia
l
izedce l
l-cel
l
ma
jorst
ructu
ralcomponentofplan
tcell junct
ion, usual
lyformedbe tweentwo
substancesinaliv
ingorganism wa
l
l epi
thel
ialcells
,cha rac
ter
ized b yden se
Broodstock: capt
ivepa rentstocki
s p
laqueso fp rote
ininto whichintermedia
te
Ch
it
in: ma
jorst
ruc
tura
lcomponentofthe f
ilamentsinthetwoad joiningcellsinse
rt
a
llowedtorea chse xua
l ma tu
ri
tyand r
ig
idexoske
leton o
finve
rteb
rates
spawningreadinessto ma keegg sand
sperm avai
lab
le De
ter
iora
tion
: theac
t
iono
fgradua
l
Choles
tero
l: aphysio
log
ical
lyimpo
rtan
t impa
irment
s
terolandiswidesp
readinthe
Brushbo rde
r: densecove ringof b
iomemb rane
m
icrov
il
liontheap icalsur
faceo fepi
thel
ia
l Det
ri
tivo
res: an
ima
lstha
tfeedon
cel
lsintheintes
tineandk idney
;the decay
ing ma
tte
r
Coenzyme : anonprote
insubstancetha
t
m
i c
rov
il
liaidinabso rpt
ion byinc
reasing takespartinanenzyma t
icreact
ionandis
thesurfaceareao fthecel
l De
tri
tus
: fragmen tsorremainsoforgan
ic
regenera
teda ttheendofthereact
ion;a ma
tte
rorotherdisinteg
rated ma
ter
ia
l
partne
rrequi
redb ysomeenz ymesto mo
vedaboutb y water

C produceenzyma

Cofac
tor: anino
t
ica

rgan
ct
iv
ity

icionorcoen
zyme
Docosahexaeno
carbonunsa
tu
icac
ra
id
tedfa
t
: (DHA
tyac
) a22
idhav
-
ingsix
Canineteeth: dog -
toothlike,oftenqu
i
te requi
redfo
renzymaticacti
vi
ty doublebonds
,anessent
ialfa
ttya c
idin
fang-
like;theyaree longatedand f
ish
subconical
,stra
ightcu rvedanda re Comp le
ted iet: feedtha
tcon tainsa
llthe
adaptedforp ie
rcing and holding essent
ial nut
rien
ts(prote
in,l
ipid, Dia
tom : asing
le-ce
lled p
lant
carbohydrate
,v i
tamins
,m inerals)requ
ired (phytop
lankton
)co vered w
ithtwo
Ca
lorie: aunitofheatorenergy;the bythean imalfor main
tenanceandg rowth over
lapping po
rousshe l
lsofsil
ica
amounto fheatrequi
redtoraise1go f
wa
terto1 °C
. Nutri
t
ional
ly
,thek calis Compoundfeed
:Afeedcomposedo
f D
ie
l: 24
-hou
r pe
riod
sometimesu sed; 1kca
l=1000cal, seve
raling
red
ien
ts
1cal=4.186joules,1joule=0
.239cal Die
t: foodregu
lar
lyp
rov
idedand
consumed
Cann
iba
lism
: ea
tingf
lesho
fone
’sown
k
ind
 Glossary G
-3
Diges
tib
il
ity: (apparen
t) the pe
rcentage
o
fthefeeds
trac
ba
ttha
ti
tuf
ftakenin
sabso
sedonthed i
f
tothed
rbedin
fe
iges
tothebod
rencebe
t
y
ive
.I
tweenthe
tis E F
amountoffeedingestedandtheamoun t Ecdysis
: thea c
to f mo
lt
ingacut
icula
r Factorsofproduc t
ion:the ma
in
inthefeces laye
r,e.g.inc
rustaceansandin
sects categor
iesofinputsusedinap roduc
t
ion
process
, namely
:land,labo
r,capi
taland
Digest
ibil
ity: (t
rue)describestheport
ion Econom iceffic
iency : afinanc
ialo r managemen t
o
fthefeedtha tisabsorbed minusthe moneta
ry-basedc ri
ter
iafo reva luat
ingthe
mater
ialsthatarelos
tb ythegu tinthe Fat:t riacylg
lycero
ls,esterso fgly
cero
lw i
th
des
irab
il
ityo fap roductionp rocesso rthe
processo finges
tion anddigest
ion threefa t
tya cids
;fatsaregene ra
lly
ut
i
li
tyofaninpu t.Itisoften mea sureda sa
consideredtobethoset riacylg
lycero
lsthat
rat
iooftheco s
tandthey ie
ldasso cia
ted areso l
ida troomtempe rature
, whereas
Digest
ibili
tycoeff
icient: va
lues toapa rt
icularprodu c
tionp rocesso rinpu
t
expressedinpe rcentagetha
tg ive
s o
ilsa retriacylg
lyce
rolstha tarel
iquidat
informationastotheava i
lab
il
ityofspeci
fic roomtempe ratu
re
Ene
rgy
: theab
i
li
tyo
rcapac
itytodo wo
rk
nutr
ientsinthefeedo rfeedingredien
ts
Fat
tyac id:a lipha t
icca rboxyl
icacids(i
.e.
Eff
luent: wate
rtha
tisdischa
rgedf
roma w
itha-COOHg roup);the me tabol
ica
l
ly
Digest
ibleene
rgy: tha
tpar
toftheg
ross
tank, pond
,aquacul
turefarmorpowe
r importantfattyac idsha vebetween2and
energyofafeed wh
ichdoe
snotappearin
sta
tion 24carbona toms(a lwaysane vennumber)
thefeces
and maybecomp letelysaturatedorhave
Endogenous
: o
rig
ina
ting w
ith
inthebod
y one(mono -unsa tura
ted)o r mo re
Digest
ion: processofbreakingdown
nutr
ientsin
tosimplechem icalcompounds (po
lyunsa turated ) C=Cdoub le bondsin
Endop lasm icre ticulum : extens ive thecarboncha in
throughtheactionofenzyme s; the
netlikelab yr
intho fb ran chingtubu lesand
breakdown,intheal
imentarytract,of
f
la ttenedsa csinthecy to solandse rvesas Fecund i
ty : thenumbe rofegg spe runi
t
complex organ
icsubstancesintos imple
r
afac toryforthep roduc t
iono fa lmo stallo
f bod y weighto ffema leb roods tock
substancessothatthey maybeu sedin
thece ll’
slipidandt rans -memb rane
metabol
ism
prote ins Feedcompos itiontable: al istoffood
Di
luent: avola
ti
lel
iqu
iduseda
long wi
th mater
ials wi
thre spect
ivenu t
rient
Enzyme : comp lexp roteinaceous compos i
tions exp ressedinpe rcentage
so
lventsincoat
ing ma
ter
ia
lsespec
ial
lyto
subs tancestha ta rep rodu cedb yliving
reducecost
cellsandb ringabou to racce lerate Feedconve rs
ione f
fic
iency(FCE ):
reac t
ionso rp rocessesinl ivingce l
ls expresseda spe rcent wet we ightgained
Disaccharides: asugarconsis
tingoftwo
monosaccha r
idesl
inked byag lycos
ide d
ividedb ytheamoun to fd ry weightoffeed
Eicosapen taeno icac id(EPA ): a20 - g
iven w i
thinafeed ingpe riod.Theh igher
bond.Thecommond ie
tarydisacchar
ides
carbonun saturatedfa t
tya c
idha v
ingf ive theFCEva lue,thebe tterthefeedi s
aresucrose(caneo rbeetsugar),la
ctose
,
doub lebond s,anessen tia
lfa ttya cidin ut
il
ized
and maltose
f
i sh
Dissolvedoxygen(DO )
: theamoun tof Feedconve rs
ionra tio(FCR ): theamoun t
Essen tia
lam inoac ids : am inoa c
id s ofdry weighto ffeedg ivenove rthe wet
e
lemen taloxygen, O2,insolut
ion unde
r
requ i
redfo rp ro
te insyn thesistha tcanno t weigh
tga ined w ith
inafeed ingpe r
iod.The
exis
ting atmospher
ic pressu
re and
besyn thesizedinthebod yand mu s
tbe lowe rtheFCRva lue,thebe tterthefeedis
tempe ra
ture. Oxygeni saddedto wate
rb y
prov idedinthed iet ut
il
ized
photosynthes
isandd iffus
ionfromthe
atmosphe re
Essen t
ialfattyac
ids: polyunsaturated Feed
ingcha rts
: thesep rov
ideonlya
fat
tyac idsthatcannotbesyn thesizedb y gene
ral gu
idetofeedintake.Tabu
lated
DNA
: deoxy
ribonuc
leic a
cid
thebod yand mustbep rovidedfrom feed
inglevelsforva
riousspeciesare
d
ietarysou rcesor
iginat
ing outs
idethe commonlyp rov
idedb yfeed manu
fa c
tur
ing
Doublebond: acovalentbondinwh
ich
organism companies
twopairsofelec
tronsaresharedbe
tween
the pa
rt
ic
ipat
inga toms
Eutrophica
tion: enrichmento fthe Feedingfrequency
: thenumbe
roft
ime
s
receiv
ing waterandox ygende f
iciencyin feedisgiveninada y
thesu r
rounding waterresul
tingfrom
aquacul
ture discha
rgesofn itrogen and Feedingra
te: theamountoffeedg
iven
phosphorus, and oxygen
-consum ing perdayandisba sedonacerta
in
substancesintolakes,r
ivers
,andsea s percen
tageofthebody we
ight

