nutritional energetics of animals

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NUTRITONAL ENEGEICS

OF NMLS

R. L Baldwin and C Bywat

Dep atment of nial Science niversity of alfoia avis aifa 66

CNTENTS

INODUION NOOG O NUIONA NGIS 3

Description of Energy Transactions in Animals xpression of Animal nergy Requireens and Feed nergy Value

ASA AOS AND AINNAN QUNS 6Soure of Variaion in Baal Meabolim 7

PODUIV POSSS AAIA ANAYSS O NY AAN DAA

INTRODUTION

Eegy geeally ee tems o potetal apaty to peom wok, s aabsato that a be measue oly ts tasomato om oe om toaothe 2). Amal uttosts ae oee pmaly wth two oms oeegy hemal a heat Eegy supple to amals hemal om a beasome to heat whe utlze suppot o vtal le poesses, stoe hemal om as gowth o fatteg, tasee hemal om to aseo amal as pegay a latato, o tasee to suougs as wok ). Quatyg these eegy tasatos amals s the ous o thee o utoa eeges

The ist a seo laws o themoyams hol that all oms o egyae quattatvely oveble to heat Classally, all measuemets o eegyasaos mae by ama utosts wee mae a epesse tems

01885/84050 101 $200

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102 BDWIN & BYWTER

o heat eegy o aoies (3 Although Klebe (31 stogy uge that thealoie be etae as the basi uit o heat eegy, the joule was aopte the teatioa System o ts a by the S Natioal ueau o Staas

as the peee uit o expesso o eleta mehaial a hemaleegy (4 Subsequety, most uttio jouas aopte the oule as thebas uit o expessig eegy. The joue s eie tes o the Iteatoal Met System as oe gmse o 0 eg. The utoa aoe isee as 44 eause al measuemets o eegy e eie tes othe uametal mei uits o mass kg) stae m) a time se) jouesae eaiy ovete to aoes a vie vesa. I ths pesetato, the aoeis the basi uit use to expess heat eegy

The majo objetive o utitioal eegetis is to evise a oveet a

auate system to pet the eegy baae o viuas base o boyweght, sex, ativty, physoogal stae, eviomet, a amout a utive vaue o ee eate (11. Commttees ommssoe by the NatoalReseah Coui NRC) a the tsh Agultual Reseah Couil ARC)have evewe the geat mass o expeimeta ata o eegy metabolsm awide range of speies inludng man, and have formlated systems to predt

the eegy a ee equemets o these speies (1 2 434. Theseommittees have pubshe summaes o the eets o boy weght, omposi

to o oos a ees ompositio o pouts suh as meat a mi age,eviomet a othe atos a have popose eeig systems thataomoate these atos atal eiiees o use o ateate utets ovaious poesses have bee osiee a ompomises aopte, ote view o appaetly oitig vews a ata (1 2 434. Some o thesystems wo exemey we whe appie aeuy a with ue osieato o mtatios impose by ompomses mae evelopmet. Mostsystems ae epeatey upate base upo ew oato a geeaysystems that ye impove peitos o eegy equiemets o a gve

speies ae aopte whe avalable (22). No uetly avaable sstem yesaate etae, eve wh a sge speies ue a oios a ouet system has uvesal applabilty Thus the geeal objetves outitoal eegets ae to evse moe geeay appliabe systems a tooulate iets that optimize pouto eeies.

The age amout o ata avaable o eiiees a ee values ssummae ompehesey AR NR a Eopean Aocaton oAma outo EAA) pubatos (1 2 43 a is ot eviewehee Rathe this eview ouses o the soues o vato eegy equiemets that limit geeal applatio o uet eeg systems, the eseahiete at ietiiatio o ueyg physiolog a metabo auses o thsvaiatio a eay attempts to quatiy a aommoate ths vaiae ompute moes o aimal gowth a metabosm

