P. W. MOE 1 US Department of Agriculture Bel t svi l l e, MD 20705 A B S T R A C T Ener gy met abol i s m research dur i ng t he past 25 yr has resol ved many uncer t ai nt i es of ener gy use by l act at i ng cows. Use of met abol i zabl e ener gy f or mi l k pr oduc t i on essent i al l y is unaf f ect ed by mi l k yi el d but is sl i ght l y i nf l uenced by its source. Est i mat es of ef f i ci ency of use f or mi l k pr oduct i on (60 t o 64%) are l ower t han earl i er est i mat es (69 t o 70%) pr i mar i l y because of l ower mai nt enance costs. Ef f i ci ency of met abol i zabl e energy f or body gain is hi gher in l act at i ng (75%) t han nonl act at i ng (60%) cows. Use of body t i ssue ener gy f or mi l k pr oduct i on is a bout 82% effi ci ent . End pr oduct s of di gest i on cont r i but e t o var i at i on in ef f i ci ency of f at t eni ng and in par t i t i on of energy bet ween mi l k and body gain in t he l act at i ng ani mal . Ener gy use in t he growi ng ani mal is i nf l uenced by compos i t i on of tissue gai ned and compos i t i on of t he di et . Energet i c ef f i ci ency of pr ot ei n depos i t i on is appar ent l y much l ower t han t hat of f at deposi t i on. A subst ant i al par t of t he l ower ef f i ci ency of pr ot ei n depos i t i on is r el at ed t o ener gy cost s of pr ot ei n t ur nover . I ncompl et e di gest i on o f mi xed di et s at high i nt ake by l act at i ng cows and me t hods t o pr edi ct ener gy par t i t i on are seri ous pr act i cal pr obl ems. In t he shor t t er m, i mpr oved met hods t o pr edi ct i nt ake effect s on met abol i zabl e ener gy of mi xed di et s will i ncrease accur acy of di et f or muI at i ons. In t he l onger t er m, met hods t o pr edi ct quant i t i es of nut r i ent s abs or bed f r om t he gut will per mi t a mor e f l exi bl e and accur at e met hod of eval uat i ng di et s and pr edi ct i ng ani mal per f or mance. Received June 26, 1980. 1 Ruminant Nutrition Laboratory, Animal Science Institute, Agricultural Research, Science and Education Administration, USDA, Beltsville, MD 20705. I N T R O D U C T I O N In 1955 Bt axt er and Gr aham ( 23) st at ed, " I n t he per i od since Kuhn and Kel l ner, t he con- sol i dat i on and ext ensi on of t he net - ener gy pri nci pl e, whi ch mi ght have been expect ed t o have occur r ed as a r esul t of such br i l l i ant work, has, wi t h several not abl e except i ons, not t aken place. Inst ead, t he per i od has been char act er i zed by pol emi cal ar gument r ar el y i l l umi nat ed by an e xpe r i me nt and har dl y ever by a cal or i met r i c trial. Kel l ner ' s ori gi nal wor k has been recal- cul at ed, re-expressed and, in shor t , sucked dr y. . . Cl earl y, in t he assessment of t he nut r i t i ve val ue of f oods, t he f ut ur e mus t i nvol ve ext ensi ve exper i ment at i on and meas ur ement r at her t han t he al most compl et e dependence on pi oneer evi dence whi ch has char act er i zed t he past 50 year s. " Progress has been consi der abl e in t he fi el d of energy met abol i s m si nce t hose wor ds were wri t t en Ef f i ci ency of ener gy use and ener gy r equi r ement s have been i dent i f i ed mor e pre- cisely. Progressi vel y mor e i nt ensi ve experi - me nt a t i on has descr i bed physi ol ogi cal and bi ochemi cal bases f or an i ncreasi ng par t of t he var i at i on in ener gy use. Al t hough energy met abol i s m has been st udi ed at many levels f r om specific bi ochemi cal t r ans f or mat i ons t o whol e popul at i ons of ani mal s, t hi s revi ew will deaI pr i mar i l y wi t h ener gy use in t he whol e ani mal . Aspect s of energy met abol i s m t hat rel at e t o t he effect i veness wi t h whi ch dai r y cat t l e consume a var i et y of di et s f or gr owt h, r epr oduct i on, and pr oduc t i on of mi l k will be consi der ed. Energy Ter mi nol ogy In t he di scussi on, t er mi nol ogy is t hat in general use. Di gest i bl e ener gy (DE) is gross i nt ake of energy mi nus ener gy voi ded in t he feces. Met abol i zabl e ener gy (ME) is DE mi nus ener gy in met hane and urine. Generally ME is an expr essi on of t he amount of ener gy avai l abl e for met abol i s m by t he ani mal , al t hough ME i ncl udes some energy, e.g., heat of f er ment at i on, 1981 J Dairy Sci 64:1120-1139 1120 METABOLISM -- 75TH ANNIVERSARY ISSUE 1121 not avai l abl e f or met abol i s m and does not i ncl ude some energy, e.g., ur i ne energy, whi ch is a pr oduc t of met abol i sm. The t er ms f or t he par t i al ef f i ci ency of ME used f or mai nt enance, l act at i on, pr ot ei n gain, f at gain, and gain in t ot al tissue ener gy are ki n, kl, kp, kf, and kg. Heat i ncr ement (HI) is t he i ncrease in t ot al pr o- duct i on of heat associ at ed wi t h an i ncrease in t he cons umpt i on of f ood. Energy uni t s are cal ori es (cal), ki l ocal or i es (kcal = 1000 cal), or megacal or i es ( Mcal = 1000 kcal ) f or t he conveni ence of t he US r eader al t hough many of t he ori gi nal paper s i ncl uded t he j oul e (1 cal = 4. 184 J; J = M2 . k g . s - 2 ) . The Situation 25 Years Ago Revi ews by Bl axt er (14, 15) and Rei d (77, 78) and maj or t ext s by Bl axt er (17) and Kl ei ber (47) summar i zed much of t he wor k on ener gy met abol i s m of dai r y cat t l e and pr esent t he mos t compr ehensi ve descr i pt i ons of t he ener get i cs of dai r y cat t l e avai l abl e in t hei r t i me. Earl i er st udi es showed t hat l act at i on was mor e ef f i ci ent t han f at t eni ng. Rei d (78) s ummar i zed t he avai l abl e cal or i met r y dat a and concl uded t hat ME cons umed in excess of mai nt enance was used t o t he e xt e nt of 69. 3% f or mi l k pr oduc t i on and 58.0% f or body i ncrease and t hat bot h were r el at i vel y const ant . Gr owt h was t hought t o be mor e ef f i ci ent t han f at t eni ng. Ener gy f r om grai ns or concent r at e feeds gener al l y was accept ed t o he used mor e ef f i ci ent l y t han ener gy of forages, especi al l y f or gr owi ng animals. Thi s di f f er ence appear ed t o be r el at ed t o t he cr ude f i ber c ont e nt of t he forage. Consi der abl e di sagr eement exi st ed r egar di ng t he r el at i onshi p bet ween ener gy i nt ake and ener gy bal ance (EB). Wor ker s gener al l y accept ed t hat t hi s r el at i onshi p was cur vi l i near when dat a bot h above and bel ow mai nt enance were i ncl uded. Bl axt er (15) emphasi zed t he curvi l i nar r el at i onshi p bet ween f ood cons umed and ener gy r et ent i on and i ndi cat ed t hat t hi s ef f ect was at l east in par t f r om a decl i ne in ME val ue wi t h i ncr eased i nt ake. Al t hough wor k had doc ume nt e d r educed di gest i bi l i t y by l act at i ng cows in compar i s on wi t h t hat of cows at mai nt enance i nt ake, t hi s ef f ect was not accept ed uni versal l y. Rei d (78) emphasi zed t he i mpor t ance of r educed di gest i bi l i t y and i ncr eased body f at t eni ng by l act at i ng cows at high f eed i nt akes in ex- pl ai ni ng appar ent di mi ni shi ng r et ur ns in f eedi ng t ri al s wi t h l act at i ng cows. Consi der abl e uncer t ai nt y r emai ned wi t h regard t o t he ef f ect of di et qual i t y or r at e of mi l k pr oduct i on or r at e of gr cwt h on ef f i ci ency of use of ME and whet her di et ef f ect s were si mi l ar for f at t eni ng and mi l k pr oduct i on. These quest i ons were of f undament al im- por t ance in pr ovi di ng accur at e and useful f eedi ng st andar ds. The si t uat i on in 1956 can be unde r s t ood mos t adequat el y by r emember i ng t hat t he l i mi t ed cal or i met r i c dat a on l act at i ng cows at t he t i me had been obt ai ned many year s pr evi ousl y. Those resul t s, al t hough i mpor t a nt and al t hough obt ai ned wi t h preci se met hods , were i nadequat e t o answer i mpor t a nt quest i ons raised a bout f act or s af f ect i ng ef f i ci ency of ener gy use by cat t l e. Those resul t s were re- cal cul at ed and debat ed in t he l i ght of newer knowl edge. New i nt er pr et at i ons were put f or war d wi t hout benef i t of faci l i t i es t o t est t hei r val i di t y. New exper i ment at i on was needed. The Last 25 Years The 25 yr since 1956 coi nci des al most exact l y wi t h an except i onal bur st of exper i - me nt a t i on in ani mal energet i cs. In t he l at e 1950' s, maj or c ommi t me nt s t o s uppor t ener gy met abol i s m research were made in a number of count r i es i ncl udi ng t he US. The i ncrease in r at e of e xpe r i me nt a t i on wi t h l act at i ng cows can be appr eci at ed f r om t he f act t hat a t ot al of 110 compl et e ener gy bal ance t ri al s wi t h 38 cows, i ncl udi ng all r epl i cat es, had been c ompl e t e d in all of t he l abor at or i es in t he wor l d bef or e 1961 (34). Si nce t hat t i me t he resul t s of 806 bal ance trials wi t h l act at i ng cows have been publ i shed f r om t he Bel t svi l l e l a bor a t or y al one. To co- or di nat e t he i ncr eased research act i vi t y and t o share research pl ans and resul t s, an I nt er nat i onal Sympos i um on Ener gy Met abol i s m was hel d in Copenhagen in 1958 under t he sponsor shi p of t he Eur opean Associ at i on of Ani mal Pro- duct i on ( EAAP) . Succeedi ng symposi a, hel d every 3 yr, have pr ovi ded a cont i nui ng f ocus f or ener gy met abol i s m research. The pr oceedi ngs of t hese symposi a, publ i shed by t he EAAP, doc ume nt a subst ant i al par t of t he ener gy met abol i s m research wi t h f ar m ani mal s dur i ng t he l ast 25 yr ( Tabl e 1). The s ympos i a in 1958 and 1961 deal t l argel y Journal of Dairy Science Vol. 64, No. 6, 1981 1122 MOE TABLE 1. Symposia on energy metabolism sponsored by the European Association of Animal Production (EAAP). 1st 2nd 3rd 4th 5th 6th 7th 8th Symposium on Energy Metabolism. Principles, Methods, and General Aspects. Copenhagen, Denmark. 