Effect of Rumen-Undegradable Protein and Energy on Growth
and Feed Efficiency of Growing Holstein Heifers1,2
G. L. BETHARD, R. E. JAMES,3 and M. L. MCGILLIARD
ABSTRACT
Thirty-two Holstein heifers were used to determine
the effect of energy and rumen-undegradable protein
(RUP) on growth and feed efficiency. Treatment diets
were in a 2 × 2 factorial consisting of two energy
concentrations (supporting 0.6 or 0.9 kg of average
daily gain) and two percentages of RUP (30 or 50% of
crude protein). Isonitrogenous diets based on corn
silage, alfalfa silage, orchardgrass hay, corn, soybean
meal, blood meal, and minerals were fed. Treatment
diets were fed until 385 d of age, constituting phase 1
of the trial. During phase 2 (until calving), heifers
were housed together and fed a common diet for 0.7
kg of average daily gain. Dry matter intake (kilo-
grams per day) during phase 1 was 4.46, 5.42, 7.38,
and 5.95 for heifers fed low energy and low RUP, low
energy and high RUP, high energy and low RUP, and
high energy and high RUP diets, respectively. Aver-
age daily gain (kilograms) during phase 1 was 0.62,
0.74, 1.01, and 0.96, respectively. Dry matter intake
and gain were higher for heifers receiving high energy
diets. Dry matter efficiency was higher for heifers fed
high energy and high RUP diets. Apparent total
digestible nutrient efficiency was higher for heifers
fed high RUP diets. Average daily gain during phase
2 was 0.53, 0.43, 0.33, and 0.50 kg, respectively.
Overall average daily gain was not affected by energy
or RUP levels during phase 1 because the slow
growth of heifers during phase 1 was compensated by
faster growth during phase 2.
( Key words: dairy heifer, rumen-undegradable pro-
tein, energy, average daily gain)
Abbreviation key: ADG = average daily gain, WH =
wither height, WHI = WH index.
Received August 21, 1995.
Accepted April 11, 1997.
1Partially supported by Virginia Agriculture Foundation.
2Conducted as part of NC-119 Regional Research Committee.
3To whom reprint requests should be made.
1997 J Dairy Sci 80:2149–2155
Department of Dairy Science, Virginia Agriculture Experiment Station,
College of Agriculture and Life Sciences, Virginia Polytechnic
Institute and State University, Blacksburg 24061–0315
INTRODUCTION
Research has shown that high RUP diets (40 to
50% of CP) can improve growth and feed efficiency of
dairy heifers and steers. Tomlinson et al. ( 2 2 )
reported improved feed efficiency as RUP increased
from 31 to 55% of CP, using blood meal as a source of
RUP. Similarly, Thonney and Hogue ( 2 1 ) reported
improved feed efficiency, but not improved average
daily gain ( ADG) , when cottonseed meal was
replaced with fish meal in the diet of growing Hol-
stein steers. Zerbini and Polan ( 2 5 ) found improved
ADG, but no differences in DMI, when fish meal
replaced soybean meal in diets of 9-wk-old Holstein
bull calves. Swartz et al. ( 2 0 ) reported that high
RUP diets (37.9 to 46.4% of CP) supplemented with
blood meal resulted in improved feed efficiency from
14 to 25 wk of age but not from 1 to 13 wk of age
when compared with a low RUP (29.7 to 32.9% of
CP) diet containing soybean meal. Other researchers
(6, 7, 9 ) found no response in ADG or feed efficiency
with high RUP diets. Response to RUP is inconsistent
and appears to vary among protein sources.
The first objective of this experiment was to deter-
mine the response of Holstein heifers to widely diver-
gent concentrations of RUP and energy from 6 to 14
mo of age. The second objective was to evaluate the
effect of growth before breeding ( 6 to 14 mo of age)
on performance from breeding until calving.
