nutritional requirements Kamal Moun and Kurt !" #osentrater$ %ort& 'entral !gri(ultural #esear(& )a*oratory$ +S,!-!gri(ultural #esear(& Servi(e Mi(&ael )" Bro-n$ ,epartment o. /ildli.e and Fis&eries S(ien(es$ Sout& ,a0ota State +niversity 1ilapia is one o. t&e most -idely (ultured .is& in t&e -orld" 'urrently$ .armed tilapia represents more t&an 234 o. -orld tilapia produ(tion 5F!6$ 20097$ and t&is (ontri- *ution &as *een e8ponentially gro-ing in re(ent years" Several .a(tors &ave (ontri*uted to t&e rapid glo*al gro-t& o. tilapia" 1ilapia are easily (ultured and &ig&ly adapta*le to a -ide range o. environmental (onditions" 1ilapia .eed on a -ide variety o. dietary sour(es$ in(luding p&ytoplan0- ton$ perip&yton$ 9ooplan0tons$ larval .is&$ and detritus" !dult tilapia are prin(ipally &er*ivorous *ut readily adapt to (omplete (ommer(ial diets *ased on plant and animal protein sour(es" :n t&e +nited States$ t&e most (ommonly .armed tilapia spe(ies are$ in order$ %ile 56reo(&romis niloti(us7$ Mo9am*ique 56" mossam*i(us7$ *lue 56" aureus7$ and &y*rids 5;reen$ 20067" 1&is pu*li(ation provides a *rie. overvie- o. environmental and nutritional requirements o. tilapia" EnvironmEntal rEquirEmEnts 1ilapia (an tolerate a -ider range o. environmental (onditions<in(luding .a(tors su(& as salinity$ dissolved o8ygen$ temperature$ p=$ and ammonia levels 5ta*le >7< t&an most (ultured .res&-ater .is&es (an" :n general$ most tilapia are &ig&ly tolerant o. saline -aters$ alt&oug& salinity toleran(e di..ers among spe(ies" %ile tilapia is t&oug&t to *e t&e least adapta*le to mar0ed (&anges 5dire(t trans.er$ >? parts per t&ousand in salinity7@ Mo9am*ique$ *lue$ and red*elly 51" 9illi7 are t&e most salt tolerant 5El-Sayed 20067" /it& t&e e8(eption o. %ile tilapia$ ot&er tilapia spe(ies (an gro- and reprodu(e at salinity (on(entrations o. up to 36 parts per t&ousand$ *ut optimal per.orman(e measures 5reprodu(tion and gro-t&7 are attained at salinities up to >9 parts per t&ousand 5El-Sayed 20067" 1ilapia are$ in general$ &ig&ly tolerant o. lo- dissolved o8ygen (on(entration$ even do-n to 0"> mgA) 5Magid and Ba*i0er$ >9237$ *ut optimum gro-t& is o*tained at (on(entrations greater t&an 3 mgA) 5#oss$ 20007" 1emperature is a maor meta*oli( modi.ier in t&ese .is&" 6ptimal gro-ing temperatures are typi(ally *e- t-een 22B ' 522B F7 and 29B ' 5?CB F7@ spa-ning normally o((urs at temperatures greater t&an 22B ' 522B F7" Most tilapia spe(ies are una*le to survive at temperatures *elo- >0B ' 530B F7$ and gro-t& is poor *elo- 20B ' 56?B F7" Blue tilapia are t&e most (old tolerant$ surviving at tempera- tures as lo- as ?B ' 5C6B F7$ -&ile ot&er spe(ies (an tolerate temperatures as &ig& as C2B ' 5>0?B F@ 5Sarig$ >969@ Morgan$ >922@ 'aulton$ >9?2@ Mires$ >9937" 6t&er -ater quality (&ara(teristi(s relevant to tilapia (ulture are p= and ammonia" :n general$ tilapia (an tolerate ta*le >" )imits and optima o. -ater quality parameters .or tilapia" parameter Salinity$ parts per t&ousand ,issolved o8ygen$ mgA) 1emperature$ 'B p= !mmonia$ mgA) range +p to 36 ,o-n to 0"> ?DC2 3"2D>> +p to 2"> optimum .or gro-t& +p >9 E3 22D29 2D9 F 0"03 re.eren(e El-Sayed 520067 Magid and Ba*i0er 5>9237@ #oss 520007 Sarig 5>9697@ Morgan 5>9227@ Mires 5>9937 #oss 520007 El-S&a.ey5>99?7@ #ed- ner and Sti(0ney$ 5>9297 S o ut & , a0ota Stat e univerSi ty G So u t& , a0 o ta 'o o per at ive e8t enSi on Servi 'e G uS,a a p= range o. 3"2 to >>$ *ut *est gro-t& rates are a(&ieved *et-een p= 2 to 9 5#oss$ 20007" !mmonia is to8i( to tilapia at (on(entrations o. 2"3 and 2"> mgA) as unioni9ed am- monia$ respe(tively$ .or *lue and %ile tilapia 5#edner and Sti(0ney$ >929@ El-S&eri. et al"$ 200?7 and depresses .eed inta0e and gro-t& at (on(entrations as lo- as 0"> mgA) 5El- S&eri. et al"$ 200?7" 6ptimum (on(entrations are estimated to *e *elo- 0"03 mgA) 5El-S&eri. et al"$ 200?7" nutritional rEquirEmEnts 1&e lo- trop&i( level and t&e omnivorous .ood &a*its o. tilapia ma0e t&em a relatively ine8pensive .is& to .eed$ unli0e ot&er .in.is&$ su(& as salmon$ -&i(& rely on &ig&- protein and lipid diets *ased on more e8pensive protein sour(es li0e .is& meal" :n addition$ tilapia are similar to (&annel (at.is& 5:(talurus pun(tatus7$ in t&at t&ey (an tolerate &ig&er dietary .i*er and (ar*o&ydrate (on(entra- tions t&an most ot&er (ultured .is&" 1o ensure &ig& yield and .ast gro-t& at least (ost$ a -ell-*alan(ed prepared .eed is essential to su((ess.ul tilapia (ulture" Slig&t variations e8ist among tilapia spe(ies$ *ut nutrient requirements are primarily a..e(ted *y t&e si9e o. t&e .is&" protein Fis& do not &ave a spe(i.i( requirement .or (rude protein 5'H7 per se$ *ut rat&er t&ey need a (om*ination o. essential amino a(ids" 1&ere.ore$ t&e pro.ile o. dietary pro- tein is important -&en .ormulating diets .or tilapia" ,ietary proteins are used (ontinuously *y .is& .or maintenan(e$ gro-t&$ and reprodu(tion .un(tions" /&en .ed in e8(ess$ protein may *e used as energy@ &o-ever$ t&e latter .un(tion is not desira*le *e(ause o. t&e e8pensive (ost o. proteins" 1&e protein requirement o. tilapia de(reases -it& age and si9e 5ta*le 27$ -it& &ig&er dietary 'H (on(entrations re- quired .or .ry 530D3647 and uvenile 530DC047 tilapia *ut lo-er protein levels 52?D3047 .or larger tilapia 5/in.ree ta*le 2" Hrotein requirements 5dry *asis7 o. tilapia o. various spe(ies and si9es" spe(ies 6" niloti(us si9e 5g7 0"3-6?"3 C3"0D26"3 0"?CD22"? C0"CD>0?"0 6" niloti(us 8 6" aureus 6" aureus S" mossam*i(us 2>"3D?