Extrusion: toshapeb yforc

ingthrougha Feedingtray
: usua l
lyny
lonsc reenand
special
lyde s
igna
ted openingafte
ra woodo rbamboos t
ripsandstones in
kers
,
previousheat
ingofthe mater
ialorofthe usedto moni
torfeedconsumption
open ing
4
-G G
lossary

Feedstuff
: oneo ra mix
tureof Growth: ano rmalprocessofin c
reasein
subs
p
m
ro
te
ine
tances wh
in
ra
,ca
ls
ichfo
rbohyd
,and wa
ra
te
rmthenu
te
,fa
rtha
t
,v
ta
i
t

reea
r
ien
tamin
s
ts
,
tenb
;

yan
s
izeo
to
de
ta
te
fat
l we
rm
is
igh
sue
to
inedb
,organ
fsamp
yba
t
,oro
ledan
ch we
rgan
ima
igh
ing
ism
lsa
.The
re
, usua
llyin
I
animala spar
to fit
sda i
lyrat
ion ataredcontaine
rwith wate
rona we ighing Immune-response:re
lat
ingtothe
scale phy
sio
log
icalreac
t
ioncharac
ter
ist
icofthe
Fi
lle
r: asubstanceaddedinthefeedto immunestate

H
comple
tethefeedformu
la
Incidencecos
t:thera
tioofthecos
t
Fish meal: driedf
isho rf
ish wa
ste incur
redtotheoutpu
tp roduced
preparedf romnume rousspecieso
ffish Hem icel
lulose
: composedo fa mixtureof
thata reu seda sanima
lfoodand Inc
iso
r:sha
rplyedgedcu
tt
ingtee
th
he xoseandpentoseunits
;anyo fvarious
generallynotdirec
tlyusedforhuman polysacchar
idesthataccompanyce llulose
consump tion Inorganic
:chem icalcompounds wh
ichdo
andligninintheskele
talsubstanceso f
notconta
inca rbona stheprin
cipa
l
woodandg reenplan
ts. Unl
ikecellu
lo se
,
Floatingfeed :p roducedb yane xtrusion e
lement(e xceptca rbona
tes
,cyanides
,
itcanbehyd ro
lyzedinrela
t
ively mi
ld
processth rough wh ichfeed ma terialsa re andcyanates)
acids
moistened,p re
-cooked , expanded(h ighe r
moisture
,tempe ratureandp ressurethan Inpu
ts:theresource
su sedtop roducean
He
rbivores
: animalstha
tfeedexc
lus
ive
ly
ordinary pel
let
ing ) andd ried
,re sul
tingin ou
tpu
tu s
ingatechnology(e.g.fryor
on p
lan
t mate
ria
ls
lowdens ityfeedpa r
t
icles.Thisis mo s
t f
inge
rl
ings,feeds
,landbu i
ltintoponds,
applicab
lefo rsu rfaceandco lumnfeede rs
Hexose: a mono sacchar
ide wi
thsix f
ishcages,labo
rand manage r
ialski
l
ls)
andi tsbeingv isiblea vo
idsove rfeed ing
carbona toms
,andhen cetheemp i
rica
l
formula C6H12O6
.Thenu t
ri
tiona
lly Insulin
:polypept
idehormonethatis
Formula tedfeed
: twoo r mo
reingred
ien
ts impo r
tanthexo se
sa reglucose
, galac
tose secre
tedb yB-cel
lsinthepanc
rea sand
propor
tioned,mixed, and p
rocessed andfructose he
lp sregu
lateglucose me
tabo
lismin
accord
ingtospec i
fica
tions animals
High
lyunsa tura
tedfa ttyacids(HUFA):
Freeenergy
: ene rgy wh
ichisle
ft fat
tyacidsthatcontainfouror mo le In
redoub teg ratedaquacu lturesys tem :system
avai
lab
leforbiolog
ica
l act
iv
ityandg row
th bonds us
ingva riousspe c
iesind i
ffe
renttrophic
af
tertheenergyrequi
rementis met levelssu cha sseaweed ,she l
lf
ish, and
Hydrogensu l
fide(H2 S
): aga sproduced he
rb ivorousandomn ivorousfishspe cie
s
Freeradicals: formedduringl
ip
id bym ic
robia
ldecompo s
itionoforgan
ic sucha sm ilk
fishandt i
lapia
perox
idat
ion,su cha s ROO,RO,OH,the se mat
ter(e.g
.e x
ce ssfeedandfe ca
l was
tes)
reactb yhydrogenremo va
landava riety deposi
tedinthepondbo t
tominthe In
t rape ri
tonea l
: thereg ionbe tweenthe
ofaddi
tionreactionsthatdamageother absenceo foxygen sk
in and mus
clelayer and organs
foodnu tr
ients mak
ingthem unava
ilableto
f
i sh Hydro
lysis
: achem ica
lrea c
tionof wate
r Isoca
lor
ic: ha
vings
imi
larca
lor
icva
lue
in wh
ichthereagentotherthan wateris

G
decomposed and h
ydrogen and hydroxy
l Isome r
ism: havethesame molecu
lar
areadded formulabutthe
irstru
ctu
ralfo
rmulad i
ffe
rin
thea r
rangementofthei
ratoms wi
thina
Glucose: a monosaccha r
ide;ahexose Hydrophil
ic: acompound wh
ichis molecu
le
(six
-carbon)sugar,ofemp i
rica
lfo
rmula so
lublein wate
r,o
raregionofa

J
C6H12O6;bas
ic moleculefo
rthesynthes
is mac
romo lecu
le wh
ichcanin
teractwi
th
ofstarchandcellulose wa
ter molecule
s

Glycero
l: atrihyd
ricalcoho
lto which Hydrophobic: acompound wh ichis Joule: un
itofene
rgyinthe me
trics
ystem
threefat
tya c
id moleculesareester
if
iedin insolub
lein wate
r,butsolub
leinl ip
ids
,or andonek ca
lisequalto4.186k
J
theformationoftr
iacyglyce
rols(
fatsand aregionofa ma cromolecu
le whichcanno
t
o
ils) inte
ra c
twi
th water
,althoughi
tdoe s

G
lycogen: ab ran
chedchainpolymerof
inte
ra c
twithlip
ids
K
g
lucose
,linkedbyalpha1-6lin
ks;the Hypervitaminosis
: anabno rma
l
Kerat
in: asu l
fur
-conta
iningp
rotein wh
ich
s
torageformofcarbohyd
rateinanimals
, condi
tionresul
tingfromthein
takeo
fan
isthep r
imarycomponen tofep
ide rmis
,
a
ss ta
rchisinplan
ts excesso foneo r morev
itam
ins
ha
ir, woo
l,hoof
,ho rn
,andtheo rganic
mat
rixofthetee
th
Golgibodies: membrane-bounded
organel
leineukaryot
iccells whe
rethe
Ke
tose
: asugartha
tconta
insake
tone
prote
insandlip
ids madeinthe
g
roupaspa
rtofi
tsst
ruc
ture
endoplasmicre
ticu
lum are modif
ied and
sor
ted
 Glossary G
-5
Ki
localor
ie: amoun
tofhea
tnecessa
ryto Marginalphysicalproduc
t(MPP )
: the Microencapsulatedfeed: ala rva
lfeed
ra
isethetemperatu
reofonek
i
logramof changeinou tputperuni
tchangeininput madeb yencapsulat
ingaso lut
ion,col
loid
wate
r1 °C ortheincrementintheuni
tsoftheproduc
t orsu spens
iono ffeedingredientmix
ture
orf
ishre su
lt
ingfromoneadditiona
luni
tof w
ithina memb raneo rcapsule;these

