Nutrition of the Domestic Cat: a Mammalian Carnivore

  u 8 p © 8 y l w  ll g 

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NUTRTON OF THE DOMESC CAT, A MAMMALAN CARNVORE M . Md d Q  gs Depatmet of Pysiologia Siees, Shool o Veteriary Mee, Uversty o Caoa, av Cafoa 

  M Depames of Pysologa Siees a Aimal Siees, vesity of afoia, Davs, afoia 66

CONTENTS INTRODUCTION  INTRODUCTION    EATING AND DRINING DRIN ING BEHAVI B EHAVIOR OR     Rhythms Rhythms n the Cat        Feeding Patten and Food Intake eulatin     . . . .     . .        . .  .  .    . . . . . . . . .    . . . . Water Requrements and Drnn Pattes Pattes . . 

ENERGY REQUIREMENTS REQUIREME NTS      abodate abodate        Lpd    . . . . .    ..   ...   ...    .  . .

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PROTEIN AND AMINO ACIDS.......................... Proten Requrement Requrement        Amno Am no Ad A d Requrements Requrements       

   . . . . . . . . . . . . .  .     .  . . . . .     .    . .  . . .  . .   .     . .     . . .

MINERALS MINERALS        .                  .            .     .         .            .               .  .        .    VITAMINS. VITAMINS ... .    . .  at-Soube Vtamns   .  .          . .  . . . .  . .     . . . . . . . . . . . . . .     .   . .    . .            . .  .   WateSoube Vtamn Vtamn       OTHER FOOD CONSTITUENTS .  

ESSENTIA FAY ACIDS. ACIDS .      Metabom  .   EA Defienc n at.  .   . .            . . .        . .    . .  .     .     .    . .  . . . . . . . . . .     EA Requrement  . . . . .  . . .  . . . . . . . . . .  . .    . . .   .  . . . . . .  . . . . .    . . . . . . .     .  . . . . . . . .  . . . .  .   

SUMMARy. SUMMARy . ..     

 

I

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545 545 546 547 547 547 55 55 552

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522

MaDONALD OGS & MOS

INTRODUCION

g r o . s w . e y i v l n e r o l a e u s n u n l a . a n o w s w r w e p m r o o F r f . d 6 e 1 d / a 6 1 o / l n 5 0 w n o o D . S 2 E 6 P 5 - A 1 C 2 y 5 : b 4 . d 4 e 8 i d 9 v 1 o . r r t p u s s N e . v c e c A R . u n n A

W or domescaed cvores eonn o he am am edae ed ae were ke n a ave or semdomesaed sae  he Eans a eas 30 eas ao . Dese ts on asocaon eween humans and cvores, rese noaon on he nurona requr requremen emenss o hese hes e mammas has een ackn a ckn un un  reave recen recen. . Ths revew ouses ouses on unque asecs o he nuton o he domes a Fels domesus, he mos hor horouh ouh suded mammaan mamma an voe. vo e. I s  s ou hess ha he domesc a s a ood moe or or sdn he eares n he nuron o cvores The ancesa orn o he domes a F domesus domesus and he rese dae o s domescaon are oen o dsue. Resoon o oh o hese qesons s hamered  he ac ha here were reave ew skeea chanes n he ca ae s domescaon domescaon The a a have een doesca doescae e  he Eans as ear as 3000 B u Zeuner (264) consders he he evdence amuous.  an case he a  a was cear cear domesaed and rea rearde rdedd as a sared anma  1600 B Cas were sared o he Ean oddess Bas o Buass and were rearded as he naon (35). Accordn o he Roman hsoan Dodous a  B he Eans ed her as on rea mk an ses o Ne sh, an he aed her anmas o her meas meas  seca souns  n he ass o mummed emamed a odes reovered n reen mes, he numer o as ke  he Eans Eans mus have een ro ro.. Durn he aer art o he nneeenh an ear weneh cenures mummed cas were exavaed a Buas Bu ass s and oher aes n E and were sread on on he and as as eer Th were so numeros ha he were aso eoed one onsn men sen o nand or erzer wehed nneeen ons (70. The Eans coeed and ke varous reconzae seces o cas nun F us he june a  F  he Aran w a However mos o he mumme skus reered o aove (0 were ound ound on examnaon o e smaer han F us u arer an F l and he domes a a   has een suesed ha he esen omesc a ma have een derve rom nerreedn o F slvesrus he Euroean wd ca, and F l  In ass a ssn n  he a, Car nne name he he domes a F us whh he desred as avn ohed a markns aaren he neeed or was ovous o he equa ommon mackere e (202, whch ame o reeve he esnaon o F oqu Thus he se name F omesus aears o e he mos arorae and ancsve name o escre he domes a The Feoae Vverrs Haens an es) vere rom he oher meers o he order Cora reave ear n her evouonar deveo men e durn he ae Eocene o he ear Pesoene. Aer he Fes dvered rom he Vverd ancesr, he undewen ra evouon o u seazed orms whh have no hane or mons o ears 

522

MaDONALD OGS & MOS

INTRODUCION

g r o . s w . e y i v l n e r o l a e u s n u n l a . a n o w s w r w e p m r o o F r f . d 6 e 1 d / a 6 1 o / l n 5 0 w n o o D . S 2 E 6 P 5 - A 1 C 2 y 5 : b 4 . d 4 e 8 i d 9 v 1 o . r r t p u s s N e . v c e c A R . u n n A

W or domescaed cvores eonn o he am am edae ed ae were ke n a ave or semdomesaed sae  he Eans a eas 30 eas ao . Dese ts on asocaon eween humans and cvores, rese noaon on he nurona requr requremen emenss o hese hes e mammas has een ackn a ckn un un  reave recen recen. . Ths revew ouses ouses on unque asecs o he nuton o he domes a Fels domesus, he mos hor horouh ouh suded mammaan mamma an voe. vo e. I s  s ou hess ha he domesc a s a ood moe or or sdn he eares n he nuron o cvores The ancesa orn o he domes a F domesus domesus and he rese dae o s domescaon are oen o dsue. Resoon o oh o hese qesons s hamered  he ac ha here were reave ew skeea chanes n he ca ae s domescaon domescaon The a a have een doesca doescae e  he Eans as ear as 3000 B u Zeuner (264) consders he he evdence amuous.  an case he a  a was cear cear domesaed and rea rearde rdedd as a sared anma  1600 B Cas were sared o he Ean oddess Bas o Buass and were rearded as he naon (35). Accordn o he Roman hsoan Dodous a  B he Eans ed her as on rea mk an ses o Ne sh, an he aed her anmas o her meas meas  seca souns  n he ass o mummed emamed a odes reovered n reen mes, he numer o as ke  he Eans Eans mus have een ro ro.. Durn he aer art o he nneeenh an ear weneh cenures mummed cas were exavaed a Buas Bu ass s and oher aes n E and were sread on on he and as as eer Th were so numeros ha he were aso eoed one onsn men sen o nand or erzer wehed nneeen ons (70. The Eans coeed and ke varous reconzae seces o cas nun F us he june a  F  he Aran w a However mos o he mumme skus reered o aove (0 were ound ound on examnaon o e smaer han F us u arer an F l and he domes a a   has een suesed ha he esen omesc a ma have een derve rom nerreedn o F slvesrus he Euroean wd ca, and F l  In ass a ssn n  he a, Car nne name he he domes a F us whh he desred as avn ohed a markns aaren he neeed or was ovous o he equa ommon mackere e (202, whch ame o reeve he esnaon o F oqu Thus he se name F omesus aears o e he mos arorae and ancsve name o escre he domes a The Feoae Vverrs Haens an es) vere rom he oher meers o he order Cora reave ear n her evouonar deveo men e durn he ae Eocene o he ear Pesoene. Aer he Fes dvered rom he Vverd ancesr, he undewen ra evouon o u seazed orms whh have no hane or mons o ears 

L NUTTON

g r o . s w . e y i v l e n r l o a e u s n u n l a . a n o w s w r w e p m r o o F r f . d 6 1 e d / a 6 1 o / l n 5 0 w n o o D . S 2 E 6 P 5 - A 1 C 2 y 5 : b 4 . d 4 e 8 i d 9 v 1 o . r r t p u s s N e . c v e c A R . u n n A

53

Th cot ratv o th Foda th Canoda whh ncud Pro cyond, Urd, urd Cand and Mutd xhbt much ratr d vrty than th Fod F oda. a. Th Urd bar ar ar w w  rconzd a omnvor omnvor . Th moar tth o th Urd ar onatd and  nam o  crown  urd a charactrtc whch ndcat an omnvorou or hrbvorou dt Th Procyond o whch  raccoon  a tyca mmbr ar ao omnvo rou. rou. Th urd, whch robaby ar an ohoot o th Urd ar rr ntd by e r and ratr anda whch ar ct hrbvor. contrat t o th Canoda a mmbr mmbr o th Foda Foda ar ar h-atr h-atr    . In contrat trct trct cvor o  o  t  car that th th ordr ordr Cvora Cvora contan mamma w w  th wdy wd y dvrnt dtary att att.. Th ordr ordr may ao rovd rarchr rarchr wth an oortunty to dtrmn whthr adhrnc to a cad dt durn vouton nducd or conbutd to chan n mtaboc athway and nutr tona rqurmnt On coud antcat that th chan mht b x rd n th Foda but not n h Canoda.  comaron o th nutona rqurmnt o th cat and th do a rrntatv mmbr o  Foda and Canoda uort th th that

D

D ECEN

Raccoon

Bar

Pada

Wasl

Gt

Ca

POCN MOCENE

OOCENE EOCENE EOCE Figure  he lationship of the cat to other members o the order Civora (Aer Res. 165, 20).

524

g r o . s w . e y i v l e n r l o a e u s n u n l a . a n o w s w r w e p m r o o F r f . d 6 e 1 d / a 6 1 o / l n 5 0 w n o o D . S 2 E 6 P 5 - A 1 C 2 y 5 : b 4 . d 4 e 8 i d 9 v 1 o . r r t p u s s N e . c v e c A R . u n n A

MaDONALD OGS & MOS

spealzaton onsstent wth the evolutonary nuene o a strt vorous det has oured n the at Examples o ths spealzaton, whh wll be dsussed more ully later, are: () The a has lmted alty to regulate atabol enzymes o amno ad metabolsm, whh auses the at to requre a hgher level o deay pen or manenane han he dog. b The at does not synthesze nan om typtophan beause o a very atve alteate degradatve pathway. (c) The dog but not the at onverts arotene to vtamn A and may thereore obtan ts vtamn  requrement rom plants (d The dog but not n ot the at) an onvet enough lnole lnole ad to arahdon arahdon ad, and an d may thereore mee ts aahdon ad reuremens rom plan soures (e) The dog an synthesze suent taurne rom ystene and an onserve taurne when lmted by usng glyne onjugaton The at has a lower apaty to synthesze taurne than the dog and s unable to use the alteate lyne onugaton. Thus unlke the at, the dog an meet ts taurne requrement om deay sulur amno ads suppled rom plan soures All he above nutrents that the at annot synthesze are ound n a vorous det.

EAING AND DRINKING BHAVIOR

hh  h  Many bologal vaables that exhbt rhythmty n most mammals (15 60) do not appear to do so n he domes a (102 189 241). The behavor pattes that do not show daly rhythmty nlude eedng and drnng (3 17246) atvty (102)and sleep-wake (59). Thus,  the domest at s allowed to eat ad lbtum, t wll eat many (-20) small eals per day, and these wll be approxmately evenly dstrbuted between the lght and dark (11171 246) Core body emperaure s normally radan n most pods (11171 mammals but t does not appear to vary on a daly bass n the at (102). Ths vrtual absene o radan rhythmy appears to be a unton o the at's natural behavor as a predator; radan rhythmty s also absent n ertan other elnes suh as the bobat. There s some evdene that these rhythms are also less rgd n he dog, whh s a less st vore than s the a (60). The la o many o the usual radan rhythms does not mean, however, that ats have no bologal loks. Instead t appears that many o ther atvtes smply ae not oupled to ther ologal loks. ats reprodutve yls depend upon day length to trger the emale estrus yle (19 63) t has been shown that he most proouned estus n the at s ndued by a perod o extended daylght (more than 12 hours aer a perod o short days 9). n nature ths ours wth the hane o seasons, whh establshes the breedng season n the late wnter and early sprng; however t an be ndued n the laboratoy durng any season by lghtng manpulatons (119) Also erebro spnal lud (F vasopressn onenaton vaes as a radan rhythm o

L UO

g r o . s w . e y i v l n e r o l a e u s n u n l a . a n w o r w s w e p m r o o r F f . d 6 1 e d / a 6 o 1 / l n 5 0 w n o o D . S 2 E 6 P 5 - A 1 C 2 y 5 : b 4 . d 4 e 8 d 9 i 1 v . o r r t p u s N s . e v c e c A R . u n n A