15-19 September 1958. Publ. by EAAP (Publ. No. 8) and Statens Husdyrugsudvalg, Copenhagen. Grete Thorbek and H. Aersoe, ed. Symposium on Energy Metabolism. Methods and Results of Experiments with Animals. Wageningen, the Netherlands. 11-15 September 1961. EAAP Publ. No. 10. E. Brouwer and A.J.H. van Es, ed. Symposium. Troon, Scotland. May 1964. Energy Metabolism. Publ. by Academic Press, New York and London. 1965. K. L. Blaxter, ed. EAAP Publ. No. 11. Symposium. Energy Metabolism of Farm Animals. Warsaw, Poland. Sep- tember 1967. Publ. by Oriel Press, Newcastle upon Tyne, England. 1969. K. L. Blaxter, J. Kielanowski and Grete Thorbek, ed. EAAP Publ. No. 12. Symposium. Vitznau, Switzerland. September 1970. Energy Metabolism of Farm Animals. Juris Verlag, Zurich. A. Schurch and C. Wenk., ed. EAAP Publ. No. 13. Symposium. Hohenheim, B.D.R. September 1973. Energy Metabolism of Farm Animals. Publ. by Universitat Hohenheim Dokumentationsstelle. 1974. K. H. Menke, H. J. Lantzsch, and J. R. Reichl, ed. EAAP Publ. No. 14. Symposium. Vichy, France. September 1976. Energy Metabolism of Farm Animals. Publ. by G. de Bussac, Clermont-Ferrand. M. Vermorel, ed. EAAP Publ. No. 19. Symposium. Cambridge, England. September 1979. Energy Metabolism. Publ. by Butter worths, London. 1980. EAAP Publ. No. 26. L. E. Mount, ed. with met hodol ogy: construction of respiration chambers, techniques of gas analysis, and potential errors in energy balance measurements. Succeeding symposia dealt increasingly with presentation of results of animal experiments, discussion of interpretations of energy balance measurements, and proposals for application of findings in practice including discussions on feed evaluation and feeding standards. More recent symposia have dealt progressively less with feed evaluation and increasingly with specific factors limiting or causing variation in energy use by farm animals including more intensive and physiological experimental ap- proaches and newer methods of describing and interpreting data. Major advances during this time have been in identifying undet ect ed sources of variation in energy use, in developing quantitative des- criptions of known sources of variation, and in developing recommendat i ons for means of implementing knowledge of energetics in practical feeding systems. Effects of A bs o r bed Nutr i ents Ener get i c ~f f i c i e nc y . A major i mprovement in understanding the causes of variation in energy efficiency of animals fed different diets was the demonstration of Armstrong, Blaxter, and their coworkers (2, 3, 4, 6, 7) that the heat increment of mixtures of steam volatile fatty acids (VFA) was influenced greatly by the proport i on of acet at e in the fattening sheep but had less effect in sheep at maintenance. The energy of VFA infused singly into the rumen of fattening sheep was used with efficiencies of 32.9% for acetic acid, 56.3% for propionic acid, and 61.9% for butyric acid. Mixtures of VFA containing 75 and 25% acetic acid were used with 31.8 and 58.1% efficiency. Other studies showed a lower efficiency (54.5%) for glucose infused into the rumen than for that infused into the abomasum (71.5%) or jugular vein (72.8%). These studies indicated the importance of end products of digestion as opposed to nutrients consumed in influencing metabolic efficiency in ruminants. Journal of Dairy Science Vol. 64, No. 6, 1981 METABOLISM -- 75TH ANNIVERSARY ISSUE 1123 These studies also suggested that variation in the efficiency with which specific end products of digestion are metabolized could account for a substantial part of the apparent difference between use of fibrous diets and that of diets containing large amounts of starch. The ap- parently low efficiency of acetate use in the fattening animal appeared to account for the depression in net energy of feeds high in fiber that had caused Kellner to introduce his "fiber correction factor" some 50 yr earlier. More recent experiments indicated that the proportion of acetate of the rumen fermentation products may not explain fully the variation in use of ME and that in some circumstances acetate may be used efficiently for body gain. Tyrrell et al. (92) reviewed several experiments in which acetate apparently was used with relatively high efficiency for body gain and reported the results of calorimetric investi- gations with VFA infusion in mature cows. They found a difference from the nature of the basal diet in the partial efficiency of acetate for body gain. Use of ME from infused acetic acid was 27% for cows fed 100% hay and 69% for cows fed a diet of 30% hay. Orskov et al. (71) reported experiments in which lambs were sustained entirely by in- tragastric infusion of VFA, protein, minerals, and vitamins. For mixtures of VFA with 450 to 750 mmol acetate/mol, efficiency of use was 57 to 64%. They also recalculated the earlier data of Armstrong and Blaxter and reported that those results predicted an efficiency of 44 to 50%. Orskov et al. (71) concluded that the effect of proportion of acetate on efficiency of energy use in growing animals was, in both instances, too small to be of practical signifi- cance. The differences in the results of Tyrrell et al. (92) and Orskov et al. (71) remain unresolved. A partial explanation may be found in the differing physiological state of the experimental animals. Data of Orskov et al. (71) pertain to growing lambs depositing substantial protein whereas data of Tyrrell et al. (92) pertain to fattening in adult cows. Under some conditions acetate is used with low efficiency for fattening although some questions remain as to exactly what those conditions are. Also, acetate can be used efficiently in many instances, especially with high concentrate diets. Partitioning of Energy. In only a relatively few experiments has the use of individual VFA in lactation been investigated. Armstrong and Blaxter (5) in calorimetric studies infused mixtures of VFA, propionate, and acetate into the rumen of goats and found efficiencies of 71.4, 72.3, and 65.0% for lactation and 50.3, 52.3, and 44.4% for energy retention in the nonlactating body. Two particularly i mport ant findings were described in this paper. First, acetic acid infusion resulted in an increase in milk fat secretion and a decline in body fat deposition whereas with propionic acid infusion the reverse was true. Second, heat production was not changed. This second finding led them to the conclusion that energy retention in the adult rumi nant is more efficient accompanied by the simultaneous process of milk secretion than in the nonlactating animal. The effect of VFA infusion on energy partition was not ed with lactating cows by Orskov et al. (70) in calorimetry experiments. Acetic acid infusion resulted in more milk energy and less gain of body energy than did propionic acid infusion. No difference in efficiency was found. Effects on energy partition in lactation have been similar with changes in diet. Table 2 shows results of an experiment by Flatt et al. (36, 37) in which cows fed a 60: 40 ratio of forage to concentrate produced more milk and lost more body tissue energy than cows fed a 20:80 ratio. The ratio of acetate to propionate in rumen VFA was also higher on the higher forage diet. In incremental studies of corn grain and beet pulp, Tyrrell et al. (89) found a greater per- centage of the increase in energy balance (milk plus body tissue) was milk when beet pulp was added to the diet than when corn was added. Sutton et al. (82) observed a reduction in proportion of acetate in rumen VFA and a reduction in milk fat yield by lactating cows when the percentage of concentrate was in- creased from 60 to 90%. At 90% concentrate, more starch reached the duodenum when corn grain was fed than when barley was fed. Live weight gain was also greater on the 90% corn diet, and milk yield was less. When corn replaced barley, the cont ri but i on of rumen digestion to overall digestion of energy was reduced considerably. Effects of percentage concentrate, percentage of crude protein, and feed intake on partition of energy in lactating cows were studied by Journet et al. (44). The partition of energy into Journal of Dairy Science Vol. 64, No. 6, 1981 1124 MOE TABLE 2. Influence of hay:grain ratio on partition of energy between milk and body tissue a. Hay:grain ratio Item 60:40 40:60 20:80 Metabolizable energy (ME) intake, Mcal 36.12 36.42 34.87 Energy balance, McaI 11.94 12.63 12.16 Milk energy, Mcal 13.94 13.17 10.41 Tissue energy, Mcal - 2.00 - . 54 1.75 Milk fat, % 3.5 3.0 2.7 Acetic:propionic 3.32 2.57 2.00 aData from Flatt et al. (36, 37). milk decreased with increasing percentage of concentrate and decreasing percentage of milk fat. They also found that when energy intake varied around the requirement, about one-third of the increment of energy went into milk and two-thirds into body gain. No relationship between major VFA and milk product i on was found, but mi nor VFA (isobutyric, isovaleric, and valeric) in the rumen and crude protein in the diet were related positively to milk pro- duction. Variation in the partial efficiency of use of the energy of VFA is considerably more im- port ant in fattening than in lactation whereas the effect in the growing animal is less certain. In the lactating cow, however, amount s of individual VFA absorbed from the gut can exert a significant effect on partition of energy between milk and body tissue. Effects of variation in amount and type of diet on energy efficiency and energy partition likely will not be explained satisfactorily without compre- hensive knowledge of amount s of specific nutrients which are absorbed from the digestive tract. I ntake and Associative Effects Intake Effects. Although intake effects on digestibility had been shown early in this century (29, 41), the practical significance of this effect by no means was accepted uni- versally. The question was debated at a sym- posium in 1965 (26, 34, 76, 79). Brown (26) cited several instances in which intake effects on digestibility appeared to be conflicting. He concluded that although