>"3 2"3D>6"6 >"?3D?"3 require- ment 54 o. diet7 C0 C0 C0 30 2? 36 C0 Siddiqui et al" 5>9??7 1-i*ell and Bro-n 5>99?7 /in.ree and Sti(0ney 5>9?>7 Iaun(ey 5>9?>7 re.eren(e !l =a.id 5>9997 and Sti(0ney$ >9?>@ Iaun(ey$ >9?2@ !l =a.ed&$ >999@ Siddiqui et al"$ >9??@ 1-i*ell and Bro-n$ >99?7" !s -it& ot&er -arm- -ater .is&$ tilapia require >0 essential amino a(ids t&at need to *e supplied *y t&e diet 5ta*le 37" Essential amino a(id requirements (an *e met *y t&e use o. a *alan(e o. *ot& plant and animal proteins$ and i. ne(essary$ *y t&e in(lusion o. synt&eti( amino a(ids in t&e (omplete .eed" lipids ,ietary lipids provide a maor sour(e o. energy$ .a(ilitate t&e a*sorption o. .at solu*le vitamins$ play an important role in mem*rane stru(ture and .un(tion$ serve as pre(ursors .or steroid &ormones and prostaglandins$ and serve as meta*oli9a*le sour(es o. essential .atty a(ids" /in- .ree and Sti(0ney 5>9?>7 .ound t&at .or tilapia up to 2"3 g$ t&e optimum dietary lipid (on(entration -as 3"24$ de(reas- ing to C"C4 .or .is& up to 2"3 g" Iaun(ey 520007 suggested t&at to ma8imi9e protein utili9ation$ dietary .at (on(entra- tion s&ould *e *et-een ? and >24 .or tilapia up to 23 g$ and 6 to ?4 .or larger .is&" !s -it& most .is&$ tilapia appear to &ave a requirement .or n-6 5linolei(7 .atty a(ids$ and to a lesser e8tent$ a requirement .or n-3 5linoleni(7 .atty a(ids" ,ietary lipids s&ould supply at least >4 o. n-6 .atty a(ids 51es&ima et al"$ >9?27" /&en dietary lipids (ontain (on- ta*le 3" !mino a(id requirements 5dry *asis7 o. tilapia"J amino a(id !rginine =istidine :soleu(ine )eu(ine )ysine Met&ionine( H&enylalanined 1&reonine 1ryptop&an Kaline a 4 o. dietary protein 6" niloti(usa C"20 >"22 3">> 3"39 3">2 2"6? 3"23 3"23 >"00 2"?0 6" mossam*i(us* 2"?2 >"03 2"0> 3"C0 3"2? 0"99 2"30 2"93 0"C3 2"20 Santiago and )ovell 5>9??7" #equirements estimated *y t&e dose-response met&od" :nitial -eig&t 5>3 to ?2 mg7 and .inal -eig&t 5>03 to>20 g7" * Iaun(ey et al" 5>9?37" #equirements estimated *y t&e -&ole *ody and mus(le amino a(id (omposition met&od" ( 'ystine 0"3C4 o. dietary protein" d 1yrosine >"294 o. dietary protein" J %ote t&at t&e met&ionine requirement depends on dietary (ys- tine (on(entration" :. (ystine is not in(luded in t&e diet$ part o. met&ionine -ould *e used to synt&esi9e (ystine and i. (ystine is supplied *y t&e diet$ t&e requirement o. met&ionine is redu(ed" )i0e-ise$ tyrosine (an *e synt&esi9ed .rom p&enylalanine and its supply *y t&e diet -ould de(rease t&e dietary requirement o. p&enylalanine .or ma8imal gro-t&" 2 sidera*le amount o. polyunsaturated .atty a(ids$ attention s&ould *e given to prevent o8idation o. t&e dietary lipids" 1&e resulting produ(ts o. lipid o8idation are to8i($ redu(e t&e availa*ility o. ot&er nutrients$ and impa(t