MATERIALS AND METHODS
Thirty-two Holstein heifers were used to study the
effects of energy and RUP on growth and feed effi-
ciency. Prior to treatment assignment, preweaned
calves were housed in calf hutches and then were
moved to loose housing pens in groups of 4 to 8 after
weaning. Mixed grass hay, calf starter, corn silage,
and alfalfa silage were fed in sufficient quantities to
promote BW gains of 0.60 to 0.80 kg/d. At 2 wk prior
to the start of the trial, heifers were placed in groups
of 8 in the research facility; groups were balanced
according to age and BW. After a 2-wk adjustment
period, treatment diets were randomly assigned to
groups. The first 210 d of data comprised phase 1 of
2149
2150 BETHARD ET AL.
the trial. Mean initial BW (kilograms) were 127, 140,
151, and 130, and mean initial ages were 176, 188,
179, and 177 d, respectively, for heifers fed low
energy and low RUP, low energy and high RUP, high
energy and low RUP, and high energy and high RUP
diets. Treatments were arranged in a 2 × 2 factorial
consisting of two energy concentrations (supporting
0.6 or 0.9 kg of ADG) and two percentages of RUP
(30 or 50% of CP). Because of facility limitations and
limited numbers of heifers of similar ages, only two
treatments could be conducted at one time. Therefore,
during phase 1, the low energy and low RUP and the
high energy and high RUP diets were fed from June
1991 through March 1992, and the high energy and
low RUP and low energy and high RUP diets were fed
from June 1992 through March 1993. Prior research
in this facility with Holstein heifers suggested that
DMI and performance were not altered by season (16,
23).
During phase 1, heifers were housed in a confine-
ment facility with a counter-slope design and were fed
a TMR through a Pinpointer 4000B computerized
feeder (UIS, Inc. Cookeville, TN) as described by
Quigley et al. (16). Phase 2 began at the end of phase
1 and continued until calving. During phase 2, heifers
were housed together with other heifers at the Vir-
ginia Tech Dairy Center (Blacksburg) in open lots
containing marginal quality pasture; the heifers were
offered a diet based on soybean meal and corn silage
that was formulated to achieve 0.7 kg of ADG. Heifers
were bred at a minimum of 14 mo and 340 kg of BW.
Thirty-two heifers completed phase 1, and 26 heifers
completed phase 2. Four heifers fed the low energy
and high RUP diet left the herd for reasons unrelated
to treatments as did 1 heifer fed the high energy and
low RUP diet and 1 heifer fed the high energy and
high RUP diet.
Body weights and wither heights ( WH) were
measured weekly during phase 1 and monthly during
phase 2. Body weights were measured on 3 consecu-
tive d at the beginning and end of phase 1 and were
averaged to achieve accurate starting and ending BW.
Dry matter intakes during phase 1 were calculated
using weekly means of daily as-fed intakes and
nutrient content of diets. The ADG was calculated for
each week during phase 1. Apparent feed efficiencies
during phase 1 were calculated using the ratio of total
intake to total BW gain. For phase 2, ADG was
calculated using BW after calving and BW at the end
of phase 1. The ADG for the first 2 mo of phase 2 was
calculated using BW at the end of phase 1, and BW 2
mo later. Overall ADG was calculated using BW at
the start of phase 1 and postcalving BW. Wither
height index ( WHI) , calculated as the ratio of kilo-
grams of BW to centimeters of WH, was used to
evaluate body condition.
Journal of Dairy Science Vol. 80, No. 9, 1997
Diets fed during phase 1 were fed as a TMR and
contained corn silage, alfalfa haylage, chopped or-
chardgrass hay (mean particle length = 2 cm),
ground shelled corn, and a 2:1 (wt/wt) mineral
(14.5% Ca, 7.0% P, and 17% NaCl) and protein sup-
plement (Table 1). The protein in the supplement
was either ring-dried blood meal or soybean meal
(44% CP). Blood meal was of swine origin and was
processed by Smithfield Packing (Suffolk, VA). Diets
were sampled weekly to monitor the nutrient content
of the TMR, but weekly ingredient analysis was used
to calculate the nutrient content of the diets. Protein
undegradability was estimated using values from the
NRC (11). Crude protein, TDN, DM, and ADF values
from the NRC ( 1 1 ) were used for blood meal and
soybean meal. Forages were analyzed at the Virginia
Polytechnic Institute and State University Forage
Testing Laboratory (Blacksburg). Total digestible
nutrients of forages and shelled corn was calculated
as follows (C. C. Stallings, 1994, personal communi-
cation): corn silage TDN = 80.4 – 0.4810 × ADF;
alfalfa silage TDN = 93.79 – 0.90 × ADF; grass hay
TDN = 100.32 – 0.653 × ADF; and shelled corn TDN =
89.8 – 0.768 × ADF.