L
input(feed
)g iventhef
ish part
ic
lescanbede s
ignedtoha veaslow
releaseofthe ma te
ria
linsidethecap sule
,
Maximumeconom icyie
ld(MEY ): the ortototal
lypreventleach
ingo fthe water-
Lablab: na
turalfoodinponds,composed produc
tionlevel whereprof
i
ti shighes
t solub
le nutr
ients
o
fcomp lexofb lue-g
reenandg reen s
in ce noaddi
tionalor marg
inalbenefi
t
a
lgae,diatoms,roti
fers
,crus
taceans
, canbega inedfroman yaddi
tionaluni
tof M
icronut
rien
ts: nu
tr
ien
tsneededinsma
l
l
insects
,roundwo rms,det
ri
tus
,and input amoun
ts,suchassomevitam
insand
p
lankton m
inera
ls
Maximumsus ta
inab leyie
ld(MSY )
: the
Lac
tose: thesuga
rofmi
lk;adisaccha
ride product
ionlevel where ou
tputperuni
tof M
icro-organ
ism: anyorgan
ismo
f
composedofgluco
seandgalactose inputishighestbutthereisnolongerany m
icroscop
icorsubmic
roscop
ics
ize
increaseinprodu c
tionbe yondth
isleve
lin
Lec
ithin
: acommonsourceo
f sp
iteo fanyaddit
iona luni
toffeedsor Mic
rophagous
: re fer
ringtoorgan
isms
phospho
lip
ids input tha
tfeedonrela
tivelysmallfoodi
tems

L
ignin: apo
lymero
fconi
fery
lalcoho
l;a Mechan ica
lfeed ing : meansofpart
ial
ly Microscope: anop t
icalins
trumentused
s
truc
tura
l ma
ter
ia
lfoundin woodyplan
ts replacinghandfeed ingtoinclude
: toexam ine minu
teob jec
tsb yg
ivingan
automa t
icfeede rs wh ichcanbese tto enlarged
, we
llresolvedimageofthem
Liming: theapp l
icationo fneutral
izing releasecon tro
lledamoun tsoffeed when
agentsspeci
ficallyca lciumcon taining ac
tivated, anddemandfeede rs whichcan Microvi
ll
i: smallslendervascula
r,f
inge
r-
compounds: CaO(qu ickl
imeo rbu rnt releaseafewpe lletsea cht
imea shapedp rocessesofthe mu cous
l
ime )
; Ca(OH )2(calciumhyd rateo rs laked tr
iggering me chan ismi sbumpedb ythe memb raneofthesma llin
tes
tinestha
t
l
ime )
; CaCO3 (agricul
turalordo lom i
tic f
ish serveintheabso rpt
ionofnu t
rien
ts
l
ime )
,toeitherpondbo ttomor wa ter
Metabo
licrate: theamoun
tofoxygen Mitochond
ria
: membrane-bounded
Linearp rogramming:compu tat
ional usedfortota
l me
tabol
ismpe
runi
toftime organel
letha
tcar
riesoutoxida
tive
methoda imedatalloca
tingscarce peruni
to fbody we
ight phosphory
lat
ionandproduces mosto
fthe
resourcesto maximizeprof
itorm inimize ATP
cost(e.g
.compu t
ingfortheleast-costfeed Metabo
lic wastes/metaboli
tes: by
­
combinationsorrat
ionthatst
il
l mee tthe produc
tof metabol
ismd ischargedf
rom Mobi
le: capab
leof mov
ingorbe
ing
nutr
ientsrequi
redb ythecul
turedf i
sh ) thebod yo
fano rgan
ism movedfromonepla
cetoanothe
r;capab
le
o
f mo v
ingorbe
ing movedabou
treadi
ly
Linolen
icac
id: an18-ca
rbonunsatu
rated Metabol
ism : theph ysica
landchem ica
l
fat
tyacidhav
ingthreedoublebonds processesb y wh
ichfeedstu
ffsare Moistpelletrat
ion : ad ietwi
thabout
synthes
izedintocomp lexelements 30% mo isturep reparedf
romd ry
Lip
ids:ab roadte
rmfo rfatsandfat
-l
ike (anabol
ism )
,comp lexsubstancesa re ingredientsandg roundfishbioma
ss,
substancesinc
luding phospho
lip
ids, transfo
rmedin tosimpleone s(ca
tabo l
ism)
, formedin toballso rpel
lets
,andfedfresh
waxes,stero
ids
, andsph ingomye
lins andene rgyis madeava i
lab
leforu seb y orunfrozenbe fo refeed
ing
ano rganism
Lumu
t:fib
rousf
ilamen
tousg
reena
lgae Molar
ifo
rmtee th: havef
lat
tened,of
ten
g
rowninponds metabolizableenergy : a mo
reexact broad
lyo cc
lusa
lsurfa
ce s
;u sedfo
r
measureo fthedietaryenergyusedfo
r c
rushing and gr
ind
ing

M
me
tabolismb ythetissues
Moldinh
ibito
r: subs
tancesaddedtofeed
s
Metamorphos
is: the ma
rkedchangeof
f thatinh
ibi
t moldg
rowth

Macronut
rients: nu
t r
ien
tsneededin shapeo rs
tru
ctu
re,pa
rt
icu
lar
lyinthe
trans
it
ionofonedeve
lopmenta
lstagein
to Monosaccha r
ide: as implesugar
,the
largeamoun tssuchasp ro
tein
s, ba s
icun i
tsf rom whichdisacchar
idesand
carbohyd
rates,orl
ipids another
polysaccharidesa recompo sed.The
Methanegas
: anodo rless
,color
le ss
, nutr
it
ionallyimpo rtant monosacchar
ides
Ma
ltose: adisaccha
ridecomposedo
ftwo arethepen toses(five
-carbonsugars)and
mo
leculesofglucose inf
lammable hyd
roca
rbon , CH4
, which
formsexp
los
ive mix
turesw i
thair
.Itresu
lt
s thehe xoses(s ix
-carbonsugars)

Manualfeeding: includesb roadcast

ing fromthedecayoforganic mat
ter
Monounsa
tura
tedfattyacids
: anyoneo
f
thefeedsintotheponda s wel
la splacing thefa
ttyac
idscon
tainingonedoub
lebond
feedsintof
loatingframes(forfloa
ting Methionine
: oneo fthee ssen
t
ialamino
feeds)orfeedingtrays(fo
rsinkingfeed s
) ac
id s
;itissul
furcontain
ingand maybe
replacedinpartbyc yst
ine
6
-G Glossary