reduced digestibility had been detected at high intakes of diets containing large amount s of concentrates in a number of experiments, additional i nformat i on was needed. Flatt (34) reviewed the information then available on intake effects on ME value of diets and concluded that such an effect could not be documented. In a study of Beltsville data, Moe et al. (54) also found no intake effect on ME values of diets containing 40, 60, and 80% concentrate for lactating cows. Not until the 1970' s did experimental results begin to appear to indicate in a relatively systematic fashion that ME values of diets for lactating cows were substantially lower than the same diets fed to nonlactating cows at a mai nt enance intake. The significance of depression in digestibility of diets at high intakes by lactating cows was established firmly in extensive digestibility measurements by Moe et al. (56), Wagner and Loosli (103), and Ekern (31). Tyrrell and Moe (86) reviewed these and other studies and concluded that digestibility of normal diets by dairy cows was reduced by about 4% for each increase in intake equivalent to the amount needed for maintenance. They also concluded that the rate of reduction in digestibility was greater for diets containing larger percentages of concentrate although this effect was less pronounced for diets based on corn silage. Intake effects are as great with corn silage-based diets as with other forages when high per- centages of grains are fed. At lower percentages of grain, however, intake effects are likely greater with corn silage diets than for those containing other forages. In a more recent review, Tyrrell (85) cites evidence that the digestibility of corn silage- based diets is improved by addition of ground Journal of Dairy Science Vol. 64, No. 6, 1981 METABOLISM -- 75TH ANNIVERSARY ISSUE 1125 l i mes t one t o t he di et (109) or by i ncr eased pr ot ei n c ont e nt (74) and in al f al f a- based di et s by i ncr eased pr ot ei n c ont e nt (88). Tyr r el l and Moe (87) r e por t e d a r educt i on in di gest i bi l i t y of cor n si l age-based di et s suppl e- ment ed wi t h ei t her cor n or bar l ey grains f or l act at i ng cows. The ME val ue of t he bar l ey di et was not i nf l uenced by i nt ake wher eas ME val ue of cor n di et decr eased at hi gher i nt akes. The decl i ne in di gest i bi l i t y was si mi l ar f or t he t wo di et s. The decl i ne in di gest i bi l i t y of cel l ul ose and hemi cel l ul ose f r act i ons was a bout 8 per- cent age uni t s and t hat of st ar ch a bout 3 per- cent age uni t s per uni t of mai nt enance i ncrease in i nt ake. Wheel er and Nol l er (110) r epor t ed t hat a par t of t he r educed di gest i bi l i t y of ener gy in l act at i ng cows consumi ng cor n grai n and cor n silage in large amount s was pr event ed by s uppl ement at i on of t he di et wi t h 2.7% l i me- st one. Suppl ement at i on wi t h l i mest one in- creased fecal pH, r educed st arch losses in t he feces, and i mpr oved f eed ef f i ci ency. They suggest ed t hat t he i ncr eased fecal pH r ef l ect ed a mor e f avor abl e i nt est i nal pH f or act i vi t y of pancr eat i c al pha amyl ase. Poos et al. (75) s uppl ement ed di et s of l act at i ng cows wi t h ei t her ur ea or s oybean oil meal (SBOM). Urea was ef f ect i ve in i mpr ovi ng di gest i bi l i t y of di et s cont ai ni ng 11.6 t o 13.6% crude pr ot ei n when added in amount s t o i ncrease cr ude pr ot ei n t o 13 t o 14.2%, but onl y SBOM was ef f ect i ve at hi gher per cent s. These resul t s suggest t hat ur ea is ef f ect i ve in i mpr ovi ng di gest i bi l i t y at hi gher cr ude pro- t ei n in l act at i ng cows t han had been shown pr evi ousl y f or nonpr oduci ng ani mal s. I mpr oved di gest i on of cor n silage di et s by l act at i ng cows wi t h s uppl ement al ur ea also has been shown by Ver i t e (101). In exper i ment s wi t h l act at i ng cows, Tyr r el l and Moe (88) f ound a gr eat er ef f ect of i nt ake on ME val ue at l ower pr ot ei n t han at hi gher. When di et s of al f al f a hay, corn, and SBOM were i ncreased f r om 14 t o 17% cr ude pr ot ei n by s ubs t i t ut i on of cor n wi t h SBOM, bot h DE and ME were i ncr eased at hi gh i nt akes. When pr ot ei n was i ncreased t o 20%, however, onl y DE was i ncreased. Wi t h cor n silage di et s, DE and ME were i ncr eased when cr ude pr ot ei n was i ncr eased f r om 11 t o 14% but not when cr ude pr ot ei n was i ncr eased f ur t her t o 17%. Addi t i on of 2.5% l i mest one had no ef f ect on di gest i bi l i t y of ener gy in t he l at t er exper i ment . Associative Lffects. Lact at i ng cows usual l y are fed mi xed di et s r at her t han ei t her all f or age or all concent r at e. The di gest i on of mi xed di et s at high i nt akes c ommonl y is c ompa r e d wi t h di gest i on of t he same di et at a mai nt enance i nt ake t o measur e " i nt ake ef f ect s". "Associ at i ve ef f ect " refers t o di gest i bi l i t y of a mi xed di et di f f er ent f r om t hat pr edi ct ed f r om di r ect meas ur ement of t he forage and concent r at e separ at el y. I nt ake effect s and associ at i ve ef f ect s are basi cal l y t he same t hi ng, t he i ncompl et e di gest i on of a mi xed di et at a high i nt ake. Thi s is i mpl i ci t in t he est abl i shed r el at i onshi p bet ween per cent age of concent r at e and i nt ake effect s. The i mpor t ance of associ at i ve ef f ect s f or l act at i ng cows concer ns addi t i vi t y of t he di gest i bi l i t y of c ompone nt feeds in a mi xed r at i on. In f eed eval uat i on st udi es, t he di gest - i bi l i t y of concent r at es t ypi cal l y is measur ed by di f f er ence wi t h nonpr oduci ng ani mal s, usual l y at a mai nt enance i nt ake. Bl axt er (20) sum- mar i zed ext ensi ve st udi es of ME of di et s cont ai ni ng 0 t o 60% concent r at e wi t h sheep at mai nt enance and concl uded t hat associ at ed ef f ect s were not a pr act i cal pr obl em. At hi gher f eed i nt akes, however, associ at i ve effect s can be real. An exampl e is descr i bed by J oanni ng et al. (43), who f ound di gest i bi l i t y of a mi xt ur e of cor n grai n and cor n silage was 11% less t han t hat pr edi ct ed f r om di gest i bi l i t y of t he com- pone nt feeds measur ed at high i nt akes. I nt ake ef f ect s on di gest i bi l i t y were obser ved on mi xed di et s, but not when ei t her cor n silage or cor n grai n was fed al one. These dat a are i l l ust r at ed in Fi gur e 1. I nt ake ef f ect s have been expr essed by Van Soest and cowor ker s (100) by descr i bi ng " di s c ount f act or s ", whi ch r epr esent t he re- duct i on in di gest i bi l i t y of single f eedst uf f s when i nt ake is i ncr eased by an a mount equal t o mai nt enance. They l i st ed di scount s f or a l arge number of feeds f r om in vivo dat a and in vi t r o di gest i on rat es and avai l abl e passage rates. They concl uded t hat feeds of hi gher cell wall c ont e nt and of a l ow degree of l i gni f i cat i on t end t o have t he l ar gest di scount s. They also st at ed t hat st arch adds t o t he cell wall ef f ect in cereal grains. The use of di s count f act or s for i ndi vi dual feeds in comput i ng t he act ual di gest i bi l i t y of mi xed di et s i ncor por at es t he s us cept i bi l i t y of c ompone nt feeds t o i nt ake effect s. Di s count f act or s pr es ent ed by Van Soest et al. (100), Journal of Dairy Science Vol. 64, No. 6, 1981 1126 MOE R E L A T I O N S H I P B E T W E E N I N T A K E E F F E C T A N D A S S O C I A T I V E E F F E C T 85 DRY MATTER DIGESTIBILITY % 65 ; M A I N T E N A N C E i N T E A N K A ~ ~ ~ ~"""'~I-~/INTAKE1 IW/ , ~. " EFFECTL~ ~ / ASSOCIATIVE A D LJBITUM EFFECT I N T A K E 100~% CORN SI LAGE~ 0 O ~ % CORN GRAIN ~ 100 Figure 1. Data of Joanning et al. (43) show an associative effect between corn grain and corn silage at ad libitum intake but not at lower intake. however, do not solve the problem of non- additivity or associative effects. A useful extension of the discount concept will be a procedure for estimating discounts for total mixed rations so that the interactions between fiber degradation and use of soluble carbo- hydrates can be incorporated. required to produce that product as in Figure 2. With several physiological processes occurring simultaneous/y, however, estimates of partial efficiency for each process are clouded by experimental error and also by necessary assumptions. The most troublesome assumption concerns maintenance. Three alternatives for estimating energetic efficiency are in Figure 3. The estimate may be derived from two or more EB measurements made above maintenance in which case the estimate is by difference or regression. The precision of the estimate is determined by experimental error and by the magnitude of the difference in energy intake and production. Alternative met hods involve an assumed amount of either ME or net energy (NE) required for maintenance combined with a single estimate of EB at some point above maintenance. These alternative methods yield lower errors because no error is associated with the maintenance estimate. An incorrect as- sumption regarding maintenance, however, will introduce a bias. Considerable variation in published estimates of partial efficiency is from differences in assumptions regarding main- tenance. Partial Ef f i c i e nc y Es t i ma t i on Efficiency is the ratio between energy in the product formed and the amount of energy E N E R G Y B A L A N C E ~AEB A M E M E I N T A K E ENERGY DIRECT MEASURE BALANCEoI / OFEFFICIENCY ENERGY BALANC% . . . . M E I N T A K E ~ A S S U M E ME REQUIREMENT FOR MAI NT E NANCE M E I N T A K E METABOLIC EFFICIENCY - 3EB A M E - k A M I L K k ; - A M E k -- A GAIN g A M E Figure 2. Partial efficiency of metabolizable energy (ME) for production. . . . . . . ASSUME NE REQUIREMENT FOR MAI NT E NANCE ME I NTAKE Figure 3. Partial efficiency may be estimated by regression (top) or by assumed