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NTRITION ENERETIS O NIMS 10

TERMINOLOGY OF NTRITIONAL ENERGETIS

ince the oina Quantitation of Combustion in a livin system y Lavoisie,many tems have een invented and appied to descie the types of eneyansactions that occu in animas ( 7) Histoical tems such as "specifcdynamic acton (of food) have appeal and ae stil used even thouh theyimply nothin aout undeyin eationships othes, includin "wok ofdiestion have specific ut incoect implications eadin unelyin elationships. Diffeent schools of nutitionists have tended to adopt and defendateative systems of nomenclatue to descie the patition of eney inanimals. The iennia symposia on eney metaolism of the EAAP (12, 18,4 55) have heped emove many dispaities fom the teminooy Recenty a

sucommittee of the ommittee on Animal Nutition of the NR wasappointed to develop a systematic teminooy fo desciption of eney utiizaton by anma whch conen wh feein em n cuen ue n he

and teminoloy appied in othe counes 45). This system is pesenteddiaammatically in iue

TOALREEAS

HEAE ()

INAKE OF EGY NOOD (I

GESE NEGY(

MEABOLZAE NEGY(

E

AS

A NEGY(

OS NEGY(

WASE NEGY: UNE (U, (O AE (

ECOVEED EY ((USEF POD

Fiue Pato o gy ama Adaptd om NRC 45)

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104 BALDWIN & BYWATER

Desription of Energy Transations in the nimalThe classical tem oss eney intake has een elaced y the em otal intakeof eney E) o inested eney wich s dened as the amont of food

consmed (I) times oss eney e unit of food (E) onvenional defnitionsof aaent diestile eney DE IE E) and metaoliale eney ME DE GE VE) ae eained (see iue fo defniions of ems). Eneyin eceta can e fom nonuilized comounds in inesed (su i) food EVE) o of endoenos (su e o metaolic (s m) oin m VE) so that e diestile eney TDE) equals IE - + E + E HE E whee HE is hea eleased in femenation din diestion. imilalye metaoliale eney (TME equals TDE VE VE. Waste oductseceted via the ills of fis (Z) o fom he eeo suface of he animal (E)

ae eated similaly o uine The em ecoveed eney RE) elaces theolde collectve tem Net Eney in Podct NE) and idenifies eneyecoveed in useful oducts such as issue (T) milk (LE) es (E)concets (YE) and wool hai o feates (E) Vse of he em RE oviaesconfsion etween NE vales assined feeds (discussed elow) and eneyansacions in the animal.

Patiton of tota heat odcton (HE into meannf hysioloic ometaolic comonens is the mos comle and conovesial asect of all

systems of nomenclaue. This is due in at o(a)

a lack of knowlede andqanticaion of the hysioloic and meaolic ocesses tha contiue ototal heat odction; (b) systematic diffeences amon estimaes of comonents of heat odcion tha shold e idenical on mathematical ouds o ydefnition ut which diffe eause alteate mehodoloies may eateeney eenditues associaed wih somewhat diffeen metaolic ocessesand c the attemt o econcile eminoloy wih academic o eseach oalsad he amaic oal that the teminoloy accommodate cuent and fefeedin systems 22 45) These olems will e elaoated uon in suse

quent sectionsomonents of total heat odcion elicily identied in the NR system(45) ae asal o fasin meaolism (HE) hea of voluntay acivity (H E)heat of odct fomaion HE) heas of diesion asotion and assimilation (HE) hea of themal eulation (HE) heat of fementation (HE) andheat of waste fomation and ecetion (HE) HE HE HE and HE can eefeed to in aeate as hea incemens (HE) of maintenance and odction

The tems dened aove allow desction of eney tansactions within anaimal in accod with the st two laws of hemodynamics. o eamle aactatin owin animal miht aition inested eney as follows: E GE VE + E HE + RE whee RE TE + E and f desied HE

H H H H H + H H. ue ao fo eame

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NUTRTONAL ENERGETCS OF ANMALS 10

of H into costs of milk lactose protein and fat synthesis, and body fat andlean snthesis is enissible as long as the equalit of inut and outut remainssatised.