t&e quality o. .is&-.les& produ(ts" 'ar*o&ydrates Fis& do not &ave a spe(i.i( requirement .or (ar*o&y- drates$ *e(ause amino a(id and .atty a(id pre(ursors (an supply t&e required glu(ose via glu(oneogenesis" 1&is does not imply t&at (ar*o&ydrates s&ould not *e in(luded in tilapia diets$ &o-ever" 'ar*o&ydrates provide a relatively ine8pensive sour(e o. energy (ompared to protein$ and t&eir in(lusion (an improve t&e quality o. pelleted .eeds" 1ilapia (an e..e(tively utili9e (ar*o&ydrate levels up to 30 to C04 in t&e diet$ -&i(& is (onsidera*ly more t&an most (ultured .is& 5!nderson et al"$ >9?C@ 1es&ima et al"$ >9?37" Fi*er is usually (onsidered indigesti*le$ as tilapia do not possess t&e required en9ymes .or .i*er digestion 5alt&oug& some (ellulase a(tivity .rom mi(ro*es &as *een .ound in t&e gut o. 6" mossam*i(a7 5Sa&a et al"$ 20067" For t&is reason$ and to attain ma8imum gro-t&$ (rude .i*er levels in tilapia diets s&ould pro*a*ly not e8(eed 34 5!nderson et al"$ >9?C7" 1&e relations&ip *et-een (on(entrations o. dietary protein and energy is important in .is& nutrition" ,iets s&ould *e *alan(ed to ma8imi9e t&e use o. protein .or gro-t& *y providing optimal amounts o. energy as (ar*o- &ydrates and lipids" 1&e ratio o. protein to energy 5H:E@ mgA K(al7 varies -it& .is& age and si9e" For tilapia t&e optimum ratio .or gro-t& varies *et-een 6? and >23$ depending on spe(ies and si9e 5/in.ree and Sti(0ney$ >9?>@ S&iau and =uang$ >9907" vitamins and minerals Kitamins and minerals are essential .or normal .is& meta*olism" Kitamin and mineral supplementation in t&e .orm o. premi8es may *e *ene.i(ial in intensive systems$ al- t&oug& most o. t&ese requirements are usually met natural- ly in e8tensive and semi-intensive pond (ultures" =o-ever$ spe(i.i( requirements are not e8a(tly 0no-n .or all vitamins" 1a*le C gives ranges o. vitamin requirements t&at &ave *een determined .or di..erent tilapia spe(ies and &y*rids" Be(ause o. t&e limited 0no-ledge and t&e un(ertainty regarding vitamin requirements$ it is di..i(ult to ma0e general re(om- mendations as to -&at t&e optimal (on(entrations s&ould *e$ *ut general minimum levels are (ommonly applied to .eeds" ;eneral mineral requirements .or various tilapia spe(ies are presented in ta*le 3" Basi( 0no-ledge o. mineral to8i(ity and intera(tions among minerals is ne(essary -&en supplementation is made" 1&oug& not spe(i.i( to tilapia$ e8(essive supplies o. (ertain minerals (an (ause de.i(ien(ies o. ot&ers and in e8treme (ases to8i(ity to .is&" For e8ample$ &ig& dietary (al(ium (an (ause de.i(ien(ies in p&osp&orus$ 9in($ iron$ and manganese" 6n t&e ot&er &and$ it &as *een s&o-n t&at 9in($ (opper$ and selenium$ at &ig& (on(entra- tions$ (an *e to8i( to various .in.is& spe(ies" 'on'luSion !lt&oug& tilapia is one o. t&e most studied (ultured .is&$ spe(i.i( dietary requirements are still la(0ing and t&e intera(tion among nutrients and -it& (ulture (ondi- tions are not (ompletely understood" :n addition$ nutrient requirements may vary among tilapia spe(ies" 1&us$ readers are reminded t&at t&is pu*li(ation is only a *rie. introdu(- tion to tilapia requirements$ most o. -&i(& -ere derived under (ontrolled environmental (onditions" More detailed in.ormation (an *e .ound in t&e additional re.eren(es listed *elo-" 3 ta*le C" Kitamin requirements 5dry *asis7 o. tilapia o. various spe(ies and si9es" vitaminAspe(ies Kitamin B> 5t&iamine7 6" niloti(us 6" mossam*i(us L 6" urolepsis &onorum Kitamin B2 5ri*o.lavin7 6" aureus Kitamin B6 5pyrido8ine7 6" niloti(us L 6" aureus 6" mossam*i(us L 6" niloti(us si9e 5g7 D D >"C6DC2"0 0"2>D3>"6 0"23D2"2 0"23D3"C Hantot&eni( a(id 6" aureus %i(otini( a(id 5nia(in7 6" niloti(us L 6" aureus Biotin 6" niloti(us L 6" aureus Foli( a(id 6" niloti(us L 6" aureus Kitamin B>2 5(yano(o*alamin7 6" niloti(us L 6" aureus :nositol 5Myo-inositol7 6" niloti(us L 6" aureus '&oline 6" aureus 6" niloti(us L 6" aureus Kitamin ' 5as(or*i( a(id7 6" niloti(us Kitamn ! 5retinol7$ :+A0g 6" niloti(us 6" niloti(us L 6" aureus Kitamin , 5(&ole(al(i.erol7 6" aureus 6" niloti(us L 6" aureus Kitamin E 5to(op&erol7 6" niloti(us 6" aureus a e requirement re.eren(e 5mgA0g o. diet7 C 2"3 3 6 >"2D9"3a >3"0D>6"3* 6 >0 26D>2>( 0"06 0"?2 %# C00 %# >000 30 C20 3000 3?30D6920d %# 323 >0. >0D23e )im et al" 520007 )im and )eamaster 5>99>7 )im et al" 5>9937 Soliman and /ilson 5>992a7 S&iau and =sie& 5>9927 S&iau and =sie& 5>9927 #oem et al" 5>99>7 Soliman and /ilson 5>992*7 S&iau and Suen 5>9927 S&iau and '&in 5>9997 S&iau and =uang 5200>7 S&iau and )ung 5>9937 S&iau and Su 520037 #oem et al" 5>990a7 S&iau and )o 520007 !*delg&any 5>9967 Soliman et al" 5>99C7 Sale& et al" 5>9937 =u et al" 520067 6M'onnell and ;atlin 5>99C7 S&iau and =uang 5>9937 Sato& et al" 5>9?27 #oem et al" 5>990*7 D 0"2>D>2"C 2"2D2?"? 0"9?D3"0 0"C>DC"> >"0DC2"2 0"3>DC"? 2"0D2>"9 0"62D6"> 0"36DC"3 >"0D>?"0 >>"CD33"> >"6D9"3 2"3DC2"0 0"29D22"2 0"C9D2"? 2D>9"2 2?4 protein diet$ * 324 protein diet$ ( .ed eit&er glu(ose or de8trin diets$ d depending on -eig&t gain o. .is&$ .ed eit&er 3 or 64 lipid diet$ . .ed a 34 lipid diet" J %#: not required" C ta*le 3" Mineral requirements 5dry *asis7 o. tilapia o. various spe(ies and si9es" mineralAspe(ies maor 5gA0g o. diet7 'al(ium 6" aureus H&osp&orus 6" niloti(us 6" aureus Hotassium 6" niloti(us L 6" aureus Magnesium 6" niloti(us 6" aureus tra(e 5mgA0g o. diet7 Nin( 6" niloti(us 6" aureus '&romium 6" niloti(us L 6" aureus >"23D>3"> 0"33DC"0 2 >C0 S&iau and )in 5>9937 S&iau and S&y 5>99?7 3">D22"> ?"3D23"> 20 30 M('lain and ;atlin 5>9??7 Eid and ;&oneim 5>99C7 20"0D3C"C 0"3D9"0 0"39 0"3 ,a*ro-s0a et al" 5>9?97 #eig& et al" 5>99>7 0"22D3"3 2D3 S&iau and =sie& 5200>7 6">D32"0 D F9 >0 /atana*e et al" 5>9?07 Kiola et al" 5>9?67 2"3D6>"3 2D2"3 #o*inson et al" 5>9?27@ 6M'onnell and ;atlin 5>99C7 si9e 5g7 requirement re.eren(e re.eren'eS !