metabolizable energy (ME) or, net energy (NE) required for maintenance. Journal of Dairy Science Vol. 64, No. 6, 1981 METABOLISM -- 75TH ANNIVERSARY ISSUE 1127 The pr obl ems associ at ed wi t h mat hemat i cal est i mat es of par t i al ef f i ci ency, i ncl udi ng t he concept s of mai nt enance, were di scussed ext ensi vel y by Van Es (95). He emphasi zed l i mi t at i ons of t he concept of net ener gy for mai nt enance and ef f i ci ency of energy use f or mai nt enance. Energy r equi r ed for mai n- t enance is pr i mar i l y for t he pr oduc t i on of ATP, and t hi s ener gy al ong wi t h t he wast ed ener gy is l ost event ual l y as heat . Heat pr oduc t i on at mai nt enance is, t her ef or e, t he t ot al of pr o- duct i ve and nonpr oduct i ve energy. Heat pr o- duced by t he fast i ng ani mal is t he r esul t of body tissue bei ng met abol i zed t o suppl y t he ener gy needed t o mai nt ai n t he ani mal . The r at i o bet ween fast i ng and mai nt enance heat pr oduct i on is, t her ef or e, an expr essi on of t he rel at i ve ef f i ci ency of body tissue ener gy and di et ar y ener gy in meet i ng t he needs for mai n- t enance. Effi ci enci es cal cul at ed in such a manner shoul d be r ef er r ed t o as " a ppa r e nt ef f i ci ency" and bear l i t t l e r el at i onshi p t o t he ef f i ci ency c omput e d f or ener gy use above mai nt enance in whi ch a measur eabl e end pr oduct is f or med. Van Es (95) also di scussed errors associ at ed with es t i mat i on of mai nt enance r equi r ement by regression. He not ed t hat regressi on anal yses t heor et i cal l y r equi r e t hat t he i nde pe nde nt vari abl e(s) be measur ed wi t hout error. If not , t he c omput e d regressi on coef f i ci ent s will be under es t i mat ed. Si nce ME and EB bot h are measur ed wi t h error, i ncl udi ng some sources of er r or such as fecal, met hane, and ur i ne losses t hat are c ommon t o bot h, t he model sel ect ed will i nf l uence est i mat es of par t i al ef f i ci ency and t he est i mat e of mai nt enance. Cr amer (28) descr i bed an or t hogonal re- gressi on pr ocedur e t hat accommodat es er r or s in bot h vari abl es. It mi ni mi zes t he per pendi cul ar di st ances f r om t he l i ne t o t he poi nt s r at her t han t he vert i cal di st ances f r om t he l i ne t o t he poi nt s as in convent i onal regression. Because of t he sources of er r or c ommon t o EB and ME, a syst em has been pr opos ed in whi ch r et ai ned ener gy ( R) is r el at ed t o gross ener gy i nt ake (G) by scaling bot h wi t h fast i ng heat pr oduct i on in a gener al i zed f or m of t he Mi t scherl i ch equat i on R = B ( 1 - e x p ( p G) ) - l , where t he par amet er s B and p are f i t t ed by an i t er at i ve pr ocedur e (19, 21, 22). Thi s t echni que avoi ds assumpt i ons regardi ng mai nt enance r equi r ement s but requi res di r ect meas ur ement of fast i ng heat pr oduct i on by each t est ani mal or a t abul at i on of fast i ng heat pr oduc t i on accor di ng t o breed, size, sex, and ot her at t r i - but es and of t he ener gy r et ent i ons associ at ed wi t h gains in body wei ght , Wi t h t he i ncr eased at t ent i on t o t he des- cr i pt i on of t he mai nt enance c ompone nt in growi ng ani mal s, Webst er et al. ( 1 0 6 ) c o mp u t e d a " bas al " c ompone nt of met abol i s m r el at ed t o live wei ght f r om EB meas ur ement wi t h gr owi ng steers and concl uded t hat t he basal c ompone nt was cl osel y r el at ed t o body wei ght t o t he e xpone nt .734. Thi s est i mat e was, however, obt ai ned by cal cul at i ng t he "basal c o mp o n e n t " f r om assumed r el at i onshi ps bet ween Q (% ME in t he di et ) and km and kf (1) whi ch i nvol ve mai nt enance of mat ur e ani mal s and ef f i ci ency of f at t eni ng. Because act ual meas ur ement of fast i ng met abol i s m in t hese ani mal s decl i ned wi t h i ncreasi ng wei ght when expr essed as kcal / kg "73 in accor d wi t h ARC (1), t he aut hor s concl uded t hat t hese fi ndi ngs cast seri ous doubt on t he val i di t y of use of measur ed fasting met abol i s m as a basel i ne f r om whi ch t o pr edi ct ef f i ci ency of gr owt h. The uncer t ai nt i es associ at ed wi t h st at i st i cal par t i t i oni ng of ener gy cost in t he pr oduci ng ani mal i nt o mai nt enance and pr oduc t i on are consi der abl e. Al t hough newer mat hemat i cal t echni ques event ual l y may per mi t a ba ndonme nt of t he concept of mai nt enance, especi al l y in t he growi ng ani mal , i t is a useful and necessary c ompone nt in di scussi ng ener gy use by ani mal s of di f f er i ng pr oduct i on rates. In t he di scussi ons t hat f ol l ow, t h e r eader is r emi nded t hat as- s umpt i ons regardi ng mai nt enance exer t con- si der abl e ef f ect on est i mat es of pr oduc t i on ef f i ci ency. Mai nt enance and Lact at i on Cal or i met r i c exper i ment s by Br ouwer et al. (25) and Van Es (94) showed t hat t he val ue of a series of hays for mai nt enance of cows was mor e cl osel y r el at ed t o t hei r ME cont ent t han t o st ar ch equi val ent . Bl axt er (16) s ummar i zed resul t s of cal or i met r i c st udi es wi t h di et s rangi ng from poor qual i t y forage t o all concent r at e and concl uded t hat ef f i ci ency of use of ME for mai nt enance was f ai r l y const ant . These resul t s s uppor t ed t he concl usi on of Ri t zman and Benedi ct (80) f r om earl i er cal or i met r i c st udi es. Bl axt er (16) concl uded f ur t her t hat t he el - Journal of Dairy Science Vol. 64, No. 6, 1981 1128 MOE ficiency for mai nt enance of ME from these natural feeds was equal to that of VFA infused into the rumen when a correction was made for the heat of ferment at i on of the natural feeds. These studies showed that the variation in use of ME for mai nt enance was low, but some variation did exist, presumably in part from losses in heat of fermentation, which was not measured in the determination of ME. Agri- cultural Research Council (1) described variation in efficiency of use of energy for mai nt enance (k m) as a funct i on of percentage of ME in the diet (Qm) as follows: k m 54.6 + .30 Qm- A major question, unresolved in earlier calorimetric investigations, was the ext ent to which efficiency of ME for milk production was influenced by the source of dietary ME. Initial lactation studies at Beltsville (27) were with diets of alfalfa hay and concentrates in ratios calculated to provide 50, 75, and 100% of estimated net energy (ENE) from the alfalfa. When maintenance was assumed to be 131 kcal ME/kg -Ts , efficiencies of use of ME in excess of maintenance were 65, 61, and 54%, respectively, for lactation. Hashizume et al. (42) found the efficiency of ME of diets containing 45 and 71% of concentrate consumed in excess of mai nt enance (116.3 kcal/kg" vs ) was used with efficiencies of 74.0 and 68.2%, respectively, for milk plus retained body tissue energy. Van Es and Nijkamp (98) reported the results of 41 balance trials with lactating cows consuming mixed diets of concentrate, silage, and variable amounts of hay. In these studies, no effects of percentage of crude fiber or of crude protein on efficiency of milk production were detected. Also, no differences in utilization of hay and silage of equal ME and protein cont ent were found. These workers concluded that ME was used for milk production with an efficiency (kl) of 54 to 58% and that 10.1 to 11.7 Mcal ME was required for the mai nt enance of a 500 kg cow (96 to 111 kcal ME/kg'VS). Van Es and Nijkamp (98) discussed problems associated with mathematical descriptions of results of experiments with lactating cows in positive or negative body tissue EB. Relation- ships between ME intake and milk energy were studied by multiple regression with separate terms for body weight, milk energy, tissue energy gain, and tissue energy loss and by applying several methods of adjusting to zero body tissue EB. The various methods yielded slightly different estimates of efficiency but all methods indicated a slight increase in efficiency of ME use for milk production with an increase in metabolizability of the diet. An extensive series of EB experiments by Flatt et al. (37) with Holstein cows producing up to 49 kg of 4% fat-corrected milk (FCM) per day and consuming diets of 40, 60, and 80% concentrate plus alfalfa hay showed no signifi- cant differences among diets in efficiency of use of ME for milk production plus tissue energy gain. These studies included cows at all stages of lactation and when nonlactating and at both ad libitum and restricted feeding. The magnitude of changes in body tissue energy status was far greater than in previous experi- ments and varied from - 20. 6 to +18.3 Mcal body tissue energy per day. Differences due to diet were in body tissue balance; cows on the highest concentrate diet mobilized less fat in early lactation and deposited more fat in late lactation than cows on the highest forage diets at equal ME intake. The regression of total EB of milk plus body tissue energy on ME intake was EB (kcal/kg "7s) = - 93.4 + .66 i .011 ME (kcal/kg'VS). This equation indicated 66% utilization of ME and zero EB at 142 kcal ME/kg "vs. Extensive mathematical analyses of these data led to several conclusions: 1) use of ME for milk or body tissue gain was relatively unaffected by milk yield, amount of body tissue gain (or loss), and stage of lactation; 2) the major difference among diets as well as among individual cows was in the amount consumed and energy partition, i.e., milk production or fattening, rather than the ef- ficiency with which ME was used; and 3) the apparently high mai nt enance requirement was not due to milk yield or to lactation per se but may have been influenced by pregnancy. The ad libitum feeding of high protein (19.5%) diets in the experiment of Flatt et al. (37), although necessary to meet the objectives of that experiment, provided substantially more protein than needed for mai nt enance plus milk production. The effects of excess protein as well as the cont ri but i on of pregnancy were studied with all available data from Behsville, which included 350 trials with lactating cows and 193 with nonlactating cows. The decrease in EB attributable to intake of nitrogen in excess of protein required was 7.3 kcal/g excess Journal of Dairy Science Vol. 64, No. 6, 1981 METABOLISM -- 75TH ANNIVERSARY ISSUE 1129 ni t r ogen (90). The a mount of ME r equi r ed dur i ng pr egnancy was descr i bed ( 57) by t he equat i on ME ( kcal / kg "7s) = 100. 