Output
: thep roduc
tofcombin
ingva r
iou
s Phosphat
idicac
id: acompoundinwhich

N inpu

f
ish
tsu
aquacul
,she
s
tu
l
ingatechno
re
l
f
,theu
ishando
logy(e
sua
lou
the
tpu
rva
.g
.in
tsa
luab
rethe
leb y
­
twofa
es
g
ter
i
lyce
f
t
tya c
idsandphospho
iedtotheth
ro
l
reehydro
r
x
y
ica
lg
c
ida
roup
re
so f
Natura
laquat
icfood
: inponds
,thesea
re produc
ts)
lablabandlumu
t Phosphatidylchol
ine: a ma
jorfo
rmof
Oxidat
ion: areac
t
ioninwh ichoxygen phosphol
ipidconta
iningchol
ineinthe
Nebu
lize
: toredu
cetoaf
inesp
ray molecu
le
combineschemica
lly w
ithanother
Neutrall
ipids: the ma
jorfo
rmo fs
torage substance
;areact
ion whichinvo
lvesthe
losso rt
rans
fero
fe le
ctrons Phospho lipid: al
ip
idinwh ichg lyce
rolis
l
ipidsthata recompr
ised main
lyof es
te r
if
iedtotwofa t
tya c
id s
,bu tthethi
rd
tr
iglycer
ides hydroxylgroupi seste
ri
fiedtopho sphate
,
Ni
t
ae
rate(NO3
robics
)
tab
i
l
: theendp
iza
tion o
fo
roduc
rgan
to
ic n
i
t
fthe
rogen
,
P andth
va
ac
r
ie
id,g
roughthephospha
tyofo
lyce
the
rol
rcompound
tetooneo
s;es
, andphosphat
te
id
rso
icac
fa
ffa
id
t
ty
it
sp resencein wate
rind
icateso rganic Part
ialbudgeting: aquan
t
ita
tive
enr
ichmen t proceduretha
tinvolvesa ma
rg ina
l Photosynthesis: thesyn thes
isof
analys
istodeterminethepro
fitabi
l
ityo
fa chemicalsubstances withtheaido fligh
t,
Ni
tri
fica
tion: theaerob
icbacte
ria
l changeinthep roduct
ionsys
tem sucha sintheforma t
iono fcarbohyd ra
tes
convers
ionofammon iaando rgan
ic (e.g
.ing reenplants)fromca rbond iox
ide
n
itrogentostab
lesal
ts(ni
tra
tes)
: Part
iculateorganic mat
ter: par
tic
leso
f and waterwiththeliberat
ionofox ygen,in
common lybyNit
rosomonas spand l
ivingo rdeadorganic matte
rtha
tare thep resenceofch lorophy
ll
Ni
trobacte
r sp suspendedin water
;planktonisaformo
f
par
ticu
lateorganic ma
tte
r Phytop
lankton:m ic
roscopicaquat
ic
Ni
tri
te(NO2 )
: anin
termedia
tep
roduc
tin p
lantssuspendedinthe wa te
rcolumn
;
the bio
log
ica
l ox
ida
tion o
fammonia(NH3
) Pea rsonsqua re me thod : a ma thema t
ical ma
joroxygen-produc
ing organismsina
tonit
rate(NO3) methodu sedinfo rmulatingfeed s pond
app l
icab leon lytofewing redientsand
Non -essen tialam inoac ids: thoseam ino w
ithfewnu trien tstoba lan ce Pinocy
tosis
: aprope
rtyo
fcel
lstha
tlead
s
acidsrequ iredfo rp rote
insyn thesistha t touptakeof ma
cromo
lecu
les
canbesyn thesizedinthebod yin Pec tin: foundp r
ima ri
lyonthespa ces
adequa teamoun tsto mee trequ iremen ts, betweenp lan tce l
l wallsand ma yalso Plankt
ivores
: animalsthatfeedon
andthe reforedono tha vetobep rovided infi
ltratethece l
l wa l
lit se
lf p
lan k
ton,the mi
croscopicplan
tand
inthed iet animall
ifein wa
terinclud
ingba c
ter
ia
Pep tidebond : thel in kbe tweenam ino
Nutrien t: anysubs tancetha ti s acid sinap rotein ;formedb y Plank
ton: the m
icroscop
icplan
tand
physio logicallyuse fulo rnece ssaryfo r condensa t
ion be tweentheca rboxylica cid anima
lli
feinthe waterin
clud
ingbac
ter
ia
grow th and deve lopmen t group( -COOH )o foneam inoa cidandthe
am inog roup(NH2 )ofano thertog ivea Plantgums: comp lex
,h ighlybranched
Nutrition : thesc ienceo fnou rishinga n -CO- NH -l in kbe tweentheam inoa c
ids re s
iduescon
tain
ing D -g
luco ron
ic and D
-
organ ism ;thesumo fthep rocessesb y galactu
ron
icacid
sa long witho the
rs imp
le
whichanan imalo rp lantabso rbsand Peris talsis: su ccessive wa veso f sugarssuchasa rabinoseandshammo se
ut
il
izefoodsubs tances .Itin
vo lvesthe invo lun tarycon t
rac t
ion pa s
singa longthe
inges tion,d iges
tion, absorpt
ion and wal
lso fthein te s
tineono therhallow Polysaccha
rides
: formedbythe
transpo rtoffoodnu t
rien
tsintobod ycel
ls muscu la rst ruc tureandfo rcingthe combinat
ionofhexosesoro
ther
andre lea seo f wastep roduc
tso f con tents onwa rd monosaccha
rides
metabo lism
pH: thepHi sde f
ineda sthenega t
i ve Predator: anyorgan
ismtha
tca
tche
s

O
loga rithmo ftheh yd
rogeniona ct
i v
ity:pH anotherforfood
=-Log (H+ )
Premix
: auni
form m
ixofmic
ro­
O
ligosacchar
ides : agene ra
ltermfo
r Phenyla
lanine
: oneo
ftheessen
t
ial ingred
ien
ts w
i
thadiluen
torcar
rie
r
po
lymerscon ta
ining abou
t3 -10 aminoacid
s;it ma
yberep
lacedby
monosaccha
rides ty
rosine Pr
ima ryproduct
ion:i tisthetota
l
increaseinb
ioma ssofgreenp lan
ts
Omnivores: an
imalstha
tder
ivethe
ir Phospha tes
: an yofsevera
lsubstances observedove
rape r
iodoftime,expres
sed
nu
tr
ientsfrombothplan
tsandanima
ls contain
ingan ysal
to resterofphosphor
ic asp roduc
t
iondiv
idedb ythepe r
iodoft
ime
ac
id(e .g
.phosphateo fcalc
ium),useda s
Organic mate
ria
ls: re
lat
ingtothe fer
ti
lizertosupplypho sphoru
s Productioncost
: thecos
to r moneta
ry
compoundso fcarbon(o
therthansomeo
f valueoftheinpu
tsusedinthep roduc
tion
itssimple
rcompounds) process
 Glossary G
-7
Product
ionfunct
ion :anexpress
iono
fthe Sucrose
: ad isacchar
ideha vingthe
techn
ica
andinpu
techno
lre
t
la
sa
logy
t
ionsh
tag i
ip be
vent
tween ou
imeusinga
tpu
ts
S formu
andf
la C1
ruc
to
2
se
H2
2O11.I
th yd
rolyzestogluco
se

Sa
lini
ty: theconcentra
tionofm ine
ral Sugar
: chem ica
lly,a monosac
charideor
sa
ltsinthe wate
r,of
tenexpressedinpp t smal
lo l
igosaccharide
. Caneorbee tsuga
r
Prof
it: thedif
ferencebetweentheva
lueof (par
tsperthou sand
) issu c
rose,ad isacchar
ideofglu
co seand
tota
lp roduct(VTP )orgrossre
venueand
thetotalvalueofva r
iab
lecost(TVC)
;or fruc
tose
Sampling method: aspeci
ficprocedure
s
imp lyrevenue m inuscost forsampling
,thedetai
lsofsett
ingupthe Supp lemen t: afeedu sed wi
thano therto
ins
trumentanduse
,andthep
roto
colfo
r imp ro vethenu tr
it
iveba lanceo r
Pro
fi
tindex: thera
t
ioo
fthene
trevenue sample p
repa
rat
ion performan ceoftheto talandin tendedto
toto
talcos
t
be( I)fedund i
luteda sasupp lementto
Sat
iat
ion: thequa
l
ityorsta
teo
fbe ing otherfeed s
;( I
I)offe
redf reechoice wi
th
P
rote
in: apo lymero
fam
inoac
idsjo
ined sa
tia
ted;theac
torprocessofa
chieving otherpa rtsoftherationsepa rate
ly
b
ypeptide bonds qual
if
ica
tion available
;o r(I
II)fur
therd i
lutedand m ixed
top roduceacomp letefeed
Polyunsaturatedfat
tyacids(PUFA): Secchid isk: adisk20cmind iameter
fa
ttyacid swi
thtwoo r morecarbon
-carbon pain
tedina l
ternatequadrantsblackand Supplemen talfeed
: feedsuppliedto
double bondsinthe molecu
le,separa
ted whi
te(attachedtoapo le w
i
th meetthenutrien
trequi
remento
ffishfor
bya me thy
lene( -CH2
)group gradua
tion s
)andu sedto mea sureligh
t ma
intenanceandg row
th when natu
ral
penet
ration,transparenc
y,orturbidi
tyin foodi sinadequate
Pur
if
ieddiet
: adie
tcomposedofpur
if
ied the watercolumn
ing
redien
tse.g
.casein
,ge
lat
in,e
tc
. Surv
ival
: thef
ina
lnumbero
fan
imalsle
ft
Semi-pur
ified
:d ie
tcomposedpar
tl
yof isdiv
idedbytheor
ig
ina
lnumbe
rof
Pyrophospha
te: acoenzymeinvo
lvedin h
ighly pu
rif
ied and p
rac
t
ica
ling
red
ient
s, an
imalssto
cked
thetrans
fero
ftwo-ca
rbonunit
s e
.g.fish mea
l

Sinkingfeed:p reparedthroughe xt
rusion
T
Q
Qual
itat
ive
: concerned w
i
th,re
lat
ingtoo
r
unde
su
p
la
rfa
chtha
i
rlylowtempe
tpel
cedin wa
le
te
tsp
r
.P rawn
ra
rodu
tu

sa
reandp
cedsin
rebottom
re
k when
ssure
Technicale
fficiency: atechnology-ba
sed
feedersanda regiventhistypeo ffeed cr
ite
riaforevaluatingthedesi
rabil
ityofa
invo
lving qua
l
ity produc
tionproce ssortheuti
li
tyo faninpu
t