525

slighly greater han 24 hours. This rhyhm is characerized by high dayime values and low nigh values 189) As in oher mamals 190) melaonin concenraions in ca CSF exhibi a prominen daily rhyhm ha is characezed by high levels duing he dark riod and low levels durng e lig hous 189) However te CSF melao nin ae splis ino several componens as he ime in consan ligh is inceased he fac ha he melaonin hym in e cas sised dung exposue o consan lig is somewha enigmaic because in mos mamals arifcial ligh o moderae inensiy suresses melaonin producion so ha no rhym exiss 190). he sling o e SF melaonin no a leas wo disinc componens durng consan ligh may repesen he disassociaion of muliple crcadian oscillaos nomally involved in he geneaion of his hormone  rhyhm. More work on he cicadian rhyhms o cas needs o be done o esablish wich rhyms ae inrinsically lined o eeding behavior and o esablsh wheher he exe he drvin orce or he behavioral pae ha esul in er naul pedaoy behavio

Fdg  d Fd k g he predaoy behavior o the cat has been sudied and descbed in grea detal 145). The eral ca es omesus eas a wide varey o prey, which varies seasonally and includes mice ras rabbis bids liards insects; and oher animals 51 72 74 154). Mos wild felids (e lion is one ecepion) ae soliay huners ha rely on craine and peed o cach heir pre  74 219) I is easy o see ha e caloric conen o each of heir meals may vary considerably ice are he mos common prey or he domesic ca (51 72), nd an average mouse consiss of abou 30 kcal 71) hus he average ca would requie  meals o one mouse each per day t is ofen hough ha domesic ca would naurally ea large infrequen meals, as do lions 74 219) Grouing all ca families ogeher, researchers usually descibed he ca feeding pae as inermien or occasion 225)  has now een shown, however ha uncaged cas  or cagd cats ed puied dies 131) o commercial dies 128 131 171 172) will ea 10-20 imes a day if hey are given he food ee-choice. As many of hese meals will be eaen duing e dar period as will be eaen during the ligh period (11 172). Normally a slighy greaer uani o ood and a lighy arger number o meas are eaen during he eriod when he animals are given fresh food whehe i is ligh or dark 131 172). I has been shown for boh caged 50 128) and uncaged 133) cas ha he ore or a ca u do o obn a ea he eer eal ad he larger he meal sie  w ea. his same phenomenon occurs in oer speces 4

132) dmec 1) showed ha pedaoy behavio akes precdence ove con summaory behavior cas will sop eaing highly palaable ood o go ill a ra

56

g r o . s w . e y i v l n e r o l a e u s n u n l a . a n w o r w s w e p m r o o F r f . d 6 e 1 d / a 6 o 1 / l n 5 0 w n o o D . S 2 E 6 P 5 - A 1 C 2 y 5 : b 4 . d 4 e 8 d 9 i 1 v . o r r t p u s N s . e v c e c A R . u n n A

MacDOAD OGS & MOS

eve houh he wi he o ac o her iia foo raher ha ea he ra Ths behavor aers to have the fncona vae of ncreasn food nt b rovdn for mte s when the oortn rses. Kaare 18 a Hrsch e a 1 reore ha whe ca ies were dued wh cefour 8 or ao 1 he cas ese a consan qant of food and herefore fewer caores raher ha es more food so that the wod obtan a constan qant of caores Kanre 18 concded hat ths mht e true because cas are cvores an the caorc coe o her oo sul wou o var ea  aure. She suese that cats ma herefore reae ther foo nae on the ass of u rather than caores Athoh both sdes consdered aatabt as a ossbe factor t was dsmssed becase tota food nte dd not decrease when there was ncreased nondestbe matera n the det Cref examnaton of the resuts of Hrsch et a 1 however, shows that he cats' food nae decrease on the frst da after the receved he hher eves of aon more than t d on sbseqent das Aso, food ntke wa reat ncreased the frst da after the cas were reue o he aofree ie whch caes ha aaa certan was one factor Mos re anmas coan a eas 45 ca dr weht; herefore, he asa et of r ca foo which coas 34 ca r weh shol e cosere area caorca e or he ca whch has a short asonestna ac an therefore a rad rans tme. I s osse tha the tota vome of det nesed was becomn somewhat mtn before the des were caorca due. Mufor  hoe 7 sowe ha whe cas are fed socaoric ets, he ecrease heir iake of a ie ha s ow  aaa  ecreas oh ea sze a ea frequec Casoua 4 showed that cas comensae effecve for water duon of he et b ncreasn ther ntak of the caorca dte dets. Kane et a 131 used oh ue a coercia ies of oo aaa u wih iffere caoric conens an showed tha cats quc adjust the qat of food the nes o aian a que cosa caoric iae Ahouh here s hh variai  ean ates amon cats ad ahoh he cat s more sesve o the hsca form and taste of ts det than re the rat, do, and man other anmas, the cat oes have mechasms o coro caoric intake. Ths is conssent wh reos ha cas have ewer rolems wh oes ha dos whe boh are ve hh aaae dets 116 71. Anan  Broec 6 showe ha cas ha are aera esoe  he venomeda hothaams VMH become herhac and obese, whereas hose esoe n the atera hothaams H) becoe ahac. Other wor ers have show ha esios i he VMH 233, 235eriaqueuca ra maer 34 rhnencehaon 161a orsa eraqueuca ra maer 36 resu n herhaa n cats Ths the neroathwas nvoved n the contro offood ntke n cats aer to be smr to those of other anmas One mht exec,

L NUTRTON


5

however, that some of the metabolc cues would be dfferent n the cat, a cvore, than n the rat, an omnvo Although ltle s known aout meabo lte medaton of satety n the cat, t has been shown hat ow oses of eoyucose (0-1 m/k body weht ncreased sho te (13 hour fo ntake fourfold (3, whch suggests that he cat may have glu costatc, shoterm conols of feedn as do other mammas. I appears, n fact, hat ower doses are requre to prouce hs effect n the cat than n other anmals; ansaton of deoxYDucose at the ose evels use for e rat (0 m/k cause vomtn (13 an prouce a sopoc effect (11, whch resulted n a ecrease n food ntake. ats respon classcally to a leeaverson paradm usn thum chlor e (11 but appear to be eneray neophc rather than neophobc ke he rat. Cats w eat a nove et raer hn a famar et uness ey e put n a new envonment or a sessfu stuaton, n whch case they wl choose the famlar et (. As a cvore, the cat mght be epected to have taste preferences that dffer from those of omnvores and herbvores Indeed, erly sdes ncated that the cat dd not detect (185 or select (40 sweet soutons. Other stues ndcate hat cats woud select sweet substaces ony f they were n duted mlk (8 or n sane solutons , because the cat has a negatve response to pure water solutons (, 8. Although t was repoed that saccharn solutons were seecte by cats (63, Baroshuk et a (3 found that cats reecd boh saccharn and cyclaate solutons. The work by Beauchamp et al (4 suppors e ea at cats are neual to sus wheer n water or sane. These workers aso foun that cats show a preference for sucrose and actose n a luted mlk souton, bu a mechansm for s phenomenon has not yet been eluate. Beauchap et a (4 coned and extene e work of Boueau  coworkers (3, 5 by showng that e cat has a postve taste preference for proten hydrolyzates, meat eacts, and ceran free amno acds such as alanne, prone, ysne, an hstdne. Ths demonsated the practca sgnf cance of the elegant work of Boudreau and coworkers, who showed that amno acs tat ect an ncrease n spke output from gencuate ganglon che moresponsve oup  unts were prefered n a preference test, whe amno acs ha ecrease roup  schae were avoe (3,  Anmal pds mprove the palatablty of cat dets  8. Acceptance by cats of dets contanng 5 fat derved from dfferent sources was reported (. These auors tested bleached aow versus sx ateate fats n a twochoce test All cats preferred dets based on bleached tallow to those mae wth butter ol or chcken fat There was a tendency for cats to prefer unbleache talow and yellow grease over beached tallow, but cats showed no preference between partally hydrogenated vegetable fat and beached tallow. When pued dets contanng 10 , 5 , and 50 yelow grease were presente to cats, ey showe

528


MacDOAD OGS

 MOS

a prefrence for the des contanng 25% yellow grease over eher 10% or 50% yellow grease he level of fat n pufed des has a maked efec on texture of the de however so he cas may have prefeed dets conanng 25% fa paly ecase hese des have a more desrale exture than hose contanng 10% fa whch ae powde n natre, or hose contanng 50% a whch ae greasy (129) In a sseuen sudy these auhors found ha cats consumed more of a det made wth 40% leached allow han of those made wth eher 12 or 25% leached tallow. xpemental dets ased solely on coconu ol as a fa source have een found o e aversve o cas. Smlay dets conanng medm-chan tr glycedes ae aversve and the averson s sronger for tglycerdes wth 81 faty acds than for hose wh 12 faty acds he aversve nare of these des may e related o an aly of he cat to deect and avd low levels of m�dum-chan faty acds ha ae normally presen n these fas (M  MacDonald unpulshed resuls).

Water Requirements and Drinkng Patterns he ca s hough to have evolved as a dese anmal n pat ecase t can concentrae ts rne mre than he dog or an can  h enale he ca o suvve on less water u he physologc processes nvolved may also conr e o he ca's ssceply o felne rologcal syndrome (US) Even hough the ca s ale to overcome a prevos waer defc when gven sea waer o dnk (262)the cat wll sponaneously dnk consderaly less pre water nder the same condtons than he dog wll.  dog wll accratly replensh a water defc p to 8% of s ody wegh n js a few mnes; he ca wll not edaely make p a waer defc of as lttle as 4% of s ody wegh (2) lthogh the mnmum water reuremen for suvval s less for the cat than for the dog he weaker rs drve of the ca results n an ad lum ntake of water that s ofen too low o prevent cysalzaton of suve n the rne, whch akes he ca paclaly ssceple to U. lhogh he actual course of US may dfer from case o case  s clea ha ncreased water nake helps prevent recuences. he only effecve way o ncrease he ca's waer ntake s to feed a hghproen and/or a hghsal det or to pu water n he de Becase a hgh sal level (4% or more) s needed n the det to povde a susantal ncrease n waer ntake 133697he only praccal way o ncrease water nake s to add water to he food f fed a normal dry ca food, cas wll drnk aou 1-20 ml of water for each gram of d food (36 131 hs propoon (21) s near he popoon of waer n he anmals that are prey for cvores hs cas need or drnk vey lle waer f hey ae fed oher anmal ssues and poaly seldo dnk  hey ae ed canned ood ha s 5-80% water 3 ndeed, he mos effectve way o trea US n cats s o feed nohng  canned ca food ecause ths ncreases war nake and heey nceases ne

 UO

529

volume. A nutrtionaly compee caned food a ontains 8-9% waer should be even more effective in controing FUS in suscepible cats aloug no such food is curenty on the make in he US Te dog drnks only durng he day uness i is deydaed ut the ca diks durng bot te ig and dak peods 3. toug te number of dikng bous ae about te same as e numbe of meas eaen no coelaion has been found beween eating and dnking paes 3 unless he ca was eaing ony one or two meas a day 28

ENERGY REQUREMENS g r o . s w . e y i v l n e r o l a e u s n u n l a . a n w o r w s w e p m r o o r F f . d 6 e 1 d / a 6 o 1 / l n 5 0 w n o o D . S 2 E 6 P 5 - A 1 C 2 y 5 : b 4 . d 4 e 8 d 9 i 1 v . o r r t p u s N s . e v c e c A R . u n n A

he matue ody sze of memes of e genus Felis anges fom  kg o less fo e blacfooted cat (F nigrips), domestic ca (F domsticus) and the frcan wd cat e.g Flybica) to mor tan  kg for te puma (F oolo) . Other members of he famiy Fedae eg ions (Pnhr lo) and tiges (P. g may aan matue body weights in excess of 2 kg. o ou knowledge here ave been so few esimaes of energy requiemens over tis range of body weigts especiay a e uppe end that no geneaied ea tonship between body mass and enegy requremens can be made. Measuremens from whc estimae of enegy requiremens can e deived ae avalabe from wo sources: soterm measuemens of gaseous excange and ongetem measuemens of food intake and body weight canges eoeicaly eite approac sou give simia vaues fo maintenance enegy equiements but acvity is eremey imied n a respation cambe o cage and anmals must be accimaed o he cambe efoe reabe measure mens can be obaied. ue, mos measuemens on cas in cambes ave been of fasig basa ea production no of mainenance energy equie mens. Haldane 96 descbed a new apparatus for measuring espiaoy exchange in anmas and repoted daa on a snge 2kg cat of unspecfied age or physoogica stae. When e usua ema equivaen vaues ae appied to e C poducion he cacuated hea poducon was 9 kcakg ody weigh day In 938 Benedic 3 comped al measuements on basa ea poducion of cats up to tha time 38, 9 wic yeded a tota of 3 indivdual measuremens from fve souces e egession of basa ea poduion on by wegt appomated neaty and gave a mean vaue of  kca/kg body weighday for cats wi weights rangig fom .93 kg is mean vaue is simiar to te ea poduction vaue e cacuated from e respiaoy ex change data of Capener 4 fo cats in e postabsoptive sate he mean  SE was 522  2.8 kcakg body weigday. A somewa ige vaue of 66 ± 2.8 cag body weighday was found by E Kane J G ois and Q. R