8 + . 567e "174t on day t of gest at i on. These dat a i ndi cat e 11.5% ef f i ci ency of ME f or f et al gain. Mul t i pl e regressi on anal ysi s were used by Moe e t al. (62, 63) t o der i ve est i mat es of mai nt enance needs and par t i al effi ci enci es of mi l k pr oduc t i on and tissue gai n in Tabl es 3 and 4. Part i al effi ci enci es of ME used f or mi l k pr oduct i on and body gain in l act at i ng cows were 64 and 75% and ef f i ci ency of mat er nal body gain in nonl act at i ng cows was 60%. The ef f i ci ency of use of body t i ssue ener gy f or mi l k pr oduc t i on by cows in earl y l act at i on was es t i mat ed by compar i ng par t i al regressi on coef f i ci ent s r epr esent i ng t he amount of ME r equi r ed f or mi l k pr oduct i on and t he a mount of di et ar y ME spar ed by body t i ssue loss. The est i mat ed conver si on of body t i ssue ener gy t o mi l k ener gy was 82% ef f i ci ent and ~i kel y refl ect s subst ant i al di r ect i ncor por at i on of body l i pi ds i nt o mi l k fat . These resul t s showed t hat t e mpor a r y st or age of ener gy as body f at in l at e l act at i on combi ned wi t h use of body f at in earl y l act at i on is near l y as ef f i ci ent as di r ect use of di et ar y ME f or mi l k pr oduc t i on (75% X 82% = 62% vs. 64%). The fi ndi ngs descr i bed in t he pr eceedi ng par agr aph were used by Moe et aI. (55) t o i dent i f y t he r el at i onshi p bet ween di et qual i t y and ef f i ci ency of mi l k pr oduct i on. Ener gy of di et s was expr essed as net ener gy f or l act at i on (NE1). Mai nt enance r equi r ement s es t i mat ed by pool ed l i near regressi on wi t hi n 32 di et s f r om t he 350 t ri al s wi t h l act at i ng cows were 122.1 and 111.3 kcal ME or 78. 9 and 67. 7 kcal NE1 per kg 7s of body weight, dependi ng on whet her ME i nt ake or mi l k ener gy was t he de pe nde nt vari abl e. Because t he average measur ed fast i ng heat pr oduc t i on in t he Beltsville l a bor a t or y (73. 5 kcal/kg' VS) wi t h nonl act at i ng, non- pr egnant dai r y cows f ol l owi ng a per i od of mai nt enance f eedi ng (35) was bet ween t he regressi on est i mat es ( 78. 9 and 67. 7) of t he NE1 r equi r ed f or mai nt enance, t he aut hor s concl uded t hat t he a mount of ener gy r equi r ed f or mai n- t enance of l act at i ng cows coul d be descr i bed adequat el y as 73 kcal NE1 ( or NEmi l k) / kg "Ts and t hat a separ at e NE t er m f or mai nt enance (NE m) was unnecessar y. Wi t h t hat assumpt i on, t he NE1 of i ndi vi dual di et s was st udi ed by r el at i ng NE1 of di et dr y ma t t e r (DM) t o ot he r expr essi ons of ener gy as f ol l ow (regressi on coef f i ci ent -+ SE): NEI (Mcal / kg DM) = - - . 19 + (. 703 + . 020) ME ( Mcal / kg DM), NE1 (Mcal / kg DM) = - - . 36 + ( . 677 + . 022) DE (Mcal / kg DM), and NE 1 (Mcal / kg DM) = - - . 12 + (. 0266 + . 0011) TDN (% of DM). The ME, DE, arid t ot al di gest i bl e nut r i ent s ( TDN) in t hese equat i ons were t hose act ual l y obser ved in t he l act at i ng ani mal , and t he aut hor s emphasi zed t hat t hose r el at i onshi ps were not appr opr i at e for meas ur ement s of di gest i bi l i t y at mai nt enance. These resul t s i ndi cat e 61 t o 64% ef f i ci ency of ME use for mi l k pr oduct i on f r om nor mal diets. In a r ecent anal ysi s of Bel t svi l l e dat a (61), resul t s of 313 ener gy bal ance t ri al s wi t h l act at i ng cows publ i shed si nce 1970 were used t o s t udy t he same r el at i onshi ps. The r el at i onshi ps bet ween NE1 and ot her expr essi ons of ener gy f r om t hi s separ at e dat a set were: NE1 (Mcal / kg DM) = - . 2 1 + (. 697 -+-+ . 022) ME (Mcal / kg DM), NE 1 ( Mcal / kg DM) = - . 4 1 + (. 673 -+ . 021) DE (Mcal / kg DM), and NE1 ( Mcal / kg DM) = - . 5 1 + (. 0315 -+ . 0015) TDN (% of DM). The ME and DE equat i ons are vi r t ual l y i dent i cal t o t hose deri ved earl i er wi t h a t ot al l y di f f er ent dat a set. The coef f i ci ent f or t he TDN equat i on is a bout 18% gr eat er t han in t he ear l i er equat i on, i ndi cat i ng a gr eat er ef f ect of per cent TDN on NE 1 t han in t he earl i er dat a set. The change in t he TDN equat i on is unexpl ai ned al t hough t he mor e r ecent dat a set i ncl uded several di et s cont ai ni ng silage f or whi ch dr yi ng losses and et her ext r act anal yses may have i nt r oduced er r or s not in ME and DE dat a. Gr eat es t rel i ance shoul d be pl aced on ME and DE equat i ons because t hey are based on di r ect combus t i on of wet mat er i al . For pr act i cal use t he ME equat i ons given above can be si mpl i f ed t o NEI (Mcal / kg DM) = --. 2 + .7 ME ( Mcal / kg DM) in whi ch ME has been adj ust ed for i nt ake and associ at ed effect s. Maintenance and Growth Measur ement of ener gy cost of gr owt h in Journal of Dairy Science Vol. 64, No. 6, 1981 1 1 3 0 MOE TABLE 3. Mul t i pl e r egr essi on anal ys i s of me t a bol i z a bl e ener gy (ME) i nt ake (Mcal ME/ da y) dur i ng 543 ener gy bal ance me a s u r e me n t s wi t h dai r y cows a, Met abol i c Body t i ssue Body t i ssue b o d y size Mi l k e ne r gy gai n l oss ( kg "75 ) (Mcal) (Mcal) (Mcal) Co n s t a n t Lact . , neg. bal ance ( N=126, R ~ =. 957, Sy. x=1. 886, ME=3 0 . 0 6 0 ,+ 9. 0 McaI) Coef f i ci ent . 153 ,+ . O12 1. 512 -+ . 034 Ave rage 114. 6 14. 882 Lact . , pos. bal ance ( N=224, R u =. 950, Sy. x=2. 025, ME=32. 726 +- 9. 0 Mcal ) Coef f i ci ent . 135 ,+ . 009 1. 576 + . 029 1. 378 ,+ . 054 Aver age 113. 0 9. 416 3. 288 Dr y cows, neg. bal ance ( N=75, R2 - . 7 0 7 , Sy . x =l . 7 3 5 , ME=I O. 401 ,+ 3.1 Mcal ) Coef f i ci ent . 050 -+ . 015 Aver age 129. 9 Dr y cows, pos. bal ance ( N=118, R 2 =. 897, Sy . x =l . 5 0 3 , ME=18. 140 ,+ 4. 6 Mcal ) Coe f f i c mnt . 089 ,+ . 011 1. 703 -+ . 058 Aver age 128. 1 3. 160 La c t a t i ng cows ( N=350, R2=. 952, Sy . x =l . 9 8 5 , ME=31. 766 + 9. 0 Mcal ) Coef f i ci ent . 141 +- . 007 1. 552 ,+ . 22 1. 339 ,+ . 045 Aver age 113. 6 11. 366 2. 101 Dr y cows ( N=193, R2=. 911, Sy . x =l . 6 7 6 , ME=15. 133 ,+ 5.6 Mcal ) Coef f i ci ent . 072 ,+ . 009 1. 677 -+ . 055 Aver age 128. 8 1. 932 All cows ( N=543, R2=. 968, Sy. x=2_075, ME=2 5 . 7 4 0 ,+ 11. 5 Mcal ) Coef f i ci ent . 104 . 006 1. 623 ,+ . 014 1. 473 ,+ . 036 Aver age 119. 8 7. 398 2. 045 1. 270 ,+ . 045 - - 2. 889 - - 5. 479 - - 1. 889 . 990 -+ . 091 6. 781 - - 2. 904 1 . 4 0 1 1. 279 + . 034 - - 2. 152 - - 1. 972 . 933 -+ . 065 3. 670 - 1 . 1 2 8 1. 234 +- . 028 . 622 - 1. 937 aDat a f r om Moe et al. (62). c a t t l e i s ma d e d i f f i c u l t b y t h e c o mb i n e d e f f e c t s o f a p p a r e n t d e c l i n i n g ma i n t e n a n c e n e e d s as t h e a n i ma l a p p r o a c h e s ma t u r i t y a n d t h e c h a n g e s i n c o mp o s i t i o n o f t i s s u e d e p o s i t e d wi t h a g e a n d l e ve l o f f e e d i n g . F a s t i n g h e a t p r o d u c t i o n o f c a t t l e wh e n e x p r e s s e d p e r u n i t o f me t a b o l i c s i z e d e c l i n e s wi t h a g e as i n T a b l e 5. Va r i o u s e x - p o n e n t s o f b o d y we i g h t h a v e b e e n u s e d t o d e s c r i b e me t a b o l i c s i z e ( 4 7 ) . E v e n s ma l l d i f f e r - e n c e s i n e x p o n e n t p r o d u c e l a r g e d i f f e r e n c e s i n e s t i ma t e s as c a n b e s e e n b y c o mp a r i n g f a s t i n g h e a t p r o d u c t i o n e x p r e s s e d p e r u n i t we i g h t r a i s e d t o t h e e x p o n e n t s . 73 a n d . 7 5 i n T a b l e 5. A s o r t o f g e n t l e ma n ' s a g r e e me n t t o e x p r e s s r e s u l t s o f EB me a s u r e me n t s b y t h e e x p o n e n t . 7 5 wa s r e a c h e d a t t h e t h i r d S y mp o s i u m o n E n e r g y Me t a b o l i s m. T h i s a g r e e me n t wa s i n- t e n d e d t o f a c i l i t a t e c o mp a r i s o n s o f r e s u l t s f r o m TABLE 4. Es t i ma t e s of ma i n t e n a n c e r e q u i r e me n t a nd par t i al ef f i ci ency o f ener gy us e in t he dai r y cow a. ME b f or Milk Ti s s ue Mi l k ma i n t e n a n c e f r om ME f r om ME f r om t i ssue N ( kcal / kg "Ts ) (%) La c t a t i ng cows 350 122 64. 4 74. 7 82. 4 Nonl a c t a t i ng cows 193 100 59. 6 aDat a f r o m Moe et al. (62). bME is me t a bol i z a bl e ener gy. J our na l of Dai r y Sci ence Vol . 64, No. 6, 1981 METABOLISM -- 75TH ANNIVERSARY ISSUE 1 131 TABLE 5. Preferred fasting heat production of cattle. Age of Body Fasting metabolism animal a weight b (months) (kg) (kcal/kg.73 a) (kcal/kg'~S c) 1 55 140 129 3 80 135 124 6 150 125 113 12 275 110 98 18 400 100 89 24 525 95 84 36 650 90 79 48 650 85 70 48 650 80 70 aFrom ARC (1). bsuggested mean body weights of growing large breed dairy cattle for corresponding ages, from NRC (66). CRecalculated from Columns 2 and 3. di f f er ent l abor at or i es. The use of body wei ght in kg 'Ts r educes var i at i on f r om mat ur e body size in fast i ng heat pr oduc t i on and pr es umabl y mai nt enance r equi r ement . The use of met abol i c body size t o par t i t i on ener gy use bet ween mai nt enance and pr oduc t i on is appar ent l y less sui t abl e f or t he young gr owi ng ani mal t han f or adul t s. Recent wor k on t he par t i t i on and ener gy cost of pr ot ei n and f at gai n and wor k on t he ener gy cost of pr ot ei n t ur nover , however, is hel pi ng t o cl ar i f y ener gy t r ansact i ons in young growi ng ani mal s. Several r ecent reviews di scuss mani pul at i on of gr owt h (32), ener gy cost of gr owt h (52), and nut r i t i on and genet i c ef f ect s on body composi - t i on (50). Thor bek (83) c omput e d effi ci enci es for body gai n in gr owi ng pigs wi t h Br i er em' s (24) est i mat e of mai nt enance needs, 196.3 kcal ME per kg -s6 body wei ght . Decl i ne in ef f i ci ency of ME f or gain was l i near wi t h i ncreasi ng per cent age of t ot al gain as pr ot ei n. Using a l i near f unct i on f or mai nt enance ME (1683 + 8.1 LW f or live wei ght s (LW) bet ween 20 and 90 kg), Thor be k (84) f ound par t i al ef f i ci enci es f or pr ot ei n and f at depos i t i on of 43 and 77%. Ki el anowski and Kot ar bi ns ka (46) s t udi ed several exponent s of body wei ght in descr i bi ng r el at i onshi ps bet ween ME i nt ake or heat pr oduct i on and pr ot ei n and f at gai n in gr owi ng pigs. They f ound t he e xpone nt . 