Qua
litycontro
l: sys
temforensur
ing Soylec
ith
in: le
ci
thin madef
romso
ya
beans Tempe ratu
re : thequan ti
tat
ivestatement
qua
li
tyo foutpu
tinvo
lv
inginspe
ct
ion,
concerninghea tu suallyexpressedin
ana
ly s
is,andact
ionto ma
kerequi
red
changes Sphingomye lins: fattya c
ide s
terso - deg
flong ree s Celsius( °C)
cha
in a lcohol andsph ingos
ine
Tetrahyd ro
fo late: the me tabol
ical
lya c
t
ive
Quan
ti
tat
ive: re
lat
ingtoo
rconce
rned
Sta
rch : apo lyme rofg lucoseun i
ts;a re formo fthev i
tam infol
ica cid
,aca rr
ierof
w
ithquan
ti
ty
usual
lypo lycycliclong -cha
in alcohols; one-carbon g roups
pr
incipalstoragefo rmo fcarbohydratesin

R p
lan

Sta
t
t

ic
s

: re lat
ingto,o ru sedinwe ighing;
T

p
igh
adja
re
tjunc
cen
vent
tep
t
ion
ithe
: ce
l
ingthepas
ia
l
l
lce
-ce
l
l
ljunc
lstoge
sageo
t
iontha
the
f mos
r,
tdis
tsea

so
ls

lved
Rancid
ity
: smel
l
ingo rta
st
ingfoul exer
tingfo rceb yrea sono f we
ighta lone moleculesfromones ideo ftheep i
thel
ial
because o
fchemica
lchange, espec
ia
lly apar
tf rome ffectsofine r
tia sheettotheo ther
duetoage
Ste
roids: compoundsder
ivedf
rom Tota
lva lueofphysica
lp roduct(TVP):the
Rat
ion
: thefooda
llowanceo
fonepe
rson cho
lestero
l(i
tse
l
fas te
roid
), mo
sto
f wh
ich monetaryvalueo
ftheou tpu
t(f
ish)thatis
oronean
imalperday are ho
rmones obta
inedb y mul
tip
ly
ingtheun i
tp r
iceof
thep roduc
ttothetota
lquanti
tyo fthe
Reci
rcula
ting
: sys
temo
fus
ingrec
ycled Stockingdens ity
: referstothenumbe ro
f product
mate
ria
l cul
turedan imalsstockedpe runi
ta rea
(e.g
.25 ,000juveni
lespe rhecta
re). To
talvar
iableinpu
tcost(TVIC)
:thesum
Res
idue: someth
ingtha
tremainsa
fte
ra Naturalpondp roduct
iv
ityisimportantin o
ftheco s
tofthevar
iab
leinput(
feed
)at
par
tistaken
,sepa
rated
,removed,o
r extensivecul
tures ys
tem whe restocking d
if
feren
tlevelso
finpu
t
des
ignated densityislow
Tox
in: anyo
fsevera
lin
tenselypoisonous
RNA
: r
ibonuc
leica
cid Substan
tia
l: cons
ist
ingo
f;re
lat
ingto
, subs
tancesp
roducedbycertainbacter
ia
shar
ingthenatureo
f,orcons
ti
tut
ing
subs
tance
8-G G
lossary

Trashf
ish
: anyofvar
iousseaf
ishestha
t Unsa
tu ra
tedfattyac
id: anyoneof Vo
lat
ile: asubs
tance wh
ichcanbeea
si
ly
havelow ma
rke
tvalueashumanfood seve
ralfat
tyacidscon
tain
ingoneor mo
re l
iberated
doub
lebond s
,e .g
.ole
ic
,lino
le
ic,l
ino
len
ic

W
Tr
ialande r
ror method: a methodusedin &arachidon
ic
ca
lcula
tingfeedformula
tion us
ing many
ingred
ientsbytr
ia
lande rro
r U
rea
: the ma
inexcre
toryend
-produ
cto
f
am
inoac
id metabo
l
ism Wate
rqua l
ity
: thelim i
t
ingconcen t
rat
ion
T
riglycer
ides
: esterso
ffa
ttyacidand
ofa wate
rconsti
tuen
t(po l
lutan
t)ordegree

V
g
lycero
l,the ma
jorformofsto
ragelip
ids
ofin
tensi
tyofsomeo the
rad ve
rse
cond
it
ion whichispe
rm i
ttedinabod yof

U V
il
li
formtee
th:a re mo
reo rle
s s
wate
r

Z
e
longa
tedcard
i
formin whichthelengthto
Unsa turated: anorgan
iccompound d
iamete
rrat
ioresemblesthatofinte
st
ina
l
containingoneo r mo
recarbon-ca
rbon v
i
ll
i
doublebond s
,andthere
forelessthanthe
Zooplankton
: smal
lanima
lsinwa ter
possible propo
rt
ion o
fhydrogen Vi
tamin: ano rganiccompoundrequ i
red mak
ing uptheseconda
ryprodu c
tionleve
l
insma l
lamoun tsforthe ma
intenanceof wh
ichdependonthe water movementfor
Unsaturatedfat
: afatformedf
romthe normalgrow th
, health
,and me tabo
l
ic locomotion
react
iono fgly
cerolwi
thanyoneo f integ
ri
ty. Def
iciencyofavitaminresul
tsin
severalunsatu
ratedfa
ttyac
ids,e.g
.ole
in thede ve
lopmen tofaspecif
icde f
ic
iency
&linole
in d
isease, whichcanbecu redo rpreven
ted
on
lyb ythatv i
tamin
 I
llus
tra
tion and Pho
to C
red
its CR
-1

I
llus
tra
tionandPho
to C
red
its
Photos andil
lustrat
ions are p
roper
ty o
f SEAFDEC Aquacu
ltu
re
Depar
tment un
les
so therw
isecred
iteda
sfo
llow
s:

Chap
ter2

Pho
to2
,p 38,Pho to2,p 40,Pho to4 ,p 42,andPho to 1,p 42
from Manua lforFish Disease Diagnosis
-II. Mar
ineFishand C rus
tacean
DiseasesinIndonesia byIstiKoesha ryani, DesRoza. Ketu
t Mahardika,
Fris Johny, Za
fran and Ke i Yuasa. Ketu t Sugama,K ish
io Hatai and
Toshihiro Naka
i(eds)
.2001 . Gondo lResea rchInst
itutefor Ma
ricul
tu re
,
Indones ia
,andJ ICA,Japan .

F
igu
re2
.23
, p 49,Figure2.31,p54andF igure2.32,p 55
from Molecular Ce
llB io
logy
, Cha r
lote J
. Ave
rs. © 1986 by Add
ison-
Wes
leyPub l
ishing Company,Inc. Reprin
tedbype rmiss
iono fPearson
Educat
ion,Inc.
,U .S.A
.

F
igu
re2
.16 p
.61photoofmilkf
ish w
ithcatarac
t
f
rom Ce
liaLav
il
la-Tor
res,Fish Hea
lthSec
tion
, SEAFDECAQD

Chap
ter5

Pho
tos : p
. 127and 129
,legum
inousseeds ©Jowaman Kha
jare
rne
tal
.1987

p
. 129
-131and 137
,hamme rmi
lls
,si
fte
r,m ixer
,pu lve
rize
randbagge
r
©Dinnissen
-BensAgr
i-Bus
inessInte
rnational Co
rp.2000

p
. 133-136
,pe
lle
tcoo le
r,drye
rsandsepa
rato
rs: ©Wenge
r
Manufac
tur
ing
,Inc. 1999

p
. 134
-135and 144
,feeds
truc
turesands
torage © DeanAk
iyama 1989

p
. 144
,Aspe
rgi
llus
flavussp
. © Ba
ticadose
tal
. 1988andLeaño2001
 I
ndex I
-1