Data were analyzed using the GLM procedure of
SAS (17). Dependent variables for phase 1 (DMI,
ADG, BW, WH, and WHI) were analyzed using the
model
Yijkl = m + Ui + Tj + ( U T ) ij + H( ij) k + Wl
+ ( U W ) il + ( T W ) jl + (UTW) ijl + E( ijkl)
where
Yijkl = dependent variable of heifer k fed RUP
i and energy j for wk 1;
m = population mean;
U1 = fixed effect of RUP i ( i = 1 or 2);
Tj = fixed effect of energy j ( j = 1 or 2);
( U T ) ij = fixed effect of interaction of RUP i and
energy j;
H( ij) k = random effect of heifer k fed RUP i and
energy j;
Wl = fixed effect of wk l ( l = 1 to 31);
( U W ) il = fixed effect of interaction between RUP
i and wk l;
( T W ) jl = fixed effect of interaction between
energy j and wk l;
(UTW) ijl = fixed effect interaction of RUP i, energy
j, and wk l; and
E( ijkl) = random residual.
Effects of RUP, energy, and the interaction between
RUP and energy were tested using heifer as the error
term.
 HEIFER NUTRITION AND GROWTH 2151

TABLE 1. Diets fed during phase 1 to heifers at 225 kg of BW. low energy diets, resulting in a lower percentage of
Low Low High High DM. Total digestible nutrients averaged 66.7% for the
energy energy energy energy high energy diets and 60.5% for the low energy diets.
Low High Low High
RUP RUP RUP RUP Acid detergent fiber was highest for the low energy
and low RUP diet, reflecting the high proportion of
( % of DM)
Corn silage 33 28 61
orchardgrass hay in that diet. Concentrations of CP
Alfalfa haylage 21 36 were intended to be similar but were lower in the low
Orchardgrass hay 73 54 20 32 energy and low RUP diet, although the differences in
Dry shelled corn 12
Soybean meal 5 7 3 CP were small and reasonable under practical situa-
Blood meal 5 6 tions.
Mineral supplement1 1 1 1 1 Dry matter intake (Table 3 ) was not influenced by
1Contained 14.5% Ca, 7.0% P, and 17% NaCl. RUP but was higher for heifers consuming high
energy diets than for heifers consuming low energy
diets and tended to differ from NRC ( 1 1 ) recommen-
dations. The DMI of heifers fed the low energy and
Apparent feed efficiencies for ADG, postconfine- low RUP and high energy and high RUP diets were
ment ADG, and overall ADG during phases 1 and 2
similar to NRC ( 1 1 ) recommendations for DMI;
were analyzed using the model
however, the DMI of heifers fed the low energy and
high RUP diet exceeded NRC recommendations by
Yijk = m + Ui + Tj + ( U T ) ij + E( ij) k
1.03 kg/d, and the DMI of heifers fed the high energy
where and low RUP diet exceeded NRC recommendations by
1.39 kg/d. Limited exercise and ad libitum feeding
Yij = dependent variable of RUP i and energy j, might have contributed to higher DMI. Low intakes of
m = population mean, heifers fed the low energy and low RUP diet might be
Ui = fixed effect of RUP i ( i = 1 or 2), attributed to the high ADF content of the diet, reflec-
Tj = fixed effect of energy j ( j = 1 or 2), tive of the high proportion of orchardgrass hay.
( U T ) ij = fixed effect of interaction between RUP i Within energy concentration, high dietary ADF and
and energy j, and
increased orchardgrass hay content apparently
E( ij) k = random residual.
reduced intake. In support of this result, Quigley et
The effects of protein, energy, and the interaction al. ( 1 6 ) found that DMI of dairy heifers decreased
between protein and energy were tested by the when ADF exceeded 21% of ration DM. Tomlinson et
residual. Differences were declared significant at P < al. ( 2 3 ) reported that maximal DMI of dairy heifers
0.05. occurred at 20% ADF.