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MaDOAD OGS & MOS

Ros (unpulshd daa). h ason fo h la valu n a han a od y an s no aan All h masumns dscussd aov f o fasn (asal) ha poduc on whch s ony on comonn o annanc f hs daa a o  ad n a fdn sysm h oh componns of mannanc ny acvy and ha ncmn mus  ncludd h mannanc ny  umn s nally on h od of 5  s h asal ha poducon If h appomaon of 1.5 ms asal ha poducon (58 kcalk ody wh/ da) s akn fo h ca h annanc n un s 8 kcal/k d whday. Vaous smas of h cas ny uns asd on masumn of food nak and funly calculad ny valus of foods (usually hyso cal fu vaus) hav n od n h lau. ycal valus (n kcak ody whday) a 616 (38) 80 (5) 6688 (68) and 60 (85 55) fo mau cas; 76 fo 3-onh-old cas (5) 9 fo 35-onh-old cas (252) and 160 (5) and 250(85, 155) fo kns h up smas fo mau cas a n asona an wh h valu of 87 kcalk/day calculad fo asal ha ducon I s dffcul o concl h l valus whch aoach asal ny ndu cnl Kndall and cks usd  dsl als  calcula h dsl ny (DE) uns of s adul cas fo annanc (136). Whn ny nak fo mannanc was ssd as a mass onn of uny h man valu was 76 kcal/day hn h mass onns of ody wh (k) w 0.5 and 067 h coffcns w 0 and 11 kcal. Alhouh no dfnv valu fo annanc n un can  dvd fom h foon daa a valu of aou 80 kcal DE/k ody wh/ day aas asonal. oad o oh aals h dffnc wn h cas mannac and asal ha oducon (.. ny ud fo acvy and ha ncn) aas cssv sall. s h cas ha losss n h dson asslaon and aols of fod al d no dff a ally fo oh amals  aas ha h acvy of h ca usd n h asumns mus hav n sla und h asal and annanc cond ons. any adul cas v h sson of no nan n unncssay cs u whh hs s an son o fac ans o  nvsad

Carbohydrates h mk of h un conans aou 35% caohyda as lacos whch accouns fo aou 0% of h aolal n of s lk (3 3 so caohyda s a majo coonn of a kns d Howv h adul ca ha consums only an anmal ssu d nss ll caohyda. Mos  al (167) pod ha adul cas ffcnly uld os coon solad ca ohydas ha w addd o a ma d Aan dson coffcns w

 UO


1

 fo a co stach 7 fo lactos an mo than 9 fo gucos sucos an n. gh concntatons of lactos pouc arha n som cats prsumaly caus thy ha naquat lactas actvty. Pat of th sapaanc of actos may hav n th rsut of fmntaton n th or gut caus th appant gstlty coffcnt of protn as uc th lactos tlaton of stach n th cal gans hat an co as sgnf cantly gat hn ths gans r fnly goun ath than coarsly oun coong aso ncas utlaton 7 Th gstty coffcnts of paally compos potato four an oo clulos  masur y Wl  uysnuyt 7 an foun to  .95 an 1 rspctvly. h valu fo potato flou s n agmnt th that pvously ct fo stach 17 ut th oo pouct caus of paa hyoyss as gst to a gat tnt han unhyrolys cluos hch as foun to  totaly ngstl h lvs of most omnvoous anmals hav to nyms gluconas an hoknas that cataly th phosphoryaton of glucos to gcos-phosphat. onas has a lo m o gucos hras gluconas has a hgh m an opats only hn th v vs a ag oa of gucos from h poal vn Th actvty of guconas n th vr of th cat s my lo has th actvty of hoknas s n th noal ang 1. vors t omnvoous tay hats .g omstc ogs possss oth nyms so t sms lly that thy hav a gatr capacty to han a hgh glucos ma han th cat hovr t os not appa that ths has n tst. n omnvoous anmas mama gluconognss occus n h postasop tv stat hch occus som hous aftr a mal n cvos ngstng ts of anma tss mama glconognss occrs n th asoptv phas m matly follog a ma caus th capacty to sto potn p s s mnma Th latonshps tn mamal guconognss an mals a smla n cvos an umnants aso guconognss s mo o lss manntly "stch on n oth goups of anmals 4 4 5. h may also  ffncs tn cats an omnvos n th atv mpotanc of vaous gluconognc pathays lvau  lan 27 sho that nhton of cytosolc phosphonopyruvat caoynas pss glu conognss fom sn n at hpatocyts ut not n cat hpatocyts hs nng suggsts that n h cat srn s convrt to glucos y a rout that os not nvolv pyruvat an th nym srn hyratas. o actvty of sn hyratas n th lvr of th cat has n pot 29 214 hch povs suppot for th noton that th s an altat pathay to pyuvat that poaly nvolvs hyoypyruvat. Rosl t al 24 suggst that hgh actvty of th st nym of ths pathay Lsrnpyuvat amnoans ras n mammals s assocat th ts of anma sh No ty qmnt fo caohyat has n monstat fo t cat Gong ns an ault cats at mantnanc poaly o not qu tay

532

MaDOAD OGS & MOS

carbohydrate if their diets fish glycerides ad adequate concenations of glucogenic amino acids. A need for dietary carbohydrat has been demon sated in pregnant Beagle bitches given a carbohydrae-free diet containing 26% o energy rom ptein and 74% rom at (211 Bitches ed his die becme hypogycemic at wheping and had depressed concenraions o acae and anine (gucose precursors) in pasma Puppies rom ypogycemic bitch es had a ow surviva rate, which may have been at eas in pr because o reduced motherng ability of the dams Wheher the queen requires a source o dietary gucose under comparabe conditions has not been deermined g r o . s w . e y i v l n e r o l a e u s n u n l a . a n w o r w s w e p m r o o F r f . d 6 e 1 d / a 6 o 1 / l n 5 0 w n o o D . S 2 E 6 P 5 - A 1 C 2 y 5 : b 4 . d 4 e 8 d 9 i 1 v . o r r t p u s N s . e v c e c A R . u n n A

Lipids Fat in he diet has hree major nctions: It provides a concenraed source of energy, it supplies essential fatty acids, and it seres as a caier or fatsolube vitamins. Fat may lso have oher roes reaed to he accepabiiy o ood nd to the provision of sensory (exural and orgnoeptic) cues for cats Dry comercial diets intended or cas usuay contain 91% crude at on a drymatter basis (129) However, exprimenta diets ranging rom ess than 1 to 64% at in he dry mater have been ed o cats Simiary a wide vriey o nm n egeae as n os ae een ncooae n exprmena es (118) ats have the abiiy o diges and utiize high eves of dietary a Mos o he measurements repord on appren digesibiity o a by cats have been derived om measurement of crude at Crude at is he materia exaced with anhydrous diehy ether; it is represented in cat oods primariy by gycerides but can also incude oher lipidsoube substances of ittle or no nuitiona value, such as pant pigments, sterols, and waxes In expanded or baked cat oods, not  o he gyceides may b eacted wih diehy ether (34, 113 unless ether exacion is preceeded by acid hydrolysis (16). Theefore, ether exracon may no provide adeuae measuremen o a o he rgycerde in these dies Apprent digesibiiy coeciens o crde a in comercia ca oods mkeed in he US have been repored o rnge rom 0.8 o 094 (179 These vues  simiar o hose repored rom he United Kingdom or apparent digesiiiies o acid eher exac in commeci ca oods, which were 0.79 in dry fds 088 in canned oods and 092 in semimoist oods (137). Apparent digesibiiy coeciens o crude a in expermen diets hat did not invove exuding or baing generaly have been higher than those reported for com mercial dies Mean values o 099 or beef and mutton fat (167) and 098 for butter, lard, unbeached and bleached alows, chicen fat, and yelow grease have en repoed (129) In one study (129), no signicant diferences we ound wn he appren digestibiiy cciens o individu ay acids ranging from C120 to C183 or adut cats.

 UO

5

PROIN ND MINO IDS


Oe of he os siing aspecs of proei and aio acid nuriio is ha he sae aio acids ae esseia for viruay a aia species acoss he whoe scae of he phyogeeic ree Our ehasis, howeve, is o wha aes he cat diffee or ique in is oei ad aio acid nuttio. he eentty of the no cid h been deerned n the rowin ie. As epeced, he esseia aio acids incude argine, hisdine, ysie, eucie, isoeucine, vaie, ehioie, hreonine, ypophan, ad vne (206. etry rne  ered for  rowth in the ery pos-weanig kitten (E. M. Kaw, J  Mor, Q. R. Roger, npb ished esus) u is o necessay afer the is few week. Taurine   dietary essentia fo he ca in ode o peen entr eina degeneaion CR, bt ecause his aio acid is o a aio caroyic acid, is o foud in otein, and   eaboie of cyseine, i is dicussed n the secio on hu ino cds beow. ro  quttive pon of vew he, nohn appeas uiqe aou he esseiaiy of hese aino acids Fo a quaniaive viewpoi, however, sevea ineesing differeces have bee foud.

Potei Requiemet  anas potein eqiee nay efects both i ina nitrogen equiee and i essent ano cid reireent. If the aino acd equees are ow he reseacher can foruate he de so ha eve he owes eve of poe esed ees a the essent ano acd eqieens; de hese codiios he iia proein eqiree becoes a easue of he ioge equiree of he aia. iewise, o deeie he esseia aino acid equiree, the niogen requireent ust be et in a diets so tht the oy tin fctor s he no cid bein ened. he eaies epeies in deerinig he proei requirees of ies ad adu cas used foodsuffs (8   o puified copones (5 1 155 ad wee done wihou ay owedge of he aino acid eqiees. Unde hose condtion he oten rereent w shown o be abot 0% of the die for the gowng kiten and bot 20% of the det for he dt cat. nderson et  (9 ed aio cid die in which the aino cid eqireents were e and cocuded hat 16% of e detry caories s poten eet he ioge equiee. When he daa fo he ae sudy ae poed, here ae no brea-poins on he ces, so it  o obvo how he author aived at their interreion. e enrn tht  no cid reqreent wee et re et  37 eted the dt cat poein reieent for antennce to e ao 12% of he die;  aey Q R. Rogers ad J  Mois (upbished esus see aso Ref. 208 have estiaed he roen eqeen

34


MacDOAD, OGS & MOS

of growng kiens o be abou 8-20% of he de whehe he ncreens of nogen in he die wee povided by free ano acids o by casein A pued des used eceny fo deenng poen and ano acid requireens of he ca have had abou 47 kcag of die. Usin he above resus and he sandard assupion ha poein and ano acd equieens vay drecy wih he enegy conen of he de we nd ha adu cas woud require abou 0% of dieary enery as poen and grown kens woud equire abou % of dieary enegy as proein Because cas accep hih-fa dies bee han ow-fa dies (especiay wh pufied dy des) he eque ens for proein and aino acds have no ye been deened wih ow-fa dies  copasons ae ade of h nogen equieens ised as he poen equeens in abe  of severa doesic aaan speces i s apparen ha hey fa no wo goups: civoes and onvoes erbivoes woud be par of he roup of onvores wih respec o poen equieens   Nuriionay he dog s a eber of he goup of onvoes even hough  is a eber of he ode Civora he ca and oher ue cvoes have subsaniay highe poein equreens han do onivoes. his higher requreen is pariy an nceased equieen for ainenance ahe han or rowh Fo exape, he ra equres hee es as uch proen o growh as fo anenance bu he ca equres ony 0% oe. f one exaines he increased increen fo owh as an absoue ncrease in proen need above ainenance he aoun is quie siar for cas and as. he eaboc eason fo he high poein equreen of he ca is he hh acvy of he nioen caaboc enyes n ca ve 209 hese enyes ae nonadapive so he obgaory nirogen oss s hh even when cas ae fed ow-pron des 207.

Tb 

Po qum

 of d) for au ad ow ama of va mamaa

c Spc

Rf

You

Adu

2 2

4 4

4 90 4 �3 78 76 6 92, 9 55 77 937; K. Sma, . R. Ro G o uud u

Do Ra Ma Ca Ca

8

5

29 7

9 0

 o

3 24



7

6

75



"Ideal potein, .e. meeing all the kown esntia amno acd requirements Ths may no  e mnk n x bVes expee a enge  e eneg.