Index
α-amy lase 88 antioxidan ts 21 , 30, 104 breeding 192 , 199
AAA 155 appa ren tp rote
ind igestib
il
ity 9
1 brokenline 53
AAS 160 appe tite 92 brokenlineana lys
is 16
abio ticfac to rs 93 aquacu lture 1 broodstock 182 , 184
, 185, 187, 188
, 190
abno rma lsw imm ing 49 aqua feeds 2 broodstockandseedp roduc
tionintanks
abso luterequ i
remen t 54 arabinose 34 201
abso rp t
ion 86 , 88 , 89 arginine 10 broodstockintanks 193 , 196
abso rptive ,s torageo rR -ce
lls 84 arousa landsea rch 94 brushbo rder 83
ace ta tefragmen ts 28 arsen ic 57 BTU ’s 161
ace tyl 47 ascites 5 1 build
ingb locks 8
acid -baseba lance 57 asco rba te-2-monophospha te 50 bulkdens ity 150
act
iva tion 58 asco rba te-2-polyphospha te 50
act
ivet ranspo rt 86 asco rbicac id 50 cadm ium 57
acylca rr
ie rp rotein 47 ash 152 , 154 ca
lcium 57
adenos ine 42 asianseabass 188 Calc
iumandphospho rus 160
adenos ined iphospha te 42 atom icem issionspec troscopy 160 ca
lciumandphospho rusdef
icienc
ies 58
adenos inet riphospha te 42, 59 ATPcon tent 28 ca
lo ries 161
adipocy tes 28 at
trac tab i
li
ty 151 canine 80
adipose 2 1 att
rit
ion m i
ll 129 captu re 94
agricu ltura lby -produc ts 4 automa ticope rat
ion 158 carapace ,hepa topanc reas 60
aldo lases 6 1 averagephys icalp roduct 212
, 214 carbohyd rate me tabol
ism 45
aldoses 33 averagephys icalp roduct(APP) 212 carbohyd rates 33 , 88
algae 82 carboxypep tidase A 87
algeb raicequa t
ion 108 Bcomp lex 45 carboxypep tidase B 87
a
lka lin i
ty 176 (3-amy lase 88 carboxypep tidases 87
alpha -helix 12 (3-oxida t
ion 29 cardiform 80
a
lte rna tivep roteinsou rces 3 balancedam inoacidprof
iles 64 carnivo res 77 , 107
a
lum inum 57 batch m ixing 138 carnivo rous 37
am ides 153 beta-shee t 12 caroteno ids 51 , 153
am ino ac idtox icity 17 bigheadca rp 185 catabo lism 29
am ino ac id de f
iciency 17 b
ile 83 ca
ta rac t6 1
am ino ac idan tagon ism 17 b
ileac ids 83 ce
llimmun ity 48
am inoac id 155 , 162 binders 104 ce
llnec rosis 51
am inoac idcompos it
ion 14, 155 bioava ilability 58, 101 ce
lls ize 60
am inoac id p rofi
le 16 biochem ica lpa thway 28 ce
llob iase 88 , 89
am inoac idtes td iets 14 b
iolog ica l 149 ce
llula r 23
am inoac ids 8 b
iolog ica lflu ids 160 ce
llula ra t
rophy 47
am inoacy lsyn the tases 59 b
iolog ica lpa rame ters 165 ce
llula r me tabo l
ism 42
am inosuga rs 153 b
iolog ica lsys tems 43 ce
llulase 36 , 88
am inopep tidases 87 b
iolog ica lva lue 19 ce
llulose 36
ammon ia 153 biomass 171 , 177 chaine longa tion 28
ammon ia-N 176 biomemb ranes 2 1 chem ica l 149
amy lase 37 , 88 b
iosyn thes is 21 , 27 chem ica lana lysis 155
anabo l
ism 29 biot
icfac to rs 92 chem ica ldeg rada tion 36
anabo l
ism o rsyn thesis 42 b
io t
in 48 chem ica lformu la 24
ana lys is 156 bl
indness 50 chem ica lstruc tu res 7
ana tomy 79 b
loodc lotting 53 chem ica l wo rk 41
anem ia 47 , 48 b
loodvesse ls 50 ch
itin 37 , 84
anima ls ize 44 bombca lorime te
r 41, 160 chlorine 57
ano rex ia 5 1 bonefo rma tion 52 choleca lci
fe rol 52
antinu t
rit
iona lfac tors 99, 126 bonet issue 50 choles te rol 22 ,5 1
antiox idan t 52, 60 breakpo int 16 choline 51
 I
-2 Ind
ex
chroma tog raphy 162 diges t
ibil
ity 14, 90 ethers 22
chrom icox ide 90 diges t
ibil
ityofnu t
rien
ts 164 evalua t
ion 149
chrom ium 57 ,6 1 digestible 33, 161 excessam inoac ids 14
chylom ic ra 89 digestibleca rbohydrate 33 excesses 7
chymo tryps in 87 digestibleene rgy (DE )41, 16
1 exopep tidases 87
chymo tryps inogen 87 digestion 43 , 86, 88, 89 exoph thalm ia 51
ci
tricac ido rkrebscyc le 29 digestiveenzymes 83 , 86 exop thalmiceye 50
c
lubbedg i
lls 47 digestivesys tem 79 exoske leton 37 , 57, 58
coba lam in 49 dipep t
idases 87 exo t
ic 3
coba lt 57 , 61, 159 directcos ts 209, 220 externalenv i
ronmen t6 1
coenzyme 45 disaccha rides 33 extraneous ma teria
ls 150
coenzyme A 47 dispensab le 9 extrusion 133
coenzymes 45 dissolved ma tte
r3 eyeles ion 5 1
co fac torso rac tiva tors 57 dissolvedoxygen 93 , 175, 176
co ld wa te r 21 distendeds tomach 5 1 fa
tt issues 2 1
common name 24 distr
ibution 92 fa t
tyac id 2 1
compe tition 92 docosahexaeno ic 3 1 fa t
tyac idcompos i
tion 21
comp leted iet 171 donkey ’sea rabalone 199 fattyac id me thyles te rs(FAME ) 157
compounds 159 doseresponsecu rve 15 fattyac ids 89
concen tra t
iong rad ients 42 dryex trusion 133 feca l 43
cond it
ion ing 132 dry ma tterbasis 106 feces 45
con juga ted p rote ins 1 1 feedadd itives 2
connec tivet issues 50 eco log ica lly-sound 4 feedapp lica tion me thods 174
coppe r 57 , 159 econom ice fficiency 220 feedb inde rs 33
cos to fp roduc tion 213 econom ice fficiencyo ffeeds 214 feedcompos itiontab les 106
cos t-effec tive 3 econom icind ica tors 216 feedconve rsion 173
cos t-efficien t4 edema 48 feedconve rsionra tio(FCR ) 177
, 214
cos tso ffeeds 209 effectivefeed ing p rog ram 1 feeddeve lopmen t2
crudeashana lys is 160 effic
iencyo ffeedu ti
lization 164 feede fficiency 37 , 164
crudefa t 153 , 154 effic
iencyo fp roteinu til
izat
ion 164 feedfo rmu la 119
crudefa to re the r-extract 152 eff
icien tfeedconve rsion 3 1 feedfo rmu la tion 104
crudef ibe r 152 eicosapen taeno ic 3 1 feed m ic roscopy 150
crudep ro tein 152 electrica l wo rk 4 1 feedpa rticles ize 174
crumb ie r 136 electront ranspo rt 60 feedpe rfo rmance 2
crus tacean 84 elonga te 29 feedqua l
ity 149
crys tall
ine L -am ino ac ids 17 emb ryo mo rtalities 60 feedsu rveys 4
cultu reinponds 184 emb ryon ico rE -cells 84 feed ing 1
cyanogens 127 endogenous m ine rals 6 1 feed ing hab its 77
cytoch romecox idase 60 endopep tidase 86 feed ing managemen t2
endop lasm icre t
icu lum 83 feed ingp rocessinf ish 92
darkco lo ra
tion 52 ene rgy 4 1 feed ingra t
ion 172
darksk inco lorat
ion 50 ene rgyba lance 4 1 feed ingt r
ia ls 18
denovob iosyn thes
is 28 ene rgybudge t 43 feeds tuffs 149
deam ina ted 14 ene rgyrequ iremen ts 4 1 ferti
lizers 170
def
icienc ies 7 ene rgyva lues 16 1 fibrousp ro te ins 1 1
def
iciencys igns 55 , 57 ene rgy -rich phospho rusbond 42 f
ishfa rm was tes 177
dehull
ing 128 ente rok inase 87 f
ish mea l4
dehyd rogenases 6 1 env i
ronmen t-friend ly 3 f
ish nu trition 1
deprecia t