Expression of Animal Energy Requirements and Feed Energy Values

Animal energy requiremens and fee energy values must be expressed inmatile tes rl in this entu the limitatins f sstems ase ungrss energ-riailit in digestibilit-and digestible energ-rabilitin urnar and gaseous losses-were reognized and several sstems basedun meliale energ nets were deele (3 mng these werethe Phsiological Fuel Value (PFV) and otal Digestible Nutrent (DN)

systems whih re still in use Speifi limitations of these MEbase systemsre at rtein is unealued in fastgrwing animals an effienies f Muse fr rous hsilgial presses ry systematially with hemical andphysical properties of feeds includin alori density (E hese limitatins led to the deeloment of net energy (N) sstems which alue feeds intes f their efien f use fr seifi resses base un the identityNE RE where can be a pitie or negatie quantity, an I is feedintke in grs In imals fe at r ew maintene (energ balane � 0)

he net energ alue of a feed for maintenance (NE) equals the amount f bodyenergy spared (TE by an increment ( f feed (NE TE n agrwing animal fe ae maintenane the net energ alue f a fee frgwth (NE is equal t the amunt f b energ stre (T) ue troision of an additional increment dI of feed (NE TE REThus net energ alues of feeds for maintenane growth fattening latationegg roduction etc can be estimated and tabulated he net energ requirements f an animal an be alulated as the sum of functions fr which energy isrequire Fr eamle fr a grwing regnt latating animal NE N

TE L Y where NE equals fasting cataolism or basal metablismplus enoenous an metabolic lsses in urne and fees and from te surfaceand b conention nonal actiity (NE HE VE E SE HjE

NE and M systems re highly cmpatible because feed alues and animalrequreents eressed accrding to oe system can be coneed readil t theother based on the eciency with which metaboliable enery from a feed isused for specific functions. Fr eample, efficiencies of ME use for maintenance (K) nd growth (K) an be calulate as K NE/M an K

N r T/(MI M) where M is he amunt f metabliale energfm a fee required t maintain eerg blane (B 0) an M ismeaoliale energy intake. Costs of igestion absortion d assimilation(HE) and of rodut snthesis (HE) can e aggregated as HE suh that M

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10 ADWIN & YWATER

= H and M /K + /K The ae fueuans povde a uanave ln amng uen feedng ssems suh as heARC ssem (2 wheen feed alues and anma equemens ae epessed n

a M ass and he RC ssems whh ae ased n (43.The ae dsussn summes uen en n nuna enees and he undeng nepua fame upn whh he eng and msuen feedng ssems ae based' The maj pbems enuneed n applan f he neps and feedn ssems e ha esmaes f pamees suhas H K K e an va gea mng anmas feeds and mehd fesman Suseuen sens eamne sues auses f hs aan

BASAL METABOLISM AND MAINTENANEREUIREMENTS

ae n he as enu Rubne ( 3 ned ha basa masm heapdun was n a lnea funn f bd wegh BW) ehe whn beween spees u ahe ed as an appae funn f suae eaBW213 Suseuen Bd (13 and Klebe (32 eped ha he bessasa beween d weh and asa esng hea pdun n ungaduls ass spees was aned wh he euan X BW0775

whee s basal hea pdun n al/da s a een wh uns fal/(da X BWO7 - 7 5 )and BWO7 .75 s meabl bd wegh (MBW)A an ea AA Cnfeene n neg Meabsm (12 he well nwneuan BMR = 70 BWO75 was adped f esman f basal heapdun n kalda n ung aduls n an nespe ass Bh Bd(13 and Klebe (32 egned and ndeed emphased ha he eanshphe eped was mahemaal empa ahe han heeal as was Runes suesn ha BW e used The as ned ha he bes empas daa whn spees wee aned wh epnens he han 0.7. Ths

ae fa has been eemphased een b Thnne e a (4) Gees &Mhels (23 and hes The desn se he epnen a 0. b nvennwas pagma ne easn was faae mpasns amng daa poduedn dffeen aaes whee epnens angng fm 00.7 wee hen nuse A send pehaps mp easn was a hee was n heeaepanan f wh he empa nespe epnen (0.7) dffes fm0.. uhe hee was n epanan f wh epnens dedued sasalfom naspe daa n basal hea pdun anged fm 0.40.8 (4. nvew f hs unean seemed ana as a s e epnen n healme euan hee fng a ane n esmaes f he effen , eamne fas ha aused he efen a and lae devse an apppae mahemaa fmula Ths appah was adped ndevepng subseuen dsussn f basal hea pdun hs s n nm