&

b true for the dat

L UO

55

Am Ad qms A summary  the amn acid requrements  the wing tten is presented in Table 2 The arginine requment is listed as 1.05%  the diet

G


the dietar level  arginine required t prevent an increase in rtic acidura which ccurs when an argininedecient diet is ed 56) Arginine decienc causes e mst dramati respnse t an nuient deciency in the cat The eeding  a single meal  an argininedevid diet may result in hperammnema in less than ne hur and causes such severe hyperammnemia within tw t ve hurs that the cats exhibit severe symp tms  ammnia intxicatin which nclude letharg emesis vcalizatin hpersalivatin hperactivit hresthesia ataxia emprsthtns ex tended limbs and expsed claws The mst severel aected cats shw

T 

unt etmate  amn acd quement  wing kiten· eerence

euirement

Amin acid

05b 030

Agnne tdne

756 10;  Quam,  G rri, Q  ger, unpublied eult 11 .  argrove Q.  ger,  G i, ubmed r publican 11 D  Hae Q  ges, G , ubmted 7;  A O'nnell, Q  ger, J. G i, unpubled eult

Ileuce

050

une

120

ne

080

etnne w exce s etnine ctne Penlalanne wt exce tine) Penlalanne ine

040

237245

075 040

217245 0  Wlliam Q  ger  G r-

00

ri, unpubled reult 0 J William Q  ge, J G  i, unpubled eult

enne ptpa Vline

070 011 0&

112247 1099 18

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+





aBed on exrintl diet contining ut 4.7 kcg diet. No work h en done to detemine wete it would vld  expe mno cd eurement of ktens s  entge of  det bed on  tnddz  cJg de Ul o a c qm  b o  mml qy of c mo cd reid in e det to gve mxmm wegt gn nd mxmum nogen non. ed on mnmzng orotc cd excreton n t urne Mxmum wegt gn  ustne by rgnne n e det "e on te dety level of dne eue fo te peventon of ctcs er evel mon Hdne t of  dt  ct  mxm og o  mml qm cold  o of  d; m w d d vl ve not n teted.



40

09%

0%

04 0%

0 0%

53


MacDOAD OGS & MOS

hypothema and badypnea cyanoss whh ae olowd by death n a low percentage o cats 12, 13, 1, 20 Ths rases th queston o why th cat s so much moe senst to agnne dcency than are othr mammas, such as rat or man. Anmas a mor snstv to ammona oad whn thy a n th postabsopte state 254), and th admnsaton o agnne geaty deceass the ammona ntoxcaton undr ts condtons  Th gnray acctd xplanaton o the peenton o hypammonma by agnne s that t acts anarotcally to stmulate urea synthss by rovdng an ntemedat o the urea cyce, as the ntrmdats ar a dptd durng the ostabsorptv state. The eason o th hgh senstty o the cat to agnne decncy s relatd to the metabolsm o the uea cycle ntemedates.  summary o th deencs btwn th cat and the rat s shown n Fgure 2 Ther apea to be at east two rasons or th dernces betwn the esponses o the cat and those o th rat: () Th cat aparenty s less abe to synthsze otne and b the cat may be ess ab to rod ctrne to th kdney o coneson to agnn. Seera lns o dence pont to mpaed othne synthess n the cat. Detay othne cometey pvnts he hyp onma o argnn dcncy n cats athough th anmas stl lose aromaty t sam amount o wght as y woud  any othr ssnta amno acd wre omted rom the det 3, , 20). Rats n conast wl gow wthout any detary argnn athough not mamay; yt the addton o othne to an agnnee det dos not mproe wght gan n ats 15) or n cats Stewat t a 242) masurd othn cune and argnn n the lers o cats ed an argnnere det. They conrmed hat the cats

GLN + CIT L U t +* SA-OR + P5CPRO

 

    

    

V

    

KY

Figur  Metaboim of ureacce inteedite in the rat and cat Enme that have been hown or  reumed to be min in the cat ae mred wth an ate GN= utamne G= gutamic acid GS= gutamc emadehde 5C= poine5carboxate O= proine ON= oithine CI= citne G= arginine

(*)

L NO


53 7

showng seee hyemonem wee the ones tht wee most seeely deleted o le one The concentons o othne n le wee lowe n cts thn n ts ed gnneee mels. Howee, the uthos ound no deence between the ct nd the t n tems o the ctty o knetc contnt o ontne mnotnee OAT o n tem o N cetylglutmte concenttons o le mtochond The de noo ynthess o othne n ts s to occu mly n the ntestne By mesung teo-enous (A-V) deences cos the used t ntestne, Wndmuelle  Seth (25, 260, 261) showed tht glutmne nd glumc cd e coneted to othne, cullne, nd olne Hezeld  R (10) showed tht OAT solted m t ntestne could be mde to ct n e othneomng decton OAT om t le could not uthemoe, e ntestne hs the enzyme glutmc cd eductse, whch oms glutmc cd semldehyde tht cn be coneted to othne nd olne ( l08, 2 1 2). ee uo dd not nd gutmte educte n ny ote t tue ex mned, ncludng the le (l08, 2 1 2) In contst, the ct my lck glutmc cd eductse n bo e ntestne nd the le Costello et l (55) he not been ble to detect t enzyme n ct ntetne o le, thu eyng tht te ct cnnot mke othne om glutmte (see Re. 164). Cllne c comletely elce gnne n the dt o the ct ( 166) s n the t but  hghe dety concenton o ullne s needed o n equlent owth te (C ohnnsen, Q. R. Roges, J  Mos, unublshed esults) In the t, ctullne syness occus mly n the ntestne (26 1); ctllne elesed om the ntestne eces the kdney whee t s coneted to gnne o use by othe tssues (76, 203) . Snce cllne s mde om othne,  lck o glutmc cd eductse n ct ntestne would eclude cllne synthess om glutmte. The ct my he ddtonl metbolc deects, such s  lck o othne nscbmylse n the ntestne (see gue 2) These metbolc deects would lo exln why te leel o cllne n ct lm s nomlly qute low (207). Polne s dsensble o the ct (206), s t s o the t In nethe the t (5) no the ct  A. Bungton, Q. R. Roges,  .  Mos, unublshed esults) does olne melote hyemmonem. Although t mght be ex cted tt olne would be coneted to otne n te le o kdney, e knetc chctestcs o OAT n thee tssues nd the substntl ctty o yolne-5-cboxylte dehydogense would eent the synthess o othne nd o the coneson o yolne5cboxylte to glutmte  olne nd glutmte wee esent t hysologcl leels. The deence · between he ct nd the t  t olne  t let one hl o he dety gnne equement o te t (204, 205); t does not se the gnne equement o the ct It seems lkely tht n e ct, OAT s  consttute enzyme lke the uecycle enzymes (20), so othne degdton would be

538


MacDOAD OGS & MOS

obligaory in e ca an us wou provi a precursor or proine synesis wether proine were in te ie or no n conras OAT is aapive in e ra an oiine caaboism is reuce wen ras are e a ow-proein ie compare o wen ey are e a ig proen e 220 Wa is e pysiological meaning of e ierences in arginine meabolism in te ca? From a eeoogica sanpoin i is aracive o posulae a as a stic civore he ca as never a a ie limiing in proein an ereore in aginine or oiine bu as aways a an ecess o aino acis muc o wic is neee or gluconeogenesis ecause anima issues are low in car boydrae us e meaoic need is no o mke oiine wen nirogen inake is ig because arginine rom ieary proein wou provie e oiine neee anaperoicaly in e urea yce as wel as proviing arginine or proein synesis raer e nee is o epee oiine urng e pos absorpive sae so a urea synesis can e markely reuce o preven ecessive caaoism of amino acids o uea is penomenon also occurs in ras bu o a lesser exen. Because e ra is an omnivore ere are imes wen i ingess a lowproein ie an aaps by conserving nirogen in pat via meabolic aapaions (220); ese aapaions i.e canges in enzyme acivi ies as a resu of dieary composiional canges apper o e muc more limied in e ca 20 Work done wi e mink  and e fee (66 inicaes a e a's arginine meaboism is no unique bu is represenaive o a o stric civores. HISTDNE ,. LEUCINE, ISOLEUCINE, VALINE, LYSNE, PHENYLALANE



&

Noing unique as een repoed aou isidine deficincy in e ca e usual epeion o emogobin as been oun an aer ongerm isiine eiciency caaracs ave een oun D. Qua J G Morris Q R. Rogers unpubise resuls. soleucine an vaine requiremens are simiar in e ca an ra bu e eucine requiremen is somewa iger or e ca D. . Hargrove Q R. Rogers 1 G orris sumied for puicaion. ese workers (unpublise resus recenly oun a ile or no rance-cain amino aci anagonism occurs in e kien Ta s 0% of any one of e ree brance cain amino acis i no ecrease e grow rae o kiens e a ow-proein ie a was soewa limiing in eac of e brance-cain amino acids. Te ca an e ra ave simiar ysine requiremens u as a percenage o e proein e requiremen or ysine is consieraby ess or e ca an e a because e ca as a iger proein requiremen ysine is limiing ony wen a cereal proein suc as wea guen is e major iery proein (122 207) Te oa penyaanine-plus-yrosine requiremen o e kien is abou 0.8 % o e ie Approximaely 60% o a requiremen can be me wi yrosine (Tae 2 wic as been sown o be a ispensale amino aci (206

FL O



e ypop equeme of e   quey m o oe o eve oe pee . Howeve ee  eggbe fow of ypop oug e quo  pwy  e  eefoe oe o yee  gf quy of  (ee eo o WeSoube Vm ee e eve uque pe of upu mo  uo  mebom  e .  e e equeme  oeby ge fo e   fo mo oe mmm (be 2 Seo e  exee  uque be upu mo  (fee  e e.  e  eque  eay uppy of ue  mo upo  o peve e e egeeo (CRD e eo fo e  g equeme fo upu mo   o ey ppe bu   bee uggee    ee o e ' k    ee o yee fee  g upu mo  equeme eu  ee mo  beg e mo mg mo   mo  e. Abou 9% m poe  eee  e e o mee e upu mo  equeme of e gowg ke  equeme wou ju bey be me   me e og eoug poe o mee e oge equeme (208. ee  o  oe quy of meoe ovee o yee Cye pe bou oe f of e meoe equeme  e    oe pee (237 24. eefoe p of e ge upu mo  equeme ppe o be fo ye  p of  fo mee pe e Cyee  ue fo fee ye  fo e ye of    my ou fo p of e ee equeme Se e  omy e  gf e pep e ee ee fo popop fo bopo  po of f  eue   ee  e ee fo Seoy meoe fo meyo eo A g ouve eve of meoe me wou o eu   ee ee fo ey meoe. A exmo of e ve of e eyme  e vou pwy ou ep fy e eo fo  omy UHUR AMNO AC


Fnn e uue of fee  ow beow CH

I

N +



HO-CH-CHC-S-CH-CH-COO





CH  be upu mo  w f epoe by D & H     6  w oe  eze by We e     Beue fee  o vbe ommey vey e wok  bee oe o e effe of e o e boye  exeo of fee Rbe

540


MacDONALD OGS & MOS

20  howvr xamnd svra aspcs offnn mabolsm and rpord that fnn vs wr hghst n ad ma ca rn q ow n fmal ca urn and ndcab n ktns of hr sx H fond an approxmay fourod vaaton n fnn xcron among a grop of  ma cas H also showd ha gvng an adul fma ca tsosron for on wk rsld n an ncras n fnn xcron o norma adu ma vls and hat h admns aton of stradol to an adt ma ca for wo wks had no ffct on flnn xcrton. Robrs (0 and Avzons  Wrston 17 boh sd sotopc ncorporaton mthods n vvo and fond ha cn was a br prcursor for flnn bosynhss han was mvaonc acd Usng smlar tchnqus Robrts fald to g any  S ncorporatd n flnn from thr [ S]mthonn or  S]cysn. Th st of fnn synhss s ncran bcas Robs 20 found som fnn n both lvr and kdny and a smar amount n boh skn and bood. Wang 25  also amptd o dtmn th sourc and prcrsors of flnn. H too was argly unsuccssful n showng flnn bosynhss n vro bt h dd fnd  C]aca ncor poraton no fln n vvo. Th mtabolc ro of flnn s a mar for spculaon. t may b a rnary componnt for trrtoral markng or prhaps s somhow nvovd n th rguaon of sro mabosm 230.  s nrsng o no th smary of flnn (Equaon   abov and sovahn COO



NH +



CHCHCSCHCHCOO



2.