*delg&any !" E" >996" ;ro-t& response o. %ile tilapia$ 6reo(&romis niloti(us$ to dietary )-as(or*i( a(id$ )- as(or*yl-2-sul.ate$ and )-as(or*yl-2-polyp&osp&ate" I" /orld !qua(ult" So(" 22: CC9DC33" !l =a.ed&$ O" S" >999" E..e(ts o. dietary protein on gro-t& and *ody (omposition o. %ile tilapia$ 6reo(&romis niloti(us )" !qua(ulture #esear(&" 30: 3?3D393" !nderson$ I"$ !" I" Ia(0son$ !" I" Matty$ and B" S" 'apper" >9?C" E..e(ts o. dietary (ar*o&ydrate and .i*re on t&e tilapia 6reo(&romis niloti(us 5)inn"7" !qua(ulture" 32: 303D3>C" 'aulton$ M" S" >9?2" Feeding$ meta*olism and gro-t& o. ti- lapias - some quantitative (onsiderations" pp" >32D>?C" :n: Hullin #"S"K" and )o-e-M('onnell #"=" 5eds7$ 1&e Biology and 'ulture o. 1ilapia" :')!#M$ Manila$ 1&e H&ilippines" ,a*ro-s0a$ ="$ K" ," ;uent&er$ and K" Meyer-Burgdor.." >9?9" !vaila*ility o. various magnesium (ompounds to tilapia 56reo(&romis niloti(us7" !qua(ulture" 26: 269D226" El-Sayed$ !"M" 2006" 1ilapia (ulture in salt -ater: Envi- ronmental requirements$ nutritional impli(ations and e(onomi( potentials" Eig&t& Symposium on !dvan(es in %utritional !qua(ulture" %ovem*er >3D>2$ %uevo )eon$ Me8i(o" El-S&eri.$ M" S"$ and !" M" E)-Fe0y" 200?" E..e(t o. am- monia on %ile 1ilapia 56" niloti(us7 per.orman(e and some &ematologi(al and &istologi(al measures" Eig&t& :nternational Symposium on 1ilapia in !qua(ulture" 'airo$ Egypt" Eid$ !"E"$ and S" :" ;&onim ">99C" ,ietary 9in( requirement o. .ingerling 6reo(&romis niloti(us" !qua(ulture" >>9: 239D26C" F!6$ Food and !gri(ulture 6rgani9ation o. t&e +nited %a- tions" 2009" 2002 F!6 year*oo0" Fis&ery and aqua(ul- ture statisti(s" &ttp:AA---".ao"orgA.is&eryApu*li(ationsA year*oo0sAen" ;reen$ B"/" 2006" 1ilapia .ingerling produ(tion systems" pp >?>D2>0" :n: '" )im$ '" /e*ster 5Eds7" 1ilapias: Biology$ 'ulture$ and %utrition" Food Hrodu(ts Hress" Bing&am- ton$ %O" =u$ '" I"$ S" M" '&en$ '" =" Han$ and '" =" =uang" 2006" E..e(ts o. dietary vitamin ! or P- (arotene (on(entra- tions on gro-t& o. uvenile &y*rid tilapia$ 6reo(&romis niloti(us L 6" aureus" !qua(ulture" 233: 602D602" Iaun(ey$ !" >9?2" 1&e e..e(t o. varying dietary protein level on t&e gro-t&$ .ood (onversion$ protein utili9ation and *ody (omposition o. uvenile tilapias 5Sarot&erodon mossam*i(us7" !qua(ulture" 22: C3D3C" Iaun(ey" K" 2000" %utritional requirements" H" 322D323" :n: M" '" M" Beveridge and B" I" M(!ndre- 5eds"7 1ilapias: Biol-