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08 ALDWIN & YWATER

liver wrk repirain, in and meablie rerpin by kidney, and nervuand endrne funin relaed inegrain and (b iue and ellular levelfunin uh a in ranp fr mainenane f membrane penial and

marmleule reynhei uver r repair) Eah ype f funin aunfr 3550% f baal energy ependiure (5, 8) Mlligan (39 wa ne f hefr emphaie ha energy ependiure in in Na) ranpr are majramuning abu 30% f baal energy ependiure, and ha hey may varyignianly wih phyilgi and endrine au. The laer uggein wa'reinfred by bervain by Imail-eigi & Edelman (29 ha energy ependiure in iue varie ignianly wih hyid au a a reul f he rlef hyrin in regulaing aiviy f he uabain-eniive Na K +)APae.Reenly, regg & Milligan 6) pariined energy ependiure in mule

frm ld-eped, w- and even-week ld lamb and laaing and nnlaaing ewe in Na,K+)-Taedependen and independen repirain.hey fund ha he inreae in repirain in mule afe epure ld walargely Na ,K)-ATPaeependen, in agreemen wih previu repr27) Repirain rae were 32% greaer in laaing han in nrl ewe and7 f hi inreae wa a,K)-aeependen. Repirain rae fmule frm w and even-week ld lamb were 50% and 20% greaer han inmule frm nrl ewe. The dereae in mule repirain wih age in helamb wa due equal prprinal dereae in he Na+ K+)APaedependen and independen mpnen. he differene in repirain raebeween he even-weekld lamb and he adul ewe wa alm enirelyindependen f aK)-ae. The pei naure f he a ,K)ATPaeindependen funin ha differ are n knwn alhugh he auhrpeulaed ha diferene in proein ynhei are nly a minr mpnen26). In a reled ere f eperimen ana e al (17) berved ha rae fpalmiae idain in live and he lie fm pregnan and laaing rawere % and 20%, and 30% and 5% greaer han he bered in ample

frm agepaired nrl ra. I i lea ha a prin f hange in apparenmainenane requiremen due plane f nuriin (25), age 26) pregnany(0 7) and laain (1) an be aibued hange in baal energyependiure in iue. uher me f hee ae due hange in aiviyf he NaK)-ae niderable addiinal wrk will be required idenify aeed funin her han in ranpr, and relae hee quaniaively hanges n whle-animal energy ependiure. Fuher, i i eenial ha diery, endrine, and her phyilgi and meabli proeeha affe energy ependiure in iue are idenified and heir ee quan

ifed 57.

VARIATIO DUE TO DIFFERECES I RELATIVE ORGA WEIGHTS Prduive funin inluding grwh pregnany, and laain are ampaniedby ignifian inreae in fd inake. Weber 60) and her

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NUON NGCS O NMS 109

(6) hav rprtd that nrgy xpndtur n vra rgan uh a lvr andgut but 2530% f baa hat prdut n might xpt that rativwght f th rga wud inra at hgh fd inta. Smarly, thratv wight f vral rgan wth hgh nrgy xpnditur pr unt maa muh gatr i yug anma than n adut. Hday t al 28 addrdh pibiity that th high mtabi rat f ifat uld b xpaid hbai f th hgh ratv wght f bran and vra rgan n nfant ampard t adut Uing th aumptn that rgy xpndtur pr unitma f tu d nt dffr btwn nfant and adult, thy tmatd that vr50% f h dffr i hat prdut pr unit MBW in infat mpad taut ud b paid th bai f diffr i rativ rga wight.Bad th am aumptin, Smith & Badwn 52 and ana t a (17)