CH sovalthn has bn found n ca urn 83 and n h rn of hyprchostr omc pants 8 Lt work has bn don th past 20 yars n ths nrstng ara of flnn synhss and funcon. Th oa qanty of dtay sulphr amno acds rqrd to provd for th synthss of flnn tha s xcrd n h rn of a ma cat shold no xcd 0.05% of th dt 575 mg. hrfor, rgadss of h nrsng aspcts that ar stl o b ld about fnn synthss and funcon t s doubfu tha t alon s th rason for h hgh sphr amno acd rqrmn of th ca spcally th kttn. aurn dfcncy n th cat has bn shown to cas cnral rtnal dgnraton ( 0 S an s a amno sfonc acd amno than sulfonc acd and as such s no prs n pron  s normaly mad from cystn n th lvr and som othr ss . g .  bran of mammals. Exct Turne

FL O


4

r it nugatin t bil ai (100 0 an it prn in a w ppti u a t rn glutarn gam-glutayl-taurin (77, 84, taurn i ntially tablialy inrt, i.. taurin rm itr ngnu ynt i r r itary intak i nt urtr tabli an i vntualy xrt a u in t urin CRD a bn rpr in at a nubr  tim (28 2, 16, 21 , 2 16, 227 br Hay an wrkr (04 0 ivr t nutrtinal au  t ia. T ivry  taurin iiny a t au  CRD l Hay an wrkr (106, 243 t xamin it yn ti an tabim in t at T t  in CRD an t un tin  taurin in gnra a bn rviw vra tim (20, 03, 106 23, but tr ar n pubi xprimnt tat larly in t taurn rquirmnt. vral abrati av rpru CRD unr abraty nitin (74,  it i lar tat t at n  taurn in t it, but t ti rquir t pru CRD var r 23 t 2 wk r lngr, an  inin  CRD in at  taurinr it vi r 2 t %

(8, 21 10. P  t unainty abut t rquirnt i au by unknwn itay r nvirnmntal atr tat at t rquirmnt r xamp CRD a bn un in kittn  mrial at  tat ntain btwn 200 an 300 mg  taurnkg ry wigt  it (7 but  labat av bn unabl t pru lin in wanling kittn atr ing pui it ntaining n tauin r n yar Q R Rgr an J G. M unpubli rut xamin t iprtan  typ  itry at, v  itary prtin, prn  ity br, an v  ity upur amin ai n t inin an vrty  CRD in at Atr xamining al t atr ty av nu tat ny t lvl  ulpur ain ai at t utm  CRD n wanling kitn wr  a it wt n taun an wit a lv  ulpur amin ai nar t rquirmnt (8 Q. R Rgr, an J. G. Mrri, unpubi rut, all t kittn vlp CRD witin 6 mnt; wn twi t upur ain ai rquirnt wa , n CRD wa un in 12 mnt Trr it appar tat t ran tat kittn  mrial at  ntaining 23 mg  taun/kg it av vp CRD i tat t tta avaiabl ulpur amin ai wr nar t rquirmnt altug vn wn kittn ar  a ig v  ulpur ain ai wi n itary tauin, a lw prntag  kittn vp CRD Tu, t rmmnatin  t Natinal Rar Cunil itt (174 tat t taurin rquirnt wn t ulpur amin ai rquirmnt i jut mt i btwn 20 mg an 1000 mgkg  ry t ti appar vali An intrplatin  xtng ata igt put t minimal rquirmnt at apprximaty  /k it A mr aurat timat  t rquirmnt ay nt b availabl r m tim bau tui  t wr v prbaby l b xtn r tr t iv yar br ty an b nir a.

4

MacDONALD OGS & MOS

Alhough Anson  a  1 1  sc h honn qumn a 0.8% o h  a clos xamnaon o h sus ncas ha kns gw jus as wl on 0.7% hronn. Fuh Tchna  a 47 an Rogs & Mors 206 oun maxma wgh gan wh honn a 0.7% o h  an somwha ss han maxma gowh a a ay v o 0.6% honn so h mnmal honn qumn appas o b abou 0.7% o h . Ktns  naqua hronn vop nuologca pobms as aly as h h ay vn hough  kns w gowng 47. Sgns nclu trmo aaxa ncoonaon ysqubum an cv ghng x. Nuolo gcal sgns w pogssv bu compy vsb All sgns nca cba ysuncon n aon o h nuologca sgns sva kns vop hoacc an plvc mb amnss assoca wh cap vaon an s movmn o h hn lmbs. Ths kns sm ucan o wak o n b no vsb pan o scomor co b nuc by vgoos palpaon o h ac lmbs an jons Th pobms w compy vs by ng an xcss o hronn. HRONN


Amo Ac Itolece Lttl has bn on o xamn h possb cs o hgh ary vs o nvual amno acs on cas.  s known ha cas a ss olan o an xcss o guamc ac han a as chckns an mos oh spcs () Alhough h ktn appas o b ab o ola 6% guamc ac n h  % o 1 %  gluamc ac suls n ncass n  gluama n pasma o h o n ms nomal lvs Ths hgh lvls o gluamc ac su n  spoac mss an vnualy caus cassca hamn cncy n ktns  44 mg o thamnkg  a lv whch s nomaly aqua 177 Th nau o h nacon bwn hs wo rnshas o bn compy cr  bu  appas ha hgh ary gluamc ac somwha ncass h cal xcon o ay hamn 6 n laon o ths  s woh nong ha mos anma prons conan ony abou ha h conn o guamc ac as ha n many pan prons. Fau   7 cny xamn h c o xcss ary mhonn n wanlng kttns. Thy oun a gowh psson n kns  2% aonal mhonn Wh 3% mhonn som cas compy jc h  Ths s n contras o cn xpmns n whch ng 10% o h  as lucn solucn o van ha no avs c (D M. Hagov Q. R. Rogs J.  Morrs unpblsh sus. Thus h ca apprs o b ss oan han som oh mammas o xcsss o som amno acs such as guamc ac an mhonn b mo oan o xcsss o oh amno acs sch as h banchchan amno acs

FELNE NTITON

43

MINERALS

g r o . s w . e y i v l n e r o l a e u s n u n l a . a n w o r w s w e p m r o o r F f . d 6 e 1 d / a 6 o 1 / l n 5 0 w n o o D . S 2 E 6 P 5 - A 1 C 2 y 5 : b 4 . d 4 e 8 d 9 i 1 v . o r r t p u s N s . e v c e c A R . u n n A

There is a paucity of data on te cats qualitative and minimal quantitative requirements for miels. Successful growth ad reproductio have bee achieved with puried diets cotaiig a mieal mixtue simil to that requied by gowig rats and pigs e.g. 0 2. The cat's quatitative requiements for essetial mieals theefore do ot geatly exceed those of these species, and the cat does not hae a requiremet for ay mieral that is ot essetil for these species However these fidigs do ot preclude the possibilities of peculiaities in the taspot, metabolism, o stoage of mieals by cats ad of a lowe quantitative requirement fo a paticula mineral. When cats consume the whole bodies of thei pey, they obtain all necessay diety mieals , but whe the so tissues re cosumed without the skeleto he diet is decient in calcium. A epo of seletal abnomalities i lage felids in he London Zoo s de by  sugeon of Mddlese ospitl as ealy as 888 (3. He epoed that "when cubs which ealy manifest signs of icets were promptly fed o boe dust ad cod liver oil they made good recoveies Pesumably hese lions wee given an all-meat diet without bones. Ths observation is of pticul interest as it pedates the demostatio by Sir dwad ellanby in  that icets could be preveted by cod liver oil. It took aother 70 yers befoe research ad cliical repots appeared describig the effects of allmeat diets on the induction of the disease now refeed to as uitioal secondy hypepaathyroidism NSHP) in both wild captive and domestic civoes 46 78  07  3 2 200 226. The metabolism of calcium ad phosphous i the cat does ot appe to diffe markedly fom that i othe mammals The fequency of occuence of NSHP in cats appers to be the result of the combination of the low calcium cotet of aimal so tissues which are highly palatable to the cat, ad th e fact that owners frequetly feed aimal tissues as the sole diet The quantitative equiemets of the cat fo magnesium have not been deined, but excessive dietay intaes of magnesium have been shown to predispose male cats to the fomatio of ury calculi composed of stvite magesium amoium phosphate) which cause subsequet uethal obstrc tion. Thus it has been suggested that high intaes of magnesium ae a etiological factor i producing FUS 70 27. The potassium requiemet of gowig ittes has bee show to be influ enced by the potein level of the diet   l l ). The estimated potassium require ment of ittes consuming a pufed diet based on 33% soy potein was 0.3% and fo 8% soy potein, 05% otassium deficiency was evidenced by hypokalemia, oexia, etrded gowth, emaciation lethagy, locomotive disordes and uempt fur

5


MacDONALD OGS & MOIS

a udoubedly equie diey ouce of odium ad chloie, bu qua iaie equieme of hee eleme have o bee expeieally deed Sa odium chlode) ha bee ued o iceae wae coupio ad hece uiay oupu o aid i peveig FS  3, 36, . The quiaive diea equieme of he ca fo io hae o bee udied alhough Spay  iddowo 29) meaued he mou of io i he ca a vaiou age of deelopme. The euieme fo coppe appea o be appoximaely 5 mg/kg die 6, which i imila o ha of ohe mammal. oppe deciec iduced i po-ealig kie euled i a deceae i he coeaio of oppe ad CuZ-upeoide dimuae SD) i plama No chage wee obeved i M-SD, packed cell olume, o coa colo o codiio i hee coppe-decie kie. Hoee, whe he deficiecy wa iduced by giig pega uee a deficie die le ha  mg coppe/kg die), hiochemical chage i he aoa of eoaal kie wee obeved ha idicaed coecie iue defec. he lae chage occued wihou de peio i iue coppe eel. hi i i coa o obevaio of weaig ke give he coppe-deficie die chage i iue coppe level wee appae i his goup. I alo appea ha he hemaopoieic yem of he adul ca, i coa o hoe of ohe mammalia pecie, i eia o chage iduced by a low coppe au. Thee hae bee wo epo (4  30 of expemeally iduced zic de fciecy i he ca. The cliical ig of he deficiecy ae imila o hoe decibed i oe mamal 24 hoee, hile e dea leio appea o be le eee i ca ha i pecie like he pig, a mild deciecy of zic, which caued o appae gowh edaio ad allowed epoducio i a female, caued defecie pemaogeei i male ca 0). Thee i o ifomaio o he availabiliy o ca of zic i aual poduc o i comme cia ca food, bu ome udie ugge ha i may have lo aailabiliy 4  . R. Roge,  G. o, upublihed eul). Iodie defciecy ha bee epoed i zoo felie (78 87 ad i domeic ca 4). liica ig clude hypeophy of he hyoid, alopecia, abomal cacium meabolim, d feal eopio. A ieacio ha bee epoed bewee calcium meabolim ad iodie 200, 222, 226). Iodie upplemea io duced cliica ig of he deficiecy i ca gie almea o ow calcium die. I ha bee aed ha euhyoid ca ca oleae elaiely high doe of iodie 223), bu ove a peiod of 42 moh, oe cliic ha foud hyehyoidim i  3  ca beee 6 and 20 yea of age 184. liical ig of hypehyoidim icluded polyphagia, iceaed aciiy, polydipia, polyuia vomiig, ad iceaed bae ciculaig level of hyoxie ad iiodohyoie  i o kow whehe he coceaio of iodie i com mecial ca food i a eioogical faco  poducig hi dieae.

L TTO

545

hee ae o pubished sudes o he equemes of he a fo magaese seeum homium ke silio vaadum molybdeum fluoe o . hese eemes ave bee show o be esseial  ohe spees (78. VIMINS

Fat-Sluble Vtamns he mos uusual aspe of fasoluble viam uo  as is ha pefomed viami A is a dey eseal he ovesio of oees o eio whh ous i he esal muosa  ohe speies ludg he dog ( 9   80 s eggble  e a (3 86 As a esul ee dea o iaveous admiiseed �-aoee a eve he develome of vami A deiey (86) Beause pefomed viami A is o pese  pla pids he a equies a soue of amal fa  paua oga meas) o saisfy is viami A equieme. oombes has epoed ha he fox aohe voe ma aso be uable o ule aoeoids as a soue of viam A (52. hee is some fomaio egadg ssue levels of vami A i as . he ideys of as oai appoimaely e imes as muh viami A as do he kideys of ohe aimas ( 48  58 howeve lve soes of viam A ae moe losely elaed o dieay saus ( 58 224  e hgh levels of viam A i a kide ae o foud  oe Fedae (   0 he quaiaive equieme of he a fo viami A as o bee peisely deed So & So (228) oluded ha appomaely 2000 /day wee eessy fo oma epoduo. Howeve ohe wokes J G. Mois Q R. Roges upublished esus) foud ha 6000 /kg de me he equeme fo oma epoduo i femae as. Vami A oy ous fequel i as ed dies ossig agely of aw ve (22 ad a be dued i kies gve suppemes of 00,000  daly fo fou weeks  ITAIN A


Viami E defey  as was fs obseved b od & Slige (54  kes fed a aed fsh-based de. Defie as develop seais  whih he depo fa beomes fm ad yellow-oage  olo. hs paifu deposiio of eoid pigme is ievesibe ad leads o deah if o eaed ody (53 showed ha he deiey ould be peveed by 20 o 40 mgda o oopheol Almos al aualy oug ases of viami  defe have bee  as fed des hgh i fsh espealy ed ua) whh oais lage amous of highl usauaed fa ads (47 7 oda al a foods omposed of ed ua mea ae fofed wh vam  AN E

Alhough vam D s esseal fo peveg kes i kes (8 he equieme s pobaby que low he kies wee fed vam ITAIN 