734 f i t best and used .75 f or si mpl i ci t y. Ener gy cost of pr ot ei n depos i t i on was 16 kcal ME/ g, and cos t of f at depos i t i on was 13 kcal ME/g. Those est i mat es cor r es pond t o par t i al effi ci enci es of a bout 35 and 71%, r espect i vel y. Part i al ef f i ci ency of pr ot ei n and f at gain in gr owi ng l ambs (30 t o 60 kg) was es t i mat ed f r om EB and fast i ng meas ur ement s of heat pr o- duct i on (10). Part i al effi ci enci es were 76% f or mai nt enance, 35% f or pr ot ei n gain, and 99% f or f at gai n when a wei ght e xpone nt of . 75 was used. Recent wor k wi t h calves (72) and wi t h bul l s and hei fers (38) also shows a l ower ener get i c ef f i ci ency f or pr ot ei n gai n t han f or f at gain. Mi l l ward et al. (52) emphasi zed t hat statis- t i cal i dent i f i cat i on of heat pr oduc t i on associ at ed wi t h pr ot ei n and f at deposi t i on, al t hough usef ul t o pr edi ct gr owt h per f or mance in ani mal s, is ar guabl y mi sl eadi ng in mechani st i c t erms. Pr ot ei n and f at depos i t i ons are not c ompl e t e l y i ndependent even t hough f at depos i t i on is l i kel y mor e mani pul abl e t han pr ot ei n deposi - t i on. I f some f at is depos i t ed as an i nsepar abl e c ompone nt of l ean tissue gr owt h, t he cost of t hat f at depos i t i on will be st at i st i cal l y i dent i f i ed wi t h pr ot ei n depos i t i on and t he ef f i ci ency of pr ot ei n synt hesi s t her eby will be under - est i mat ed. The c ont r i but i on of pr ot ei n t ur nover t o t he appar ent l y high cost of net pr ot ei n synt hesi s and t he hi gher r at e of met abol i s m in young gr owi ng ani mal s has been t he subj ect of r ecent i nt ense st udy. Thi s t opi c was consi der ed in det ai l by Wat er l ow et al. (105). They pr es ent ed Journal of Dairy Science Vol. 64, No. 6, 1981 1132 MOE evi dence f r om rat s t hat pr ot ei n t ur nover decl i nes wi t h age and t hat pr ot ei n degr adat i on is gr eat er in ani mal s gr owi ng mor e sl owl y. Edmunds and But t er y (30) pr esent ed dat a showi ng subst ant i al di f f er ences among speci fi c tissues in t he f r act i onal r at e ( per day) in pr ot ei n synt hesi s; . 058 f or l ean tissue, . 475 f or brai n, and i nt er medi at e r at es f or ot he r tissues. They also i ndi cat ed t hat t he r at i o of synt hesi s t o deposi t i on was 3 : 1. The cl ar i f i cat i on of t he rol e of pr ot ei n t ur nover in t he gr owi ng ani mal will, hopef ul l y, be of consi der abl e val ue in par t i t i oni ng ener gy cost s in young growi ng cat t l e. Anot he r i mpor t a nt f act or in i dent i f yi ng ener gy needs of t he growi ng ani mal is t he ext ent t o whi ch body compos i t i on is i nf l uenced by nut r i t i onal mani pul at i on. Many di et ar y effect s have been summar i zed conci sel y by Bl ack (12) for t he growi ng l amb. His dat a suggest t hat ani mal s gr own at hi gher nut r i t i on will have hi gher body f at t han ani mal s gr own mor e sl owl y al t hough t he di f f er ence in com- pos i t i on becomes progressi vel y less as t he ani mal appr oaches mat ur i t y. Pr ot ei n cont ent of t he body i ncr eased progressi vel y as pr ot ei n cont ent of t he di et ( per cent of ME) was in- cr eased f r om 6% t o 10, 15, and 20%. Thi s response t o pr ot ei n st eadi l y decl i ned wi t h i ncreasi ng body wei ght . Pr ot ei n above 10% of ME had l i , t l e ef f ect on compos i t i on of l ambs wei ghi ng over 30 kg. Tyr r el l et al. (91) par t i t i oned gai n of Her ef or d hei fers i nt o pr ot ei n and f at by car bon and ni t r ogen bal ance in ani mal s in f ast and when given mai nt enance and ad l i t i bum i nt ake. Regressi on of f at depos i t i on on t ot al ener gy depos i t ed i ndi cat ed t hat 95% of t he change in EB was f r om change in f at r et ent i on and onl y 5% f r om change in pr ot ei n r et ent i on. The l i mi t at i on of ener gy r et ent i on al one as t he expr essi on of pr oduct i vi t y in gr owt h is a ppa r e nt f r om resul t s of Tyr r el l and Wal do (93) and Wal do and Tyr r el l (104). In cal ori - met r i c and gr owt h st udi es t hey fed di r ect cut or char dgr ass silage wi t h or wi t hout a mi xt ur e o f . 12% f or ma l de hyde and .14% f or mi c acid. Each silage was fed al one and s uppl ement ed wi t h f or mal dehyde- t r eat ed sodi um casei nat e and fed t o Hol st ei n steers. Tr e a t me nt of t he silage or s uppl ement at i on wi t h casei n i mpr oved ni t r ogen r et ent i on but di d not i nf l uence ener- get i c ef f i ci ency. Increasi ng i nt ake of i nsol ubl e pr ot ei n i ncreased t he pr opor t i on of gai n as pr ot ei n f r om 38 t o 51% of t ot al cal ori es gained. Di s c r e t e E f f e c t s on E n e r g y Use Many sources of ener gy loss are i ncl uded in t he di scussi on of ener get i c ef f i ci ency pr esent ed above. Many of t hese have been s t udi ed speci fi - cally, and t he i nf or mat i on gai ned has i mpr oved our under s t andi ng of t ot al use of ener gy by cat t l e. Webst er et al. (107) di scussed several component s of heat i ncr ement (HI) i ncl udi ng t he cost of eat i ng and r umi nat i ng, t he heat pr oduced by r umen f er ment at i on, and t he i ncreased heat pr oduced by t he tissues of t he gut and t he liver. He ci t ed a range in ener gy cost s of eat i ng of 2.5 cal / kcal ME for grass pel l et s and 36 cal / kcal ME f or fresh grass. He concl uded t hat t he energy cost of r umi nat i ng coul d be di s count ed as a cont r i but i on to HI. Webst er et al. (108) es t i mat ed heat of f er ment at i on in vivo in sheep and f ound 68 cal heat pr oduced per ki l ocal or i e of di gest i bl e energy f r om forage diets. He f ound no di f f er ence due t o di et sour ce in t he a mount of heat pr oduced by t he tissues of t he gut , but heat pr oduc t i on i ncr eased exponent i al l y wi t h in- creasi ng ME i nt ake. At an i nt ake of 143. 4 kcal ME/ kg "Ts, heat production in t he tissues of t he gut was 27 kcal / kg "Ts per 24 h. Fast i ng heat pr oduct i on of gut tissues was 15 kcal / kg "vs. The HI due t o f eedi ng in t he gut was, t her ef or e, 12 kcal / kg "Ts of whi ch 7 kcal was f er ment at i on heat and 5 kcal was aer obi c met abol i s m in t he gut tissues. Webst er et al. (107) concl uded t hat processes of ingestion and di gest i on can account for a bout 25 t o 30% of t ot al HI and t hat mos t of t he var i at i on in t ot al HI mus t be f r om t he nat ur e of subst r at es made avai l abl e by di gest i on as suggest ed by Ar ms t r ong and Bl axt er (2, 3). Envi r onment al t emper at ur es i nf l uence t ot al ener gy use in several di f f er ent ways. Young ( 112) revi ewed ef f ect s of col d envi r onment on ener gy use and emphasi zed t hat t her mal stress is descr i bed t oo f r equent l y in t er ms of t empe> at ur e al one. He ci t es Lee' s (49) compi l at i on of envi r onment al vari abl es ( t emper at ur e, humi di t y, air movement , r adi at i on, pr eci pi t at i on) , ani mal char act er i st i cs (species, age and sex, br eed and t Tpe , met abol i c st at e, coat , accl i mat i zat i on, nut r i t i on and hydr at i on, der angement and disease, i ndi vi dual var i abi l i t y) and cri t eri a of ef f ect ( pr oduct i vi t y, gr owt h, r epr oduct i vi t y, Journal of Dairy Science Vol. 64, No. 6, 1981 METABOLISM - 75TH ANNIVERSARY ISSUE 1133 physi ol ogi cal response, pat hol ogi cal pat t er ns) t o i l l ust r at e t he c ompl e xi t y of descr i bi ng or pr edi ct i ng envi r onment al effect s. Thi s t opi c is covered ext ensi vel y in a separ at e paper in t hi s issue, so I will not pur sue t he t opi c here. Most ener gy met abol i s m research has been under condi t i ons of " t her mal ne ut r a l i t y" so t hat envi r onment al ef f ect s mus t be consi der ed in appl yi ng t he dat a t o ext r eme condi t i ons. One speci fi c effect , however, shoul d be ment i oned here because i t appear s t o oper at e over a wi de range of envi r onment al condi t i ons. Thi s ef f ect is a r educt i on in di gest i bi l i t y wi t h decreasi ng t emper at ur e. Young (112) ci t es several experi - ment s in whi ch t he mean r educt i on in DM di gest i bi l i t y was 1.8 per cent age uni t s per 10 C r educt i on in t emper at ur e. Kennedy et al. (45) r epor t ed t hat decreasi ng t emper at ur e had t he ef f ect of i ncreasi ng r at e of passage of r umen ingesta, whi ch decr eased organi c ma t t e r diges- t i on but i mpr oved ef f i ci ency of synt hesi s of mi cr obi al pr ot ei n. Improvements in Research Techniques I nnovat i on or devel opment in r el at ed fi el ds has had a pr of ound ef f ect on t echni ques avai l abl e t o researchers in ener gy met abol i s m. I mpr oved surgical t echni ques and devel opment of i nt egr at ed el ect r oni c ci r cui t s and i nexpensi ve comput er s have had a pr of ound ef f ect on col l ect i on and anal ysi s of per t i nent dat a. Recent reviews doc ume nt t he devel opment and use of several t echni