timatd that abut 50% f th nra n apparnt mantnan rqurmntin atating a mpard t natatng w and rat ud b th rut finra n ratv wght f vr gatonttna trat and har Kngand hi agu 34, 35 hav rprtd a r f prmnt wth rat,pg and hp dgnd t brng amal f ah pi t th am fawght at th am ag at dffng plan f nutrtn Fr xamp n th pgxprmnt, n grup HL wa fd to gan 19 g n th frt 35day fdngrid and t 5 g durng a nd 35-day prd a nd grup MMwa fd t gain 7 g durng ah prd a thrd grup LH wa fd t 5 g

th rt prd and t gain 19 g n th nd prd Fating hatprdutin/MBW FHP n th MM and H grup wr 20% and 40% gratrthan n th H grup, arly lutratng th magntud f th fft f pan fuiti baa mtabim Lvr wight in th MM and LH grup wr20% and 45% gratr than in th HL grup. Pitv and gnfiant ratn btwn FHP and wght f lvr gatrnttna trat, panra, anddny wr fund. brvatn wth rat and hp wr mparabl 35Pathway anay ndiat that a arg prprtn f varan in FHP an b

autd fr by hag in rativ wight f vital rgan ad pibyatd with hang in rprat pr unt wight f tu L. J. Kng,rna mmuniatn. If th ft f pan f nutrtin upn vital rganwight ar ntay nar and hghy rratd wth FHP, a uggtd byth rpt f Kng t al 34, 35 ad ana t a (6 ) rlutn fbrvd vaia n apparnt mantnan rqurmnt ud b traghtfrward. Uftunaty, many additinal fatr ntribut Efft f ag, phyigi, ad ndrn tat n nrgy pndtur pr unt wght f tuwr dud abv. Fard t a 19 rprtd that dt ha gnfant ff t

n th inra n livr wght that ampany atatn and/r th attndanthigh fd nta hi mpatbl with rprt f ntratn btwn ditand FHP 18, 25 Wbtr 6), Frl t al 20 and rmr t al 59hav rpd igniant x ana and brd fft upn FHP. Untla f th aua ratnhip that ad t varatn n appart mant-

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110 BADWN & BYWATER

nanc ha bn d u and quanfd, ur f cun fdn ym wllb rcd carful lcn f mannanc ma b ud fr hanmal hy wh fd

PRODUIVE PROESSES

Accura ma f anmal rfmanc bad un cndran f fdn, bdy wgh, nnmn, hylgcal a, and dc fncnrqur n nly accura ma f mannanc rqurmn, bu alaccura ma f ffcncy f cnrn f mablzabl nrgy rduc. h quan rnd ab NE = E and Kg = E ME,r h ba fr uch ma, wh h rrcn ha h w an f

fd na ( and ) and f mablzabl nry na (ME and ME)ud calcula and ME ar a r ab h amun rqurd frmannanc Chan rcrd nry (E) can b mad n wway. ncy dnn, ME = E + HE, n can maur HE undrc r ndrc calrmry (1, 15) and calcula E by dffrnc, r ncan maur drcly (3). hr ar adana and dadananhrn n ach arach. qumn fr bh drc and ndrc calrmry n; an nar ha a lmd numbr f un, h ma ar

uually h n duran and nl lmd numbr f anmal. Drcmaurmn f E n rwng anmal, fr aml, rqur ha E blrg rla rrr n marmn f na and fnal bdy ny cnnh arach can b nn and lar numbr f anmal can b d,u d mu b lng n duranw n ra and mnh n cal.Unfrunaly, rul band ng h w mhd nd dr. Cmrn h A 2 and N fdn ym fr rwn cal whch bad n ndrc calrmry and cmra laughr daa, rcly,llura h C ma f Km fr aru fd cd NC ma

by 015% whl ma f Kg ar 2530% rar. A rul f hdffrnc a rcmmndd nry rqurmn fr 200- and 00-cal rwn a h ra f day calculad accrdng h NC ymr 18% and % grar, rcly, han h calculad accrdng hAC ym (23) Sm h drnc can b land n h ba dffrnc n clma, lc manamn racc n h w cunr,nry l frm urfac SE), nnbl nrn l n accund fr nndrc calmry, and acy drncmmn mwha rcd n ndc calm chamb w, mch f h dncrman b land nc h quan ME E + HE s hrcally nd and abl n can cula ha h dffrnc l n dran fh w y f ud ladn amn f mwha drn hylc r abc rc h rc ncld ac f ur fmacrmlcl and hylgcal adaan chan n lan f nurn