54


MacDONAD. OGS & MOIS

Decent dets tt contned 1 % clcu nd % pospoous only one ct out of seven ws stll lve e 1 onts (89. en te det contned % clcu nd .5% pospoous, sx out of nne cts suvved. ckets de veloped n vtn Ddecent cts fte 5 onts, but n tose cts tt suvved e st 1 onts ofte study tee wee spontneous essons. In these studes coleclcfeol (5  twce  week) pevented ckets Scott (4 epoted tt clncl cses of ckets occu ely nd only n kttens kept n the d ves et l ( 19 epoted tt levels of vtn D n pls of vtn Defcent kttens selded fo ultvolet lgt declned ntlly but ten tended to ncese wt ge. ey epoed no sgns of bone dsese fte 9 onhs. xcess vtn D (.5 g/kg det) cused clcfcton of soft tssues ( 197 wc ndctes tt lge doses of tn D e toxc fo cts s tey e fo ote ls e essentlty of vtn K fo cts s not been estblsed. ebe & o (188 obseved tt n dted beef det, wc cused eoge n wenlng ts dd not ve dvese effects on cts. Te lvel of vtan K ws cculed o be 6 gg of sods ITAIN K

WtrSb Viti In omnvoes, te equeent fo ncn cn be et by detay typtopn Te ptwy o yptopn etbos n ve s  bnc pont t -no- cboxyuconc-seldeyde nd ts nteedte cn be conveted ete to qunolnc cd wc s then etbolzed to ncn o to pcolnc cd In ost ls te ctvty of pcolnte cboxylse s qute g so ost typtopn s dveted wy fo te ncn-syntesng ptwy e ctvty of pcolnte cboxylse n cts s 35 tes ge tn n ts, so h  lost no nh of nn  44.  o kn    lckng ncn develop sgns of defceny tt cnnot be evesed by te dnstton of ddonl typton (1 DSlv et l ( found tt wen yptopn ws dnsteed olly o subcutneously n doses of up to  g/dy tee ws no sgnfcnt ncese n uny N-etylncotnde exceton. Lekle et l (143 gve ge doses of typtopn (1 g/kg body wegt/dy) o kynuenne ( g/kg body wegt/dy nd found  slgt ncese n N-etylncotnde exceton but te ncese ws only ppox tely 3% of tt seen n ts. ese utos lso sowed tt less tn  % of  dose of -[C]typtopn could be ecoveed s ncn etboltes n ct une (14 ee ppe to b ote pecultes n typtopn etbols n cts n tt 8% of te doctvty n une of cts gven lbelled typtopn could not be dented s ny of te known typtopn etboltes. e efcency of coneson of typtopn to ncn vaes wdely ong speces: t s getest n ts l so n un, nd neglgble n  Pn

L UO


547

& Combs (186) epoted tat picoinate caboxyase actiity is een ige in out and samon tan in cats Despite te act tat cats do not conet yptopan to niacin, ei equiement o niacin does not apea to exceed tat o oe species; howee, te cat depends on anima tissues as a souce o niacin, because nicotindes ae pesent in owe quantities in pant tissues e cat equies tiamin (73), iboain (141), pantotenic acid (88), pyidoxine (87 ), biotin (39), oacin (60), and itamin B  ( 135) Ascobic acid is syntesized in adequate amounts and is not a dietay essentia (59) Vitamin B is ow in diets o egetabe oigin, and an aegetabe diet woud necessitate suppementation wit tis itamin (135) Deciencies o some itamins occu wi eatie equency: tiamin deiciency is easiy poduced by eeding s poducts (especiay cap and eing) in wic te tiaminase as not been destoyed (238) Choine i s equied by kittens wen e ee o dietay methionine is ust enoug to meet te metionine equiement (218) ee is eidence tat te choine equiement o the kitten, as o othe species, can be met in pat by excess diety metionine (12), but it is not known wethe dietay coine is competey dispensabe ee ae epots that inosito is equied by cats (224), but is as not been inestigated oougy OHER FOOD CONSENS

Benzoic acid, a ood peseatie, is exemey toxic to cats Bedod & Cake (26) sowed tat doses o benzoic acid in excess o 0 45 g/kg body weight ed to yeestesia, muscua emos, depession, and oten, deat In many se cies, benzoic acid is detoxied with gycine to om ippuic acid and with gucuonic acid (258) Cats ae abe to om gycine conjugates but tey ae ow ees o gucuony ansease in ie (71) is may aso expain te toxicity and sow metaboism o many dugs by cats (33, 253) Cats excete unusua conjugates in wic quinoine2caboxyic acid, a typica aomatic acid, is combined with gycytauine o gycygycine ( 1 25, 26) e sign cance o tese eactions is not known ESSENAL. AY ACDS

Mtim Te essentia atty acid (EFA) equiements o most mmas can  met entiey by inoeate In the ie, inoeate is coneed to aacidonate by ateating desatuation and eongation (Figue 3) he st indication tat tis sceme migt be dieent in cats came in 975, wen Ries, Sincai & Cawod (199) epoted tat cats ed puied diets containing inoeate but acking aacidonate ad extemey ow ees o LA F �6 DESAAN



DND, RGR & RRI

  S

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� 1 8 : 2 - 1 8  3  20  3  20    2 2    2 2  5 1

 A

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ARAH A

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02,11,1420 Patways of linoleate metabolism in mammals. In the liver of the cat there is almost no Figue A6-desatuase att, s aahdnate s a deta essental E= elngatn, A= desatuatn

3


aacidonae in lama liid. Te auor uggeed a ca were unable o ynee aaconae om noeae ey ao preced a e ca wou require aracidonae in e die Haam e al 0 owed a e enzyme defec wa a e L6deaurae, e  e n e converon n ca given njecon of[- C]lnoleae, ver le raoacvy wa ncoorae no acdonae Sncla e al 2 conrmed ee reul and owed a le an 0% of e radioaciviy from Clnoleae recovered om iver id wa aociaed wi fay acd avng fou oube bon Mo of e nfomaon abou EFA meabolim in ca a been obained by udying e eec of dieren die on e ay acid comoion of blood and iue liid Wen ca e fed uified die conaining alowe eed oil a e only ource offa e eel of acdonae n boo d ecreae w ime 80. A e ame ime linoleae comrie nearly 0% of e fay acid in lama liid 0, , 2. In ca fe a e low n noleae, e eel of lnoleae n paa nd erroce decreae wi ime and evel of n7 and n fay acid 6: n7, 8:n, 20:n, and 20:n increae 0. Tee fay acd are yneized de novo in liver and an ncreae in ei concenaion i a ign of FA decency   4 Alou e amoun of 0n yneed n linoleae decien ca i ve mall comared o ra, for eamle e ynei of 20n from 8n uggeed a e ca mig  abe  yneie ome racidonae from linoleae becaue e ame enzyme are reonble for bo converion. erefore, Sinclar e  22 uggeed a eary linoleae could rovide aracdonae n ca, werea River & Frankel 6 concluded a e moun of acdonae a coud be yneied w ul be ngni ca f linoleae were convered o aracidonae, e incluion o a ource of noleae n an oewe ecen de would reu i ncreaed level of aracidonae in ca iue. owever eaminaion of e fay acid comoi ion of ama, eryrocye and liver liid of ca fed die wi or wiou afower eed oil owed a ere wa no ne increae in aacidonae wen afower ee o wa ncue n e e 0. u,  appea a e

L NUON

549

actty o te 6-desaturase s so low as to be nearly saturated wt subste een n lnoleate decency so ere s nelble syness o acdonate n cat ler. Ts conerson oweer may cur n cat testes. eels o aracdo nate and loner-can n-6 atty acds  er n testes lds o cats ed a det contann lnoleate tan n tose ed a lnoleatedecent det  152 How eer te aracdona n testes s aarenly nt released nto te blood sream  Snclar and coworkers 231 sowed tat a A5desaturase s resent n cat ler Tey njected cats wt 20:3n6 labelled wt C n te 2oston and ound radoactty n ler aracdonate Ts nd cated tat 203n6 could be conerted to aracdonate. Te resence o a A5-desaturase results n te aerance o 5 1 1  1203 n e tssues o cats ed dets contnn lnoleate 1; see Fure  eels o 20:2n6 and 5  1 1 1420:3 n bld ncrease as te leel o lnoleate ncreases 149 150 ltou oter mammals can also syntesze 5 1 1  1420:3 248 ts atty acd usually s not detectable n anmal tssues. n uman erytrocyte lds te leel s only 0. 1 % o total atty acds . L acDonald unub lsed results n cat erytrocyte lds te leel s almost 3% 150 Te syntess o relately  amounts o 5  1 1420 by cats robably results rom te exemely low actty o te A6desaturase wc allows lnoleate to accumulate and to be sunted toward 5  1 1  12 03 Snclar 23 1  232 suested tat te cat mt be able to syntesze aracdonate by A8-desaturaton o 202n6 1 1  1202 to orm 20n6 8 1 1  1420:3 but snce tere s nelble syntess o aracdonate by any route bot te A8- and A6-desaturases aarently ae exemely low act tes n cat ler. Te resence o a unctonal elonase and A5-desaturase 231 means tat 1 8:3n6 sould be a usel recursor o aracdonate. rankel & Rers 81 used eenn rmrose ol wc contans 18n6 to sulement cats tat ad reously been ed a det contann 25% salower seed ol as te only source oat ter e days tere were ncreases n te leels o 18:3n6 and 20 3n6 n lasma osatdylcolne but leels o 204n6 were uncned. t tt tme te autors concluded tat cats lack a A5desaturase and were unable to coner 20:n6 to 204n6. Howeer later studes by Frankel 80 sowed tat e leel o aracdonate n lasma lds ncreased only ater eenn rmrose ol was ed or 10 weeks or more Snclar et al 23 1 sowed tat wen te metyl ester o 18:3n6 was ed to cats te leels o 203n6 and 204n6 n erytrocyte lds ncreased wc s consstent wt e A5-desaturaton o 20n6. 5-DESATURATION


0

DND, RR & RRI

F Defcien in ats

Rivers and coworkers 19 19 19 199) reorted signs of FA deicienc in cats fed a diet ig in linoleate but lacking in aracidonate e defciency signs included oor growth "staring fur skin lesions roun te mout an on te ocks slow woun ealing, fatt liver, increased suscetibility to infections oor reroductive erformance underde veloed testes an cleft alates and oter defects in kittens bo to efcient dams Later Frankel 80) sowed tat growt was not abnormal as long as resratory infectons were revente ese erl studies were done witout te beneft of more recent knowledge of te nutrtional requrements of cats. Te diets used by Rivers et al 199 lacked taurne nd -tocoerol. Stean & Haes 20 suggested tat te deciency signs could ave resulted from vitamin  decienc. e defcien cies of taurne nd vitamin  were corrected in later studies 19 but levels of oter nutrents ma ave been marginal. xamination of te diets tey used  19, sowed tat level of inc in te mineral mix was only 2% of te NRC requrement 177; te iets also lacked several oter trace elements. Snce many signs of inc ecienc re similr to tose descred aove e.g. oor woun ealing, eroral skin lesions, rerouctive failure, n cleft alate 10  Kane Q. R Rogers   Mois unublised results) it was difcult to determine wic signs were attributable to aracidonate defciency alone Altoug sulementation wit evening rmrose oil resulted in araci donate sntesis and caused emae cats to come into esrus (81 reroductive erformance was not imroved 80 wic suggests that tee may ave been a decienc of noter nutrent MacDonald et al  1  1 , 12 also found tat cats fed diets containing linoleate % safflower seed oil) but lacking aracidonate become aracidonate defcient. Platelet aggregation was imaired in aracidonate-deficient cats and tere was occasional tromboctoenia 11. Female cats ad few litters; kttens were bo dead ad undereveloed  12 but tere were no arent birt efects. Wen te same cats were sulemented orally wit etyl raci donate, eoductive efonce imoved geatly; kitten suvival inceased frm 0 to 7% n cats fed a uried diet wi animal fat added to rovide aacidonate at 0.0% of dietry kilalores, rerouction was essentially norml Wen oter diets containing as little as 001 % of kilocloes as racionate wre used, reroductive erformance was not lways comletely normal but was better tan it was wen aracidonate was absent Reroductive failure in females is terefore an exemel sensitive index of aacidonate deciency, and te aracidonate reuirement fr normal reroduction aears to be vey low 12) AAHDONA DFNY


FELINE NUTRITION

551

Because tee is negigibe syntesis of aacidon ate fm ineate in cat ie te cat is a usefu mode f studying te secifc physioogica oes of tese two faty acids Lees of inoeate can be ateed witut caging ees of aacidnate in tissues. Te esuts of studies b y Fanke & Ries (8 1) ed tem to concude tat "18:2n6 acks EFA actiity in te cat and tat the syndome encounteed was a dietay deciency in sme o al of the deied FA They noticed that te condition of te sin and coat was wose in cats fed diets containing ydogenated coconut oi (acking inoeate) th in those fed diets containing saowe seed oi (igh in linoeate) owe e, they atibuted tis to accentuation of aacidonate defciency by te s y acds i the cocout oil (9. Sincai et a (232) epoted tat cats fed a diet cntaining satuated beef fa t deeloped signs of FA deciency i. e. lethagy sow gowth dy scay skin ad skin esions. Tey aso epoed tat saowe seed oi eesed te defciency signs 15 but e data ae not been ubised Macond et a (150) found tat cats fed a diet ow in inoeate (03% of tota kiocaoies) ad ow feed efciencies  scay skin ai oss enaged fatty ies, and makedy inceased ates of wate ss tou te skin Necosies eeaed degeneation of te kidneys testes and adenas nd yeeatosis of te skin (149 152) Tese obems wee peented by dietay inoeate (5% safowe seed i) wic demonstated tat inoeate is an EA fo te cat