ques. The i ncr eased use of i nt est i nal l y cannul at ed ani mal s has al l owed i dent i f i cat i on of si t e of di gest i on and di sappear ance of nut r i ent s f r om speci fi c segment s of t he gut (51). The i ncr eased avai l abi l i t y and use of mar ker s has pe r mi t t e d syst emat i c s t udy of t he dynami cs of f ood par t i cl e degr adat i on and passage t hr ough t he gut (33). Met hods of measur i ng bl ood f l ow have been used t o measur e quant i - t at i vel y upt ake of nut r i ent s f r om t he gut (9, 48). I mpr oved anal yt i cal sensi t i vi t y and aut o- mat ed anal ysi s have i mpr oved t he abi l i t y t o i dent i f y and quant i f y i nt er medi ar y met abol i t es . As t hese mor e r ef i ned t echni ques i ncrease our under s t andi ng of t he met abol i s m of speci fi c nut r i ent s and i ndi vi dual tissues, mor e sophi st i - cat ed mat hemat i cal t echni ques are needed t o i nt egr at e t hi s i nf or mat i on i nt o descr i pt i ons of energy use in t he whol e ani mal . Si mul at i on and model i ng can be power f ul t ool s in eval uat i ng hypot hes es a bout nut r i ent use and ani mal pr oduct i on (8, 13, 40). Al t hough cal or i met r y has pr ol i f er at ed dur i ng t he past 25 yr and some novel appr oaches have been used, accur acy has not been i mpr oved over t hat in t he ear l y wor k of Ar ms by and Kel l ner. I ndi r ect cal or i met r y pr ovi des a mea- s ur ement of r es pi r at or y exchange and, in- di r ect l y, heat pr oduct i on. Cal or i met r y serves as a poi nt of r ef er ence in char act er i zi ng ener gy use by t he whol e ani mal . I t is one t echni que among many t o t est hypot hes es regardi ng ener gy use by animals. The mos t ef f ect i ve use of cal ori - met r y will be in exper i ment s in whi ch heat pr oduct i on is measur ed s i mul t aneous l y wi t h rat es of met abol i s m of speci fi c nut r i ent s, Feed Evaluation and Feeding Standards Feedi ng st andar ds in use in 1956 were t he t ot al di gest i bl e nut r i ent s ( TDN) and es t i mat ed net ener gy (ENE) syst ems in t he US and st ar ch equi val ent s (SE) in much of Eur ope. The ENE syst em (64, 65) was based on NE of feeds f or gr owt h and f at t eni ng in compar i s on wi t h t hat of cor n grai n whi ch was assigned 2. 08 Meal NE/ kg dr y mat t er . The st ar ch equi val ent syst em ( 111) was based on t he ear l i er wor k of Kel l ner in whi ch 100 i b of t est f eed was descr i bed in t er ms of pounds of st arch equi val ent . Wi t h a few except i ons , none of t hese syst ems pr ovi ded separ at e values for f at t eni ng and l act at i on. Al l syst ems i mpl i ed t hat rel at i ve val ues of feeds were si mi l ar f or f at t eni ng and l act at i on. As t he def i ni t i on of ener gy r equi r ement s became mor e preci se and as f act or s i nf l uenci ng t he ener gy val ue of feeds were unde r s t ood bet t er , new pr opos al s were advanced f or use in pr act i cal f eedi ng si t uat i ons. A syst em based on ME i ni t i al l y was pr opos ed by Bl axt er (17) and descr i bed in det ai l by ARC (1) and Bl axt er (18). The mai n pr ovi si ons of t hi s ME syst em were: 1) Ener gy r equi r ement s of ani mal s and ener gy val ue of feeds shoul d be expr essed in an ener gy uni t , t he cal ori e. 2) The basi c t a bul a t i on of t he ener gy val ue of feeds shoul d be t he ME, det er mi ned at mai nt enance nut r i t i on. 3) ME r equi r ed for mai nt enance is 1.35 t i mes fast i ng heat pr oduct i on. Journal of Dairy Science Vol. 64, No. 6, 1981 1 1 3 4 MOE 4) Efficiency of ME for mai nt enance and body gain can be expressed as a funct i on of ME concentration. The ME system, although nearly universally accepted as the most scientifically sound system available, was not used widely in practi- cal feeding systems. The most common com- plaint was that it was too complex. Although not widely used directly, parts of the ME system were included in nearly every feeding system developed since that time. The system currently used in the United Kingdom is an ME system expressed in joules described by the Ministry of Agriculture, Fisheries, and Food (53). It is a modification of the ARC (1) system in which intake effects are ignored and ME use for milk production is assumed to be a const ant 62% for all diets. Nehring and coworkers (67, 68, 69, 81) introduced a system in which requirements for maintenance, growth, and lactation are ex- pressed in terms of a feed uni t for fattening. The energy values of diets are comput ed from digestible nutrients and adjusted for digestibility of the total diet. Intake effects in the NRC (66) systems are incorporated into requirements for milk pro- duction in the DE, ME, and TDN systems and into the values of feedstuffs in the NE 1 system. The NEI at 3 x maintenance are computed from 1 x TDN by the equation (60), NE 1 (Mcal/kg DM) = - . 12 + .0245 TDN (% of DM), which assumes a reduction in TDN of 4% per multiple of maintenance. In the Netherlands (97, 99), ME is computed from digestible nutrients and is assumed to decline by 1.8% per multiple of maintenance. The NE1 are converted to a feed uni t (1 VEM = 1.65 kcal NE1) that corresponds to the value of 1 g barley. Vermorel (102) developed a similar system for France in which NE1 also is converted to a feed uni t based on barley (1 UFL = 1.73 Mcal NE1). The NE1 is computed from ME after adjustments for intake and associated effects according to percentage concentrate in the diet and forage quality. In Switzerland, Bickel and Landis (11) described a system that is basically the same as those for the Netherlands and France except that energy units are expressed in joules instead of calories. Energy systems for lactating cows have taken on a variety of outward appearances, especially with regard to the units which are used at the farm. Both calories and joules are used, although use of the joule is increasing as a result of its adoption by most European scienti- fic journals. The ME, NE1, NEg, and various feed units are used. This proliferation of units of expression has occurred despite attempts to avoid confusion by trying to identify a single uni t that could be used on a world wide basis for feed evaluation and formulation of diets for cattle. A working group of the EAAP was established to develop recommendations for standardization of terminology (96). Activities of that group and a workshop sponsored by the International Union of Nutritional Sciences (73) led to emphasis on the different requirements of units for feed evaluation than for practical feeding systems. Feed must be evaluated in such a wa y that the potential value of that feed for animals is identified. If information about feeds is to be compiled from many sources, the measurement should be repeatable and should reflect feed quality rather than effects of the animal or technique used for the measurement. The most descriptive and reproducible measurement of feeds is ME determined at the maintenance intake, as suggested by Blaxter (17). Net energies are suited less well in measuring the value of a feed because such measurements involve animal effects, intake, associative effects, and differences in methods of mea- surement (58). In contrast to the need to use a uniform term (ME at maintenance) to describe the value of feeds, units used in practical feeding systems need not and probably cannot be standardized to the same degree. The uni t used, whether a feed unit, NE, or ME should be understood by the user, should be adequate to describe energy needs of the animal, and should be estimable from ME. Nearly all of the systems introduced recently are similar in that ME at maintenance is used as the starting point. The accumulated knowledge regarding intake and associated effects and efficiency of energy use by the animal then is used to develop working re- quirements and feed values for use in form- ulating rations. Working units such as NE should not be viewed as fixed attributes of feeds but must be upgraded continually as Journal of Dairy Science Vol. 64, No. 6, 1981 METABOLISM - 75TH ANNIVERSARY ISSUE 1135 addi t i onal i nf or mat i on is gai ned a bout how ME of di et s and ME use change wi t h f eed i nt ake and t ype of ani mal pr oduc t f or med. Al l maj or f eedi ng st andar ds t r eat ef f i ci ency of ener gy use pr i mar i l y as a f unct i on of con- cent r at i on of ME in t he di et . A maj or gain in t he useful ness of f eedi ng syst ems, especi al l y f or pr edi ct i ng ani mal per f or mance, will be possi bl e when suf f i ci ent i nf or mat i on is avai l abl e on t he r el at i onshi p bet ween amount s of speci fi c end pr oduct s of di gest i on and per f or mance of ani mal s (58, 59). A maj or gai n in feed evalu- at i on will come t hr ough i dent i f i cat i on of i mpor t a nt f eed at t r i but es t hat i nf l uence nut r i ent avai l abi l i t y ei t her t hr ough t hei r own i nher ent pot ent i al or by t hei r i nf l uence on t he en- vi r onment wi t hi n t he di gest i ve t ract . I nf or ma- t i on on feed at t r i but es i nf l uenci ng nut r i ent avai l abi l i t y will al l ow accur at e pr edi ct i on of i nt ake and associ at ed ef f ect s and also per mi t st rat egi es to mi ni mi ze t hose effect s. The Next 25 Years The mean mi l k pr oduc t i on per cow per year in t op her ds has i ncr eased f r om about 8, 000 kg t o 11, 000 kg in t he past 25 yr. Thi s gai n has been possi bl e t hr ough genet i c i mpr ove me nt and appl i cat i on of f eedi ng and management syst ems t hat are responsi ve t o t he cow' s nut r i ent needs. I am aware of no evi dence t hat pr ecl udes progressi ve i mpr ovement s in t hese areas such t hat her ds averaging 14, 000 kg mi l k per year may be seen by t he year 2006. The f act t hat one cow, Beecher Ar l i nda Ellen, act ual l y pr oduced over 22, 800 kg of mi l k in 305 days i ndi cat es t he bi ol ogi cal possi bi l i t y of pr oduc t i on at sust ai ned high dai l y rat es. What r oadbl ocks mus t be r emoved t o al l ow cows t o achi eve such pr oduct i on? I f r equent l y have hear d t he c omme nt t hat Ellen mus t have had an "unus ual l y ef f i ci ent me t a bol i s m" because she coul d not ot her wi se possi bl y have cons umed enough f eed t o pr oduce at t hat rat e. More l i kel y, El l en' s success was because of t wo fact ors. Fi r st is t he f or mul at i on of a di et t ha t coul d be cons umed in suf f i ci ent amount s wi t hout over l oadi ng her physi ol ogi cal abi l i t y t o mai nt ai n condi t i ons wi t hi n her di gest i ve t r act f or ma xi mum r at e of f er ment at i on, di gest i on, and absor pt i on. El l en is r epor t ed t o have consumed up t o 26 kg of t op qual i t y al f al f a hay and 25 kg of a concent r at e mi xt ur e per day. Second is t hat she was bl essed wi t h an ext r a- or di nar y abi l i t y t o synt hesi ze mi l k f r om avai l abl e nut r i ent s at a high r at e (up t o 88. 