Tema 10 - Material de Lectura 5 - Moe

DAI RY PRODUCT I ON
Energy Metabolism of Dairy Cattle

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.
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
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
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),
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.
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 -
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
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
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
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,
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.
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
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
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 .
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J our na l o f Dai r y Sci ence Vol . 64, No. 6, 1981

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Energy Metabolism of Dairy Cattle

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