OEATE DECNY


EFA Reiements Te EFA equiements of cats ae not yet been ecise detemined . Atoug inoeate at 6.7% of kicaies is moe tan adeuate f gowt and noma skn condition (150) te minimum equiement is pobaby ess tan tis. acuations f te doube-bond index f ie iids (19) based on compaison to data fm ats ( 1 15 157) indicate tat te inoeate equiemen t of te cat may be ne 25% of dietay enegy in te absence of dietay aacidonate. Te discussio of quantitatie equiements is compicated by two factos ( Aacidnate aone wi pobaby meet te tota EA equiement ofte cat  as it does in te at  15  157) so te inoeate equiemet o the c wil be negigibe if tee ae ig ees of aacidonate in te diet. b Wit inoeate at 4 8% of dietay enegy and aachidonate at 004% eoductie efo  mance is essentiay noma (152) wic indicates tat 0.04% of kiocaoies is nea te minimum equiement fo aacidonate. Howee ow ees of dietay inoeate woud be exected to incease te aacidonate equiement . e eason fo this is tat bot inoeate and aacidonate ae actie in main taining membane integty   7 . eefe  atug a souce of aaci donate is indispensabe fo te cat inoeate migt sae  educe te

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MDOALD ROGRS & MORRS

acidonte euiement Fue studies ae needed to untitte te inoete and cionte euiements wen ot ae pesent in te iet n osetion tt desees ute study is tt toug inoete peents lie enlagement ue to tty inttion te neutl t content o lie emins ige tn noml ( 49)  ecuse lees of inoete in tissues e nea maim wit diets contining % sowe seed oi ige ees o dietay linolete woud not e epecte to e ute eneci eects is ises te uestion o wete  souce o tty cid tt is moe poyunstuted tn linoete is needed to minimize te ccumution o neut t in ie In ts ie fy cid syntesis is eguted in p by ees of poyunstuted fty cids (), and acidonte my e moe eectie n inoete (4) ietay acidonte o ote polyunstuted tty cids my e euied y te ct o nom ie ft metoism Befoe  minimum acidonte euement can e ssessed  study is needed o te mount o neut t in te ies of cts fed difeent dietay ees nd combintions of inoete nd acidonte e lck o 6destution wic peents te syntesis o acidonte woud so peent te conesion o 8n (inoente) to onge-cin n3fty cids. ies et  (99) suggested tt tese ftty cids my so e euied by cts Becuse n3-ftty cis cn be coneed to eicosnois tt can e aious pamcoogic effects te ct woud be useu in te study o te pysioogic oes o tese tty cids e domestic ct is not te ony cioe tt cks te biity to destute ftty cids t te 6 position e ion (98) te tubot ( cioous s) (  8); and een n insect cioe te mosuito (8) e simia iniities to syntesize cidonte is metoic pecuiity ppes to e coeted wit species t ae sict cioes egadess of tei position on te pyogenetic tee. UARY

Fom te foegoing discussion o te nuttion euiements nd some o te metoic nomies o te ct it is cea tt te ct is dpted to eting  cioous diet It my owee e ess cbiity tn omnioes nd eioes to dpt to wide nges in diety omposition o empe te ck of iity to syntesize sufficient itmin A fom cotene oitine om gutmic cid acidonte om inoete nd tuine om cysteine esuts om  compete deetion o seee imittion o te enzyme o ptwy tt mkes ec nutient. Ote nuient euiements suc s te bsoute euie ment o nicin nd te ig potein euiement ppe to esut fom te ig ctiity of one o moe enzymes nd te fct tt tese enzymes e not dptie in te ct Fo empe te ct cnnot decese picoinic caoyse in ode

I UTRITI


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to fo tytoan towad t niain-yntti atway 2 no an it ca t ua cyc nzym wn dty potin i cad n t t n o to con niogn (209. Indd t at appa to a  aabiity to adat to mot ang in itay omotion bau it annot ang t uantiti of nzym in od in t mtabo atway (209. T outionay dopmnt a ultd in mo tngnt nuttona umnt fo at tan fo omno uc a t at dog an man Wat itt dnc xt fo ot cio i ad u to uggt tat ti att may w b ommon among ot it co. T mtaboi diff� btwn t at and omno id t  c wit a ufu nma mod fo tudying t biomica ba of om nunt umnt. Fo xamp bcau t  no gnficant conon of inolat to aacionat n cat i (0  0 23  t pyoogical funtion of inoat an b dtmind indndnt of it aing a  a a puo of aidonat ( 0 . T a not bn poib wit ot pc It i anticipat tat fut tui of t nutton of t at wi inca ou undtanng of mtaboc aaptation ad nutnt functon. ACOWEEME T auto wi to xp ti aiation to  Kan n fo  inauab atanc in t ppaton of ti w. Literature ited I dae  E. 176 The nteaton o hnge and peyng n the doest at (Feis cats n adaptve heahy? B . Bio 18:67  dolph  S 17 Toleane to heat and dehydaton n seveal spees o aals A J Psio 1515675  had B 11 The ate o aotene ate absopton n the ana ogans ice. J 511510  ken T D Shnep L E, Fobes  M Cobn E 178 The eetoa owzn de t on the gowth and skn on dton n ats Prc t A Meet A c. Ai. ci Midet ect. p 7 (bstr 71) 5 llson J B   Mlle S   MCoy  Bsh  K 16 Stdes on the ntton o the at N A Vet 78 6 nand B K   Bobek J  151 Hypothala onto o ood ntake n ats nd ats ae J Bio Med 1

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10168 8 ndeson   Bake D. H   Cobn J E Helpe L C 17 Bohea esons assoated wth tane deeny n the at J Ai ci 17  ndeson P  Bake D H Shey 180 Ntogen e P  Cobn qeent o the ktten A J Vet Re

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phenylalannetyosne nd tryptophan eqeents o gowth o the yong kt ten' J Ai ci. 5078 ndeson P  Be D  H Sery 180 P    Teete  G Cobn heonne solene valne and ene eqeents o the ong ktten J Ai ci 506671 1 ndeson P   Bke D H   Sherry   Cobn J E 17 Cholne ethonne nteelatoshp n elne ntton J Ai ci 57 1 ndeson  S 18 Wate balane n

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DND RR & RRI

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te dog and a. J.  Ai Prat 2358898 14 od A Shad J S. 954. Nirogen baane tude wih dog on aein or ethonneeene aein J. Nutr 5325-73 15 Aho J 1979 Ciadian rhthm: genera feature and endorinoogia a. In Edorie Rts ed. . T. eger  1-61. New York Ra ven 1 Aiaion of Ofia Agruura Che 1980. Oia Metods of Aasis Wahingon C: A O. Ag Che. 13h ed 17  Avion P V  Wion J C 959 On he bioynhei of feinine Bioi Biops Ata 34:27981 18. Baard F. J. 195. uoe uiizaion aaian iver Cop Bioe Psio 14:437-43 1 9. Bnk  R  Sabnfed   19 83. Proan in the Ca: I iua pate and phooriod effe. Bio Reprod 28933-39 20 Baa A  Hxabe R J 978 aurie ad Neuroogia Disorders New York: Raven 2 1 . Bet . C  Burger I H 980. Taune defiieny einoahy in he a J a Ai Prat 2:521-34 22 Barohuk  M Hed M A Prk  . 1 97 1  ae of waer in he a Ee on uroe preferene i ee 171:99-70 23 Batohk L M  Jaob H L  Niho .   Hoff L A   Rykmn J J. 1975. ae rejeion of nonnutriive weeener in a. J Cop Psio Pso 89:971-75 24 Beauham  K   Maer Roger J.    Jr. 1977 Favor preferene in a Feis ats Patera p. . Cop Psio Pso 1827 25. Beaer B 8 Veterar Aspets of Feie Beaior pp 40 S. Loui C V Moby 2. Bedford P.  Carke   C. 19 72 Eperiena nzoi aid poion ing in he a Vet Re 905358 27 Beivea  P  Freedand R A  982 Meaim of erine gine and thonine in ioaed a hepaye Feis Doestia Cop Bioe Ps  7B: 1318 28 Beho, R W Agui  . nd Beho M B. 1974 Feine enra reina degeneraion Iest Opta o 13:08-1 29. Beho R W. Fiher C A 1970 Feine enra reina degeneraion  A Vet Med Asso 57:842-49 30 Benedi F  938 Vita Eergetis A tud i Coparatie Basa Metabo

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is Cegie In Wah Pb No 503 25 pp Band Suton J 1888. ike in mon ke ion ars and bird.  Cop Med urg  Boudreau J C 1 974 ea ening in a geniuae gangion ongue uni. Ceia eses ad Faor 14-5 Brodie B  B   Reid W   197. Some haoogia onequene of  ie variaion in rae of meaboim Fed Pro 2: 102-70 Budde E. F. 952. he deerminaion of fa in baked biui  dog food J. Asso O Agri Ce 35799 Bdge E A W 199 e ods ofte Egptias 244 New ork over Burger  H   Anderon R. S . ome . W. 1 980 Nutiiona faor affeing waer baane in he dog and a Ntri tio ofte Dog ad Cat ed R S Ander on pp. 1455. Oford Peagon Burger I   Baza S E Kenda P. T 98  . The proein reuiremen of adu a Pro Ntr o 4002A. Abtr Cnon W B   Newon H  F  Brgh E M   Menkin V   Moo R M. 929. Soe ape of he phyioog of ani a uiving ompee exuion of ympahei nere ipue A J Psio 89:84-107 Carey C J. Moi J  977 Bioin defien in he a nd he effe on hepai popion CoA rboae J Ntr 07:3334 Caner J A 195 Sie differ ene in ae preferene  Cop Psio Pso 49 1 39-4 Crner T M. 94 he effe of uga on he repirao exhange of a J. Nutr 28315-23 Caongua . W 198 iear diu ion and inake in he a Psio Bea 275479 Chew R. M 195. Waer eaboi of maa In Psiooga Maao g, Vo 2 ed W V Maer R.  Vaneder pp 4-17 New ork Aademi Cark L  Seawigh A A 198 Skeea abnomaiie in he hindimb of young a a a reu of hypeiaminoi A Natre 2177-7 Carke B A   Carke S . 19 82 Sp preion of ra iver fay aid ynhei by eioa5 81 1- 14eraynoi aid wihou a reduion in ipogeni en zyme J. Ntr 12:122-19 Cok E V 959. Oeodroph in Siaee kien Vet Re 748 Coffin    ozworh J. 954 "Ye ow fa in wo aborao a aidfa pigmenaion aoiaed wih a fihbae raion. Coe Vet 43-7

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MacDONALD, OGES & MOS

glumylurin (liorlon) on h m phibin mmorphoss Co Bio ch Phio C:7-7 8 inns R. N T Grhmons O. 90 Stdi of  nuriionl dis (oodyophi ibro) of young lion ocid wih chng of h klon nd ympoms of muculr wkn. Proc. Zool Soc Lon 3357-9 79. Fox,  A. D   nsn G R  Knox, K L 973. ffc of vriions in proin quliy on growh PR PR nd P in gowing kins Ntr Rep Int 7:2-3 0 nl , T. . 980. Essential/a ci cinc in th ct (Fi tu ) h D  hi olfon Col  niv Cm bdg, .K.  Frnkl, T , Rivr, . P.  97 Th nuronl nd mbolic impc of linolnic cid 83) on c d pivd of niml lipid Br J. Nutr. 39227-3 2 . Frngs, H 1 95   Sw s in h c nd h scrum. Exrinti 7 424-2 3. Fukuom K. 9  Biosynhis of iso vlhin in h c J Boche (Tokyo) 49444-45 84. Furk A. Sbsyn F Fur L Hovh, A., Hrcl ., Ormi, S Bnyi B 90 Iolion of gmm glumyl urn om h proinfr quous xrc of boin phyoid powdr ct Biochi Bioh c ci Hun 539-47 85 Grhoff S N 19   Th nurionl rqurmns of cs. Proc ni Cr Pn   49-52 8 Grshoff S N  Ands S B . , Hgsd, D M., Lnini,  A 957 Vimin A dficincy in c Lb. Int 227-40 87 Grshoff, S N Frgll, . . Nl son, D A, Anrus S B .  959 Vi min B dfcincy nd oxl nphrl cinoi in h c  J  277280   Grhoff S N  Golib  S 19 nohnic cid dfcincy in cs J Nutr 235-3 9 Gshoff S N   gg M A O'Con nor F. , Hgd. D. M 957 Th ffc of vimin Dficn dis con ining ou CA rios on cs J Ntr 3:79-93 Gssr, C F   H 95 ro in in h nuiion h gowing dog Nutr 5:452 91. Gdmn, D S, Hung H. S., Kni, M   Shiror, T 197 Th nzymic conrsion of llrn�cron ino inl J Bio Ch 242354354 92. Gs P 195 Proin nd clor rqums of h flin n Canne and