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1 BLW BW

yeas o a esouo of poblems of vaaos maeace a poucveececes. e fs appoac s e use of eseac moes o evauae cuecoceps a aa fo quaave a yamc aequacy a o efy a

epe ccal epeimes s appoac a e uely pysoloca meaboc pocesses a e appopiaely cosee aayses of amaleey epeues a paro of uies wee evewe ecey 7, 853. eseac moels caacescay ae ly eoecal (causa acopoae coseable pysoloc a meabolc eal. As a esu ey aeoo compe a epesve o use as pecve moels o as compoes offee o maaeme sysems. Howeve ey ca be use o efy acaacere elemes fo corpoao o moe emprica pecve moeso use ue appe coos 7 14 4)

Te seco appoac as bee eveome of pecve moels of eeyasacos amas ese moels ae moe ly aeae a empical a esec moes bu e copoae pimary eema eemes oe ee a ey ae fa ess empcal a moels upo wc cuefee sysems ae base. Tese moels eealy cose e eso offee compoes a meabolsm of absobe uies eplcly ae a e aeae as eey a crue poe (9, 24 48; M ll pesoacommucao. ao ey oe cose e uove of macomolecules e amal a pao o uies amo ssues a fucos w e ama 8 1 4. ee elemes eable accommoao of aae poo of vaace appae maeace a poucve effcecesue o e composo composo of boy eey a o loss ae apysolocal sae of e amal. Above al ule cue sysems sucmoels ae yamc a eefoe may coa euaos o epese eeffecs of pae of uio ove me o basal ea pouco 5) y alae e plosopy a meooloy fo evelopme a vefcao oforous pecve moels e elavey ew a e ew moels avaable

eque fue eieme (9, 14). Howeve we eou s ow abouamal sysems a esposes pecve moes coa povsos foal eae vaiace ama ey epeues a ececes wbeome a eal pa of fee sysems mpov bo e accuacy aeea applcably of cue syses () Te avalably a eavelylow cos of mcocompues ae ely o aceleae eveopmes s aeaa o prove e ecessary vece fo paccal use of suc moels. We musbe cauous abou mpleme ese moes oweve e eecy s oaccep em ealy because suc sysems ae eeay moe sou o

acaemc ous oweve e capacy o euce eos of pecocompae o cue sysems soul be emosae befoe ey ae aope2) A mpoa feaue of e omeclaue 45 scusse above s a s compabe w e ouco of peco moes so as e laemaue a ae pove e ca be copoae ea.

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NUTRTON ENERETCS O NS 113

terture ited

Agrculural Research Coucil 1965.Th Nu Rqums of Fam

L�ok. Nos. 1& 2 ondon Ag. ResCoc2 Agculal Reseach Cocl. 1980

Th Nu Rqums of Rumavso pp. 4117 Fanham Royaluey: Gesham 2nd ed.

3 Asby P 1917 Nuo of FamAa New Yok Macmlla

4 Armsrong G, Blaxer K L. 1983n Po I Syp Hbvo Nu Subops ad ops ed Glchrst M Macke R Johaesburg A.. o pess

5 Baldw, R , mh N E 1974.Molecular corol o energy meabolsmn Mabl ed. pp 17-25 versy Par Pe.ae v Press

6 Baldwin, R , mh, N. E, aylor,J.harp M 1980 Manipulaig meabolcparameers o mprove gh rae andmlk seceo J Am S 5141628

7 Baldw R Bywaer A. C. aylorJ .• Muphy M 1983 negraon omeabolsm n he umen ad hos aimal. ee Re 4

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