ion 210 env i
ronmen talfac to rs 2 1 f
lamespec trome t
ric me thods 160
der
ived p roteins 11 enzyme 37 f
lav inaden ined inuc leot
ide 46
desatura te 29 epibranch ia lo rgan 79 f
lav in mononuc leotide 46
desatura t
ion 28 epithelia lke ra tin ization 5 1 f
loa tingfeeds 174
desmosomes 83 ergoca lc ife rol 52 fol
icac id 50
det
ri
tivo res 77 erosion 47 foodava ilab i
lity 92
det
ri
tus 82 errat
icsw imm ing behav ior 46 foodpassagera te 90
dext
rin 36 , 88 esophagus 79 ,8 1 foregu t 79 , 81 , 84
dext
rinase 88 essen tia l7 fragilee ryth rocy tes 53
d
iet 26 essen tia lam inoac idindex(EAA I) 156 freeam inoac ids 14
d
ietcompos ition 53 essen tia lam inoac ids 10 freeene rgy 41 , 42
d
ietaryin take 62 essen tia lfa ttyac ids 2 1 freefa t
tyac idva lue(FFA ) 157
d
ietarynu tr
ien tleve
ls 53 essen tialnu trien ts 1 freerad ica ls 30 , 52
d
ietaryp recu rsors 28 essen tialo rind ispensab le 9 frequencyo ffeed ings 175
d
iets 99 esterif
ica t
ion 157 freshwa te r 26
 I
ndex I
-3
f
ruc
tose 34 impa iredco l
lagenfo rmat
ion 50 linolenicac ids 28
impa iredfa t me tabo
lism 51 lipases 89
ga lactose 34 impa irmen t3 1 lip
idex trac t
ion 157
gallb ladder 79 inv it
ro90 lip
idox ida t
ion 54
gasch roma tog raphic 156 inv it
roassays 9 1 lip
idranc idi
ty 157
gas tr
icemp ty
ingra te 90 inv ivo90 lip
idsou rces 27
gas tr
icemp ty
ingt ime 51 inapp rop r
iatefeeds 1 l
ipid -solub le 45
gene ralformu la 33 , 34 incidencecos t 214, 217
, 220 l
ipids 21 , 83 , 89
gene ralstructu re 63 inciso r 80 l
iqu idfue ls 161
gizza rd 79 indiges tible 33 l
ive r 28 , 83
globu larproteins 1 1 indirec tcos ts 209 , 220 loca t
ionandiden ti
fica
tion 94
glucose 34 , 88 inhibition 29 lordos is 50
glutathione pe roxidase 60 inorgan ic 57 losso fappe t
ite 47
g
lyce rol 89 inorgan ice lemen ts 57 low -dens ityex tens
ivecu l
ture 53
g
lycogen 36 , 83 , 88 inorgan ic m ineralsal
ts 154 lumu t 170
golgibod ies 83 instab il
ity 46 lysine 14
gossypo l 162 insu l
in 84 lysozyme 48
gossypo llevel 162 intens ivecu lture 171
graze rs 78 interrup tedsys tem 24 mac ro,m icro ,andt race m inerals 57
grinding 129 intestine ,live r 79 mac rocy ticanem ia 50
grossene rgy 4 1 inve r
teb ra
tes 82 mac ronu t r
ien ts 45
grow -out 182, 200 iodine 57 , 159 magnes ium 57 , 159
grow -outculture 186 , 188, 189
, 191
, 193 iron 57 , 159 main tenanceo fce l
lin tegri
ty 58
grow th 42, 164 , 173 ironpowde r 90 majo rfam i
lies 25
grow thresponse 18 malfunc tion 14
gus tat
ion 94 jerky mot
ion 47 maltose 35 , 36 , 88
joules 161 mamma lian 37
hard -
tissue ma tr
ices 57 junctions 83 manganese 57 , 159
headgu t 79 , 80 mang roveredsnappe r 192
hema toc r
itva lue 60 ketoses 33 manua lfeed ing 174
hem ice l
luloses 36 keyissues 3 marg inalphys icalp roduc t 214
hemocyan in 60 k
ilocalo
rie 41 marg inalphys icalp roduc t(MPP ) 212
hemog lobin 13 , 49 k
ilocalo
ries 43 marg inalp roduc t 212
hemog lobincon tent 60 k
ilojou
le 41 marg inalra teo fsubs t
itut
ion(MRS ) 218
hemo lymph 60 , 84 k
jeldahl me
thod 153 marinef ishes 26
hemo rrhages 46 maximum econom icy ie
ld(MEY ) 214, 220
hemo rrhag iceyes 47 L-asco rbicac id 50 maximum p roteinsyn thesis 44
hemo rrhag icg i
lls 53 lablab 170 maximumsus tainab leyield 211
hemo rrhag ick idneys 5 1 lactose 35 maximumy ield 214
hepa topanc reas 57 , 84 larvaeinha tchery 194 measu remen t4 1
herbivo re 32 larvaeo rfry 182 measu remen to fl
ip idqua li
ty 156
herbivo res 77 , 107 larval 186 mechan icalene rgy 42
herbivo rous 37 larvalandnu rseryrea r
ing 188 mechan ica lfeede rs 174
hexane 128 larvalrea ring 184 , 190 mechan ica l wo rk 4 1
hexoses 34 larvalrea ringandnu rsery 196 melan ism 50
h
ighdens ityin tensiveculture 53 leach ing 54 memb rane 27
h
ighreso lution 158 leach ingra tes 58 memb ranef luidity 27
h
igh lyunsa turatedfattyacids 23 lead 57 mercu ry 57
h
indgu t 79 , 83 , 84 leas t
-cos tcomb ination 217 metabo l
icene rgy 2 1
his
tidine 10 leas t
-cos tfo rmulation 119 metabo l
icpoo l 14
hormones 22 lensca tarac ts 47 metabo l
icp rocesses 57
HPLC 158 lensde forma t
ion 5 1 metabo l
icra te 42
HPTLC 162 lesionsandedema 47 metabo l
ic was tes 4
hydroch lo r
icac id 153 lesionsintheco lon 49 metabo l
ismchambe rs 4 1
hydrogensu l
fide 176 lethargy 50 , 52 metabo l
ism , nerveand musc lecon t
ract
ion
hydrolys is 34 , 46 leucaenaleucocepha la 127 58
hydrolys isis 34 l
igh texposu re 44 metabo l
izab le 161
hype rirr
itabili
ty 48 l
igh tintens ity 93 metabo l
izab leene rgy (ME )41, 16 1
hype rplasia 5 1 l
ignin 37 meta l
loenzymes 60 ,6 1
hypoch rom ic 60 l
im it
ingam inoac id 14 meta l
lop rote ins 57
hypoch rom icanem ia 49 l
inea rp rog ramm ing 108 , 219
, 220 meth ionine 10 , 14 , 16
l
ino leic 28 methy lene( -CH2 -)in ter
rup tedsys tem 24
l
ino lenic 25 m
icrob ialac t
ion 37
 I
-4 Index
m
ic rob iolog ica l 149 phate 47 phospha te 52
m
ic rob iolog ica l me thod 162 ni
tr
ite-N 176 phospha tidic 2 1
m
ic robound 106 ni
trogen -
freeex trac
t 154 phosphok inases 59
m
ic rocy ticanem ia 60 nomenc latu re 24 phospho lipases 89
m
ic ronu t rien ts 104 non-essen tial7 ,9 phospho lipids 2 1
m
ic rophagous 78 non-l
ip idsou rces 27 phospho rus 3 , 57
m
ic roscope 150 non-p roteinene rgy 41 phospho rusava ilabili
ty 59
m
ic rosomes 28 nonpro teinene rgy 43 pho tophob ia 47
m
ic rovill
i 83 nonrum inan ts 34 phys ical 25 , 149
m
idgu t 79 , 82 , 84 nucleicac ids 153 phys icalac tiv i
ty 42
m
ig rato ry 26 nutr
ien tleach ing 17 phys icalp rope r
ties 53
m
ig rato ryf ishes 26 nut
rien trequ iremen ts 1 phys iolog icfunc tions 57
milkfish 178 nut
rien tre ten t
ion 163 phys iolog ica lfunc tions 45
milkfish b roods tock 178 nut
rit
iona lde ficiencydiseases 45 phys iolog ica ls ta tus 44
milkfishf ry 180 nut
rit
iona lde ficiencysigns 45 phys iology 79
milkfishing row -outponds 180 nut
rit
iona lva lue 41 phytop lank ton 181
milkfishla rvae 179 nut
rit
iona lly 2 p
inocy tos is 86
mimos ine 127 nut
rit
iona lly-ba lancedfeed 1 p
lank t
ivo re 78
mine ralcompos it
ion 160 nut
rit
iveva lue 64 p
lank t
ivo res 77
mine raleva lua t
ion 159 p
lan tgums 37
mine ralfunc tions 57 offeed ings 175 p
lan t ma te r
ia ls 160