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Fin Nutrition qirnt d. O Grhmons pp. 3345 London P gmon 93 Grvs .  Sco  P.  90 Nu ion of h c. 3 ron rquirmns for niogn qulibiu in dul c minind on  mixd di Br J. Nutr 43-9 94 Grs, P , Sco M. Sco, P P 1959 Thyroid chngs in c on  high proin d rw h. Phio 14 7P 95 Grnsin P., ini, M Gullino P  Bbum S M  , Oy, M. C 95 Sudis on h mbolsm of mino cids nd ld compound in vivo. I  vnon of mmoni oxiciy by ginin nd rld compounds rch Bioch Bioh 342-54 9 Hldn 892 A nw form of ppr u for muing rproy xchng of niml . Phio 34930 97 Hmr, D, Cho .  C Dysr M I Rich, L 97 ffc of s um chlorid in prnion of ximn lly producd phoph uolih in ml cs  ni Ho oc 2 5-7 9 Hrdy A Morrs G Rogrs, Q. R 977 Vlin rqurmn ofh grow ing kin J Ntr 07308-2 99 Hrgrov D M   Rog Q R  Mor  G 193 Th rypophn rquirmn of h in Br J Nt 5047-93  Hslwood, G. A D. 94 Th biolog icl ignifcnc of chmcl diffrnc in bil l Bio  3953774 0   ssm A G  Rirs, P . Crw ford M A 977 Th fur of h c o dsr linolic cid is nuronl im plicion Ntr tb 2322 02 Hwking   obbn M C   Gm mg, K., orm M   97  . Circdin rhyhms ciiy mrur urn, nd microflrl) in dog, c hn duck, hnou nd Grbi J Intri cpl Cycle Res 245573 03 Hys, K C 97 A riw on h biologicl ncion of un Ntr Rev 345 04 Hy, K. C  Cry, R  , Schmid S. Y 975 Rinl dgnrion id ih urn dcincy in h c. Scinc 88949-51 05 Hys K Rbin A. R. Bron,  L. 975 An ulucurl udy ofnuti onlly inducd nd rrsd rinl d gnrion in cs.  J Ptho 7: 505-24 0 Hy K Surm  A 198 Turn in mbolism. nn  Ntr 4025 07 ndson,  . B Kyoo E K.

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zymes of oithine metalism in dult nd developing rat intestine Bchm

Bh. ct. 428610  lO Heywd R 1967 Vitmin A in th� liver nd kidney of some felidae Br. et  123:3996  l   Hils D Morris J G Rogers  R 1982 Potssium requrement of ktens s ected by dietry ptein  Nutr 1 12: 2122 112. Hisch E Dubose C acobs H 1978 Dietry conl of f intake in cts Phl. Beha 20:28795 113 Homn H H 1953. Repr on cde f in aked dog food  Aoc. . Agrc. Chem. 36:208 11 Holman  T 1960 The rio of

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iological ctiities of polyunsatuated ftty cids T le Fat n Human Nutrtn ed A J Vergrsen pp 3315 1 . New York Acdemic 118. Humphys E R Sco P P 1962. he ddition of herng nd vegetable oil to the diet of cts Prc Nutr c. 21:18 19 Hui H 1981. Dy length and reeding in he domestic ct b nm. 1522

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1

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6


MacDONALD ROGERS & MORRIS

h nuition of th cat Poc Nt. Soc 33A (A) 8 Rivs, J. P W . , Hasa A. G ., Craw 19 ford, M A  , Brall, M . R. 1976 The niiy of te ion, anthera Leo, to ut linolic ci. FBS Le. 729-70 199 Rives, J. P W Sincla, A. J. Craw ford, M A. 19 75. nability of th at to aut sntial fay aci . Nate 2517173 2. Ros A. H   Sott P. P. 1961  Nutri tion of h cat 5 The inuene of al i ad iin spplnts to a at iet on he retetion of nigen, clcium an hoo. Br. . Ntr. 15732 201  Ros, R. N. 196   sty offelnne and ts exceton by the cat Ph.D. hesis Stat Univ N. uffalo 202 Roinon, R 197 1 . Genetcs fo at Breeders Oxfo Pgon. 191 . 20. Rog Q R Fedlnd R. A. Sy ons, R. A. 1972. n vivo synthesis nd utlization of agnn in th rat. Am. . hysol 223230 2 Rog Q. R  , Cn, . Y , Hr A. E 1970. The ipo of dispnsable amino acids for maximal gowth in the rat Poc So xp Bo ed 141 22 205 Rog Q R  Hr A E 195 Ami no aid diets d ial growth in the rat. . Nt 87:267-7 20 Rogers, Q. R Mo, J. G. 1979. Eentiality of mno is for th gow in k n  Ntr 1097182 207 Rogrs Q. R. Morrs, J. G 1980. hy ds he cat require a high potin dit? S Rf. 36 pp 45-66 208 Roges Q. R .  Mos J. G. 1 982. Do cats ally ned more otin? n Walth am Symposm, No , ecent Adances n Felne Ntrton, d. A. . B. dny, . Small nm act 2321-32 209 Rog Q R   Mos . G   Freland R. A 19 77 . Lak of hepatic nzyati adaptation to low and high lvls of diet ary ptein in th adult cat. Enzyme 2234-5 210 Ror A S. 1966 Vtebate Pleon tology Chicago: Univ of Chiago Press. 468 pp d d. 2 1 1  Rosos, D. R.  Palmer H. J. , Muir  L, Bennink M R. 1981. Inunc of a low bohydrate diet on fo e of regnnt ad latating dogs.  Nt. 68-89 2 1 2 Ros, G . , Dunn, D. , Jone, M E 197 ihin ynthi om glutamat in rat intstinal mucosa hoognats Evi dence for the duction of glutamat to glutamyl smialdehyde. Bochem. Bo phys. Res. Comm. 5:1407

III

2 1 . Rowland N. 1981 . Glucogulatoy feeig in cts. Physo/. Beha 269013 2 14. Rowsll, E. V.  Cie J A ahbi, S D. , Ai, A  .  Ro, K  . 1979 in hyat an eny vat minorasfra ctivitis in ir ent animal scis. omp Bochem. Physol. 65455 215. Ruin, L. . 193. Ay of  n con in th ct etin. . Am. Ve. ed. ssoc 14214120 216. Rubin, L. F , Liton D. E. 197. Rtinal degnrtion in ittns.  m Vet. ed ssoc 12479 217. Schaffr M. Rogr, Q R . o,  G 198 Methionin ruent of th growing kittn in the absenc of diet ry ystine.  Nt. 1 12962-71 21. Sceffe,  C ., Roges, Q. R  , Mo  192 h choin rumnt of the growing kittn in the rsene ofjust adequate ditary ethionin. Nt Res. 2:2899 219 . Schller G. B. 1972 The Seren et Lon. Chicago niv. Chicago P. 480 pp. 220. Shimk R. T. 1962. Adaptive harac terstis of ura yl enzyes in the rat . Bo hem. 237459- 221  Schwa R S   rhm S  192 Effet of dietry polynsatrated fatty aids on the ativity and contnt of fatty aid synthetase in ous liver. Bochm Bophys. ta 711312 222 Sco P P 1959 Clcium  ioin diny in atfd ats with fer ence to ostognsis imrfecta. Poc B Small nm Vet ssoc, . 84 223. Scot P. P. 1 9. Nution n Dis. In Felne edcne and Sge, ed. E. J. Catott pp. 11-144. Snta rbra, Caif.: Am. Vet. 2n. d. 224. Scott P. P. 1965. Minral nd vitmi in flin nution. S Rf 92 . 788 225. Sott P. P. 1 968. Th scial featurs of nuttion of ats, with obations on wild lidae nuttion in th London Symp Zool Soc. ondon No 21, p 21-6 226. Sott P. P. , Gravs, J. P. , Scott M G. 191. Nurition of the ct 4. Clcium and iin dcincy on a mat dit. Br.  Nt 15:5-51 227. Scott P. P.  Graves J P. , Sott, M G. 19 Nutrtional blindness in the at. xp. ye Res. 357 228. Scott P P  Sco M G 196 Vitmin A nd repoduction in the cat  epod. Fetl. 8:27-71 9. Sawright A. A  , English, P B  , G ner, R. J. W 1970 Hyitaminoi A



G

Z

FELNE NTRITON o the cat. Adv Vt. i Comp d 14:1-27 hapio I L 1962 In vivo studis on th mtaoli rlationship twn flinin and srum holstrol in th domsti at hD. thesis. Uiv. Del.  Newark iclair A. J   Mca J G, Moger, E A 1979. Metalism of ioleic acid i the cat Lipids 14:93236 icar, A J   latte W  Mca, J G  Moger E A 198 1 . Essetial atty acid deciecy ad evidece or aach doate sythess  the cat Br J. Nutr 46:93-6 kltety, M. 1966 Chages i caoic itake oowig rai stem esos i cats. II Eects of lesios i medial hythalaic regios Arh Nurol 14:541-52 ktety M. 19 Chages i caloic itake olowg ra stem lesios 

Decker R H  Hedeso L M   Hak es  V 1957 cies varatio i the metalis of 3hydroxyathraiate to pyridiecarxyic acds Bio. Chm 228973-82 245 Teeter R G  Baker D H  Cori J E. 1978. Methioie ad cystie qi mets o the cat J Nutr. 108291-95 246 hoe C J 1982 eeig havior i the catrecet advaces. I Waltham ymposium No , Rnt Advans in Flin Nutrition, ed A. T B Edey J mall Ani. Prat. 23:55-62 247. itchea C A. Roges, Q R., I diei, R J. Mois J G 1980 hreoe mbalace deficiecy ad erologic dysctio i the ktte J Nutr. 1 10:2444-59 248. Ullma D , precher H 197 1 . A  vitro ad i vivo stdy o the covesio o eicosa   14dieoic acid to eicosa

qedcta gray atter ad rostral hy thaams Arh. Nurol 14:670-80 235. ktety . M. 1969 Alteratios o caloie itake  cats folowg lesios of the hythaas ad midbrai Ann. N.Y. Aad i 17861-74 236 kltety, . M  Gary . M. 1962 Ex imeta hyhaga i cats ollowig destctive mdbrai lesios. Nurol 12394-401

sio of ecosa   eoic acid to eicosa 5   dieoic acid i the rat Biohim Biophys Ata 248:186-97 249. Ude E. J 1977. ra Elmnts in Human and Animal Nutition New Academic 4 p 4th ed 250. Vessey, D. A 1978. he biheca basis for the cojgatio o ie acids with either gycie or tarie. Biohm J. 174621-26 21  Wag, T. M  1964 Futhr studis on th iosynthsis of flinin h.D. Thesis. Uiv. De Newark 252. Waterhose H. N Caer, D . 1962. Growth rate fo ad calorie cosmp tio of laratory cats Pro. Anim Car Panl 2:27174 23. Welch, R M. , Coey, A H.  Bs J J 1966. The metaism of acetopheet di ad Nacetylpamiopheol i the cat. Biohm Pharmaol 15:521-31 254. Wergedal E., Harr A. E. 19 Metaolic adaptatios i higher aimals IX. Effect of high protei itake o ai o itroge catabolism i vivo J Bio Chm 239: 1 1 663 255. Westall, R G 1953 he amio acids ad other ampholytes o ie. 2. The isolatio o a ew slphrcotag ao acd f cat re Biohm. J. 5:2448 256. Deleted i pr 257. White, T D, Boda, J C. 1975. Taste prefereces of the cat or ero physiologically active copods. Phys io Psyo 4-1 258 Williams R. . 1967 Comparative pat tes of dg metaoism Fd Pro 26102939 259. Widmeler H G .  paeth A E. 1974 Uptake ad metalism of pas

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kites gve aio acid dets cotaig

adequate cystine

Br.

J

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I7

238. mth D C. rott  M 1944 De veopmet of thiaie deficiecy i the cat o a det of raw fsh Po. o p Bio d 56:13 239 pray C M   Widdowso E. M. 1950. The effects o growth ad developmet o the comsito o mammals. Br J. Nutr 4:332-53 240 tepha Z  Hayes K C 1978. Vitami E defciecy ad essetial fatty acd EA stats of cats. Fd Po 37: 2588 241  tea M B .  Kass T  ehma, D. Cemete C. D. 1965. Circadia seep nd wking pattes in the laba toy cat Eltronph. Clin Nuro physiol 19:509-17 242. tewa, . M  Batshaw M.  Valle, D. Walser M 1981. Effects of argiie fre eals o eageesis i cats. Am J Pysio 41:11 243. tma J. A Hayes K C 1980 he iology of tare i tritio ad developmet. Adv. Nutr. Rs 323199 2. hadolk R. teves C.

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