7 kg per day) , not necessar i l y mor e ef f i ci ent l y but at a gr eat er rat e. In short , t hi s cow was abl e t o deal wi t h t he stress of hi gh f eed i nt ake and pr oduce mi l k. Much of t he progress in genet i c i mpr ovement in dai r y cows will be t hr ough i mpr ovement s in cows' abi l i t y t o mai nt ai n homeost asi s wi t h regard t o t he envi r onment wi t hi n t he di gest i ve t r act t hr ough sal i vary secret i ons, i nt est i nal secret i ons, and a r educed sensi t i vi t y t o t hose effect s such as r umen fill and bl ood met abol i t es t hat ma y t end t o i nhi bi t f eed i nt ake. Abi l i t y t o cope wi t h t he stress of high feed i nt ake and t o synt hesi ze mi l k bot h will l i kel y be i mpr oved in sel ect i ng ani mal s f or hi gher mi l k yi el d. The rol e of t he nut r i t i oni s t is t o f or mul at e di et s t hat mi ni mi ze t he st resses t hat t ax t he cows' abi l i t y t o consume and di gest large amount s of f eed and t o ef f ect i vel y absor b and met abol i ze t he needed nut r i ent s. To f or mul at e t hese di et s we need t o unde r s t a nd l i mi t at i ons t o i nt ake and di gest i on of feeds and met abol i s m of nut r i ent s. Wi t hout benef i t of a "cr ys t al bal l " I pr opos e t he f ol l owi ng as areas l i kel y t o yi el d t o research in comi ng year s in physi ol ogi cal , not pr i or i t y, or der : 1) Remove cur r ent l i mi t at i ons t o r at e of mi cr obi al degr adat i on of st r uct ur al car bo- hydr at es in t he r umen. Convent i onal wi sdom recogni zes t he need f or subst ant i al amount s of r api dl y f er ment ed f i ber in di et s of l act at i ng cows. New chemi cal t echni ques are needed t o char act er i ze t he resi st ance of f eeds and di et s t o high rat es of degr adat i on. Con- di t i ons wi t hi n t he r umen for opt i mum r at e of f e r me nt a t i on mus t be i dent i f i ed. 2) I mpr ove ef f i ci ency of synt hesi s of mi cr o- bial pr ot ei n. The cur r ent l i mi t at i on in use of hi ghl y degr adabl e pr ot ei n, nat ur al l y occur- ri ng nonpr ot ei n ni t r ogen (NPN), and sup- pl ement al NPN by l act at i ng cows and young growi ng cat t l e is t he unf avor abl e r at i o of mi cr obi al pr ot ei n t o t ot al end pr oduct s of f er ment at i on. Evi dence f r om in vi t r o st udi es suggests t hat hi gher effi ci enci es of pr ot ei n synt hesi s are feasi bl e. 3) Prevent unnecessar y di gest i ve losses of energy. I dent i f y opt i mum physi cal - chemi cal condi t i ons wi t hi n each segment of t he gut Journal of Dairy Science Vol. 64, No. 6, 1981 1136 MOE f o r ma x i mu m r a t e o f f e r me n t a t i o n , di ge s t i on, a n d a b s o r p t i o n . I d e n t i f y f e e d f a c t o r s t h a t i n f l u e n c e g u t e n v i r o n me n t . I d e n t i f y me c h - a ni s ms f or h o me o s t a t i c r e g u l a t i o n of g u t e n v i r o n me n t t h r o u g h s e c r e t o r y or t r ans - f er pr oces s es . 4) I d e n t i f y me a s u r a b l e f e e d a t t r i b u t e s or c h a r a c t e r i s t i c s t h a t , wh e n a ppl i e d t o a mi x e d di e t wi t h u n k n o wn i ngr e di e nt s , p e r mi t p r e d i c t i o n of a mo u n t s of s peci f i c n u t r i e n t s l i kel y a b s o r b e d f r o m t h e gut . Suc h c ha r - a c t e r i s t i c s i n c l u d e n o t s i mp l y c o n t e n t of i mp o r t a n t n u t r i e n t s b u t a t t r i b u t e s t h a t a f f e c t n u t r i e n t avai l abi l i t y, i. e. , s u s c e p t i b i l i t y t o hi gh r a t e of f e r me n t a t i o n a n d di ge s t i on, a nd a t t r i b u t e s t h a t i n f l u e n c e g u t e n v i r o n me n t or me c h a n i s m of a b s o r p t i o n a n d t h e r e b y i n f l u e n c e va l ue o f t o t a l di et . 5) I d e n t i f y q u a n t i t a t i v e r e l a t i o n s h i p s be- t we e n e n d p r o d u c t s of di ge s t i on a n d a n i ma l p e r f o r ma n c e , i. e. , t he r ol e of s peci f i c n u t r i - e nt s or g r o u p s o f n u t r i e n t s i n l i mi t i ng mi l k yi e l d or g r o wt h a nd i n f l u e n c i n g p a r t i t i o n o f n u t r i e n t s b e t we e n mi l k a n d b o d y gai n or c o mp o s i t i o n of b o d y gai n. No si ngl e ar ea o f r e s e a r c h wi l l yi e l d mo r e l a s t i ng i mpr ove - me n t s i n a n i ma l p e r f o r ma n c e a nd e f f e c t i ve us e of t h e a va i l a bl e f e e d s u p p l y t h a n an u n - d e r s t a n d i n g of h o w a n i ma l s r e s p o n d t o var - i a t i ons i n a mo u n t s o f ke y n u t r i e n t s a b s o r b e d f r o m t h e gut . 6) De ve l op pr a c t i c a l f e e d i n g s y s t e ms b a s e d on i n f o r ma t i o n f r o m i t e ms 4 a n d 5 above. Ef f e c t i v e n e s s of di e t f o r mu l a t i o n a nd pr e- d i c t i o n of a n i ma l p e r f o r ma n c e c a n be i mp r o v e d gr e a t l y b y d e v e l o p me n t o f ma t h - e ma t i c a l t e c h n i q u e s t h a t r e l a t e b o t h di e t s a n d a n i ma l s ' p e r f o r ma n c e t o q u a n t i t a t i v e d e s c r i p t i o n s of n u t r i e n t s a b s o r b e d f r o m t h e gut . Dy n a mi c mo d e l s t h a t a c c u r a t e l y p r e d i c t i n c r e me n t a l c ha nge s i n a n i ma l p e r f o r ma n c e r e s ul t i ng f r o m c ha nge s i n di e t ar e n e e d e d t o r e pl a c e c u r r e n t l y u s e d s t at i c mo d e l s c on- s i s t i ng of t a bl e s of n u t r i t i v e va l ue o f f e e ds a n d n u t i e n t r e q u i r e me n t s of ani mal s . R E F E R E N C E S 1 Agricultural Research Council. 1965. The nut r i ent requi rement s of farm livestock. No. 2. Rumi nant s. Agric. Res. Counc., London. 2 Armst rong, D. G., and K. L. Blaxter. 1957. The heat i ncr ement of steam-volatile fat t y acids in fasting sheep. Brit. J. Nutr. 11: 247. 3 Armstrong, D. G., and K. L. Blaxter. 1957. The ut i l i zat i on of acetic, propi oni c and but yri c acids by f at t eni ng sheep. Brit. J. Nutr. 11: 413. 4 Armst rong, D. G., and K. L. Blaxter. 1961. The ut i l i zat i on of t he energy of car bohydr at e by rumi nant s. Proc. 2nd Symp. Energy Metab. EAAP Publ. 10:187. 5 Armst rong, D. G., and K. L. Blaxter. 1965. Effects of acetic and propi oni c acids on energy r et ent i on and mi l k secretion in goats. Proc. 3rd Symp. Energy Metabolism. EAAP Publ. 11:59. 6 Armst rong, D. G., K. L. Blaxter, and N. M. Graham. 1957. The heat i ncrement s of mi xt ures of steam-volatile f at t y acids in fast i ng sheep. Brit. J. Nutr. 11:392. 7 Armst rong, D. G., K. L. Blaxter, N. M. Graham, and F. W. Wainman. 1958. The ut i l i zat i on of t he energy of t wo mi xt ures of steam-volatile f at t y acids by fat t eni ng sheep. Brit. J. Nutr. 12: 177. 8 Baldwin, R. L., and J. L. Black. 1979. Si mul at i on of t he effects of nut ri t i onal and physiological st at us on t he growt h of mammal i an tissues: Description and evaluation of a comput er pro- gram. Anim. Res. Labs. Tech. Paper No. 6. CSIRO, Melbourne. 9 Bergman, E. N. 1975. Product i on and ut i l i zat i on of met abol i t es by t he al i ment ary t r act as measured in port al and hepat i c blood. Proc. 4t h Int. Symp. Rumi nant Physiol. Pages 277--291 in Digestion and met abol i sm in t he rumi nant . I. W. McDonald and A.C.I. Warner, ed. Univ. New England Publ. Unit. 10 Bickel, H., and A. Durrer. 1973. Energy ut i l i zat i on by growing sheep. Proc. 6t h Symp. Energy Metab. EAAP Publ. 14 : 119. 11 Bickel, H., and J. Landis. 1978. Feed evaluation for rumi nant s. III. Proposed appl i cat i on of t he new system of energy evaluation in Switzerland. Livestock Proc. Sci. 5: 367. 12 Black, J. L. 1974. Mani pul at i on of body com- posi t i on t hrough nut ri t i on. Proc. Australian Soc. Anim. Prod. 10:211. 13 Black, J. L., N. M. Graham, and G. J. Faichney. 1976. Si mul at i on of prot ei n and energy ut i l i zat i on in sheep. Pages 161- 166 in From pl ant t o animal prot ei n, ed. J. R. McWilliam and R. A. Leng. Univ. New England, Armidale, NSW (cited by Baldwin and Black, (8)). 14 Blaxter, K. L. 1950. Energy feeding st andards for dairy cattle. Nutr. Abstr. Rev. 20: 1949. 15 Blaxter, K. L. 1956. The nut ri t i ve value of feeds as sources of energy: A review. J. Dairy Sci. 39: 1396. 16 Blaxter, K. L. 1961. The ut i l i zat i on of t he energy of food by rumi nant s. Proc. 2nd Symp. Energy Metab. EAAP Publ. 10: 211. 17 Blaxter, K. L. 1962. The energy met abol i sm of rumi nant s. Charles C. Thomas, Publ., Springfield, IL. 18 Blaxter, K. L. 1969. The efficiency of energy t r ansf or mat i ons in rumi nant s. Pro. 4t h Syrup. Energy Metab. EAAP Publ. 12:21. 19 Blaxter, K. L. 1974. Metabolizable energy and Journal of Dairy Science Vol. 64, No. 6, 1981 METABOLI SM -- 75TH ANNI VERS ARY I SSUE 1 13 7 f eedi ng s y s t e ms f or r u mi n a n t s . Proc. Uni v. No t t i n g h a m Nut r , Conf . Feed Manuf . , But t er - wor t hs , Engl and 7: 3. 20 Bl axt er , K. L. 1980. 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