mine ralp rem ix 6 1 off-odors 150 plan tnu trien ts 160
mine ralrequ iremen ts 57 oleic 25 , 28 po l
lu tan ts 2
mine ra l
-de fic ien tdie ts 61 olfact
ion 94 po lyme rs 63
mine rals 57 omn ivo res 77 , 107 po lypep t
ide 8
mine ralso rino rgan ic 159 omn ivo rous 37 po lysaccha r
ides 33
mitochond ria 28 , 29 , 83 op t
icalisome rs 35 po lyunsa tu ra ted 23
mo isture ba lance 153 op t
ima lforag ingtheo ry 93 pondcu lture 198
mo isture /d ry ma tter 152 op t
ima lp ro
te in 19, 41 poo rg row th 47 , 60
mo lari
fo rm 80 op t
ima lp ro
te inlevels 19 poo rha tch ingra tes 60
mold Aspe rg illusflavus 162 op t
imum p roteinrequ irement 18 pos tlarva lse t
tlemen tand nurse
ryrear
ing
moldinh ib ito rs 104 orange -spo ttedg roupe r 190 200
moltingo recdys is 57 organicche lates 58 po tass ium 57
molybdenum 57 osmo regu lation 57 , 58, 59 po ten ttox ins 162
mono -glyce rides 89 osmo ticp ressu re 57 precu rso r 10 ,5 1
monohyd rica lcoho ls 22 osmo tic wo rk 41, 42 preda tors 78
monophospha te 42 oxidat
ion 21 prima ry 12
monosaccha rides 33 oxidat
ivephospho ry
lation 53 prism monoch roma tor 160
monounsa tu ra ted 23 process ing 126
mou th 79 , 80 palatablera tion 43 produc tioncos t 210
MSY 214 palm i
tate 28 produc tionfunc t
ion 211 , 213, 220
mucus 8 1 palm i
toleic 25 proenzyme 86
mudc rabs 196 palm i
toleico rhexadeceno icacid 25 pro f
iles 2 1
mufflefu rnace 154 panc reas 79 , 84 pro f
it 214
musc lea trophy 46 , 49 panto then icac id 47 pro f
itindex 214 , 217 , 220
muscu lardys trophy 60 parasites 78 prom isinging red ien ts 4
myo inos itol5 1 part
iallyd iges tible 33 prope rty 25
pearson ’ ssqua re method 108 pros tag land ins 21
nat
iveca tfish 186 pect
in 37 proteases 86
naturalorin tactp roteins 16 pect
inase 37 proteine ff
iciencyra t
io 38
naturalproduc t
ivity 4 pel
leting 132 prote inqua l
ity 7 , 155 , 156
naturaltoxins 161 pepsin 86 proteinrequ iremen ts 7
netene rgy(NE ) 161 pepsinogen 86 proteinsou rce 100
necrosis 47 peptidases 6 1 proteinspa ring 33
negativee ffect 32 pept
idebonds 8 protein S truc tu re 12
nerveimpu lsetransm ission 58 per
itonea lcav ity 48 proteins truc ture 12
nerveimpu lses 51 peroxida t
ion 2 1 proteins 7 , 86
nervousd iso rde rs 48 peroxideva lue 157 proxima teana lysis 152
NFE 101 , 161 peroxides 30 , 60 proxima tecompos i
tiono ffishsamples
n
iacino rn icotinicac id 47 pH 176 (
initialand 165
n
ickel 57 pharyngea ltee th 79 purif
iedo rsem i-purified 15
n
icotinam ideaden ined inuc
leot
ide 47 pheny lalan ine 16 pyloriccaeca 83
n
icotinam ideaden ined inuc
leot
idephos­ phospha tases 59 ,6 1 pyloricceca 79
 I
ndex I
-5
py
lorus 79 slugg ishness 47 transformationo fenergy 45
py
ridoxa
lphosphate 48 sod ium 57 transparency 176
py
ridox
ine 47 so f
tt issues 57 transport5 1
py
rophosphatases 59 so l
id ma tter3 trehalose 88
sou rceso fene rgy 100 tria
lande rror 108
qual
itat
ive 7, 53 sox tec 154 trig
lycerides 2 1
qual
ity 2 spas ticconvu lsions 49 truep rote
ind igest
ibi
l
ity 91
qua
litycontro
l 149 spec ies 53 truep rote
inva luedetermina
tion 155
quanti
tat
ive 7, 53 spec tropho tome te r 158 trypsin 87
quanti
tat
iveana lys
is 161 spec tropho tome tric me thod 160 trypsinogen 87
quaternary 13 spec tropho tome trically 162 tryptophan 10
sph ingomye l
ins 23 tyrosine 10
Rg roup 8 spo ttes t 159
rabbitf
ish 184 starch 83 , 88 ul
trav
ioletrad ia
tion 53
radioac tive
lylabeled 15 statis t
ica ltoo l 16 undigested ma ter
ial 37
rancidity 150 step -w ise manne r 29 uneatenfeed 4
recommendedd ietaryleve
ls 17 stereom ic roscope 150 union
izedammon ia 93
reco rdkeep ing 171 stero id ho rmones 2 1 unsaturated 23 , 28
redb loodce l
ls 49 stero ids 22 unsaturatedfa t
tyacids 2 1
regulationo f metabolism 45 stero ls,phospho lipids 2 1 unsaturat
ion 25
relat
ivegu tlengths 82 stockenhancemen t 201 urea 153 , 161
re
la t
ivehum idity 144 stock ingdens ity 170 ureaandammon ia 14
renalca lc
inosis 59 stomach 79 ,8 1 ureaseac t
ivity 161
reproduc t
ion 42, 43 storagefo rm 36 ur
inary 43
respiration 43 straine rs 78 ur
ine 45
returnsonfeeds 216 , 217 struc turalcon f
igu ra t
ion 34 u
til
izat
iono fca rbohydrates 33
r
ibo f
lavin 46 sub -ce llula r memb ranes 23
r
icke ts 52 sucke rs 78 vanadium 57
suc rose 35 vascula
rtissues 53
salinity 26 , 176 sulfu r6 1 v
il
li
form 80
samp ling me thods 171 sunsh inev i
tam in 52 v
iscousfluids 37
sapon if
ica t
ion 157 supp lemen ta
l 99 v
ision 94
satiation 92 supp lemen ta
lfeeds 171 v
itamin A 51
satu rated 23 supp lemen taryfeed ing 4 v
itaminand m inera
lmix
tures 104
scolios is 50 surviva l 172 v
itaminde f
iciencysymp
toms 54
scu rvy 50 surviva lra te 165 v
itamin K 53
seaho rses 200 susta inab il
ity 2 v
itamins 45
seasona lva riation 27 susta inab ley ield(MSY ) 220
seaweeds 37 swa llow ingo rre jec t
ion 94 warm wa ter2 1
secch id isk 176 symp toms 45 , 57 waterflowra te 44
seconda ry 12 systema ticeva lua tiono ffeeds
tuf
fs 165 waterqua l
ity 172 , 176
sec reto ryo rB -cel
ls 85 waterqua l
ityands t
ress 44
selen ium 57 tasterecep to rs 94 waters tabil
ity 3 , 151
selen ium -dep leted 52 tastetes t
ing 94 watertempe ra ture 44
sem ipu rified d iets 61 TBA 158 watertempe ra turefluctua
tion 93
senseo fsme ll 150 tempe rature 26 , 176 water-soluble 45
senseo ftas te 150 terres trialan ima ls 25 waxes 22
sens itivity 158 tertiary 13 wel
l-balanced 3
sens itivitytoshock 46 tetrahyd rofolicac id 50 wetex t
rus ion 133
series 25 thiam in 45 wholebodyp rotein 16
sex 22 thiam inase 128 w
indow -paneoys ter 201
sho rthand abb rev
iat
ion 24 thiok inases 59 woundt issue 50
shrimpd iet 210 t
ige rsh rimp 193
shrimpsing row -outponds 195 t
igh t 83 xy
lose 34
si
licon 57 t
ilap ias 181
simp led iffusion 86 t
in 57 z
inc 57
, 159
simp le p roteins 1 1 t
issuep roteinre tent
ion 18 z
incbioavai
lab
il
ity 61
simp lesuga rs 34 t
issues torage 55 zoop
lankton 181
sinkingfeeds 174 t
itration 160 zoop
lanktons 82
s
izeo rage 53 ti
trime t
ric me thod 157, 160
skele talabno rma l
it
ies 60 tocophe rol 52
skind iso rders 49 tocophe rolace ta te 52
skinles ions 49 ,51 totalphys ica lp roduct 211
, 212
, 214
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