Heifers consuming low energy diets responded to
high RUP by increasing DMI, but the DMI did not
RESULTS AND DISCUSSION
increase for heifers consuming low energy and low
Treatment diets differed in DM, TDN, ADF, and RUP diets. Conversely, heifers receiving high energy
RUP as expected (Table 2). High energy diets con- diets decreased DMI in response to high RUP, result-
tained more silage and less orchardgrass hay than did ing in a significant interaction between energy and

TABLE 2. Nutrient composition of treatment diets fed during phase 1.

Low energy Low energy High energy High energy
Item low RUP high RUP low RUP high RUP
X SD X SD X SD X SD
DM, % 70.5 6.0 63.6 2.0 55.6 2.5 52.8 2.9
CP, % of DM 13.3 0.9 14.2 0.8 13.8 0.4 13.8 0.5
TDN, % of DM 59.7 0.6 61.4 1.1 67.1 2.1 66.3 1.1
ADF, % of DM 36.2 1.0 30.9 1.7 27.3 2.7 24.5 0.1
RUP,1 % of CP 34.2 1.6 50.0 1.5 26.6 1.4 51.9 1.9
n 30 30 30 30
1Estimated from the NRC (11).

Journal of Dairy Science Vol. 80, No. 9, 1997

2152 BETHARD ET AL.
RUP. Therefore, high RUP diets apparently result in
decreased DMI at 67% TDN but result in increased
DMI at 60% TDN. At a moderate energy concentra-
tion (64% TDN), Tomlinson et al. ( 2 3 ) reported that
DMI decreased as RUP increased from 31 to 55% of
CP, which was not true for heifers fed the low energy
diets in this trial but was true for heifers fed the high
energy diets. In contrast, Mantysaari et al. ( 9 ) and
Zerbini and Polan ( 2 5 ) reported no change in DMI as
RUP increased in the diets of heifers fed a high
concentration of energy (75% TDN). Baumgardt ( 1 )
postulated that DMI is controlled metabolically by
energy balance, implying that DMI may be reduced at
high energy concentrations as the animal attempts to
metabolically seek energy balance. Therefore, energy,
and not RUP, might limit intake at high energy con-
centrations (> 75% TDN). Mantysaari et al. ( 9 )
reported similar DMI to those reported in this trial
(5.66 kg at 210 kg of BW and 6.76 kg at 263 kg of
BW, respectively). Calculated TDN intakes, using
75% TDN, were higher for heifers in the study of
Mantysaari et al. ( 9 ) than for heifers in this trial
(4.25 kg at 210 kg of BW and 5.07 kg at 263 kg of BW
respectively), supporting the conclusion that energy
and not RUP might have limited intake. Swartz et al.
( 2 0 ) found that DMI was highest at moderate percen-
tages of RUP (37% of CP) and lowest at high percen-
tages of RUP (46% of CP) when diets contained 68%
TDN. However, those researchers ( 2 1 ) found that
intake of diets containing 78% TDN was not in-
fluenced by RUP percentage. Therefore, RUP may
negatively influence DMI at moderate energy concen-
trations (64 to 68% TDN) but not at low (< 60%
TDN) or high (> 75% TDN) energy concentrations.
The mechanism to explain this is unclear. It is possi-
ble that at low energy concentrations, a lack of fer-
TABLE 3. Least squares means and standard errors for weekly measures of DMI, BW, average daily gain (ADG), wither height (WH),
and wither height index (WHI) during phase 1.
Low energy Low energy High energy
Item low RUP high RUP low RUP
DMI, kg/d 4.46 5.42 7.38
BW, kg 189.6 221.1 267.2
ADG, kg/d 0.62 0.74 1.01
WH, cm 104.4 107.2 112.9
WHI4 1.79 2.04 2.34
n 240 240 240
1RUP effect.
2Energy effect.
3Interaction between energy and RUP.
4Kilograms of BW/centimeters of WH.
Journal of Dairy Science Vol. 80, No. 9, 1997
mentable substrate may limit microbial growth and
DMI, thus minimizing any influence RUP may have
on DMI. At high energy concentrations, DMI and
microbial growth may be maximized, possibly result-
ing in sufficient AA flow to the small intestine, thus
masking any affect of RUP. Another possible factor
determining DMI may be palatability. Animal by-
products, such as blood meal, in combination with low
quality, first-cutting orchardgrass hay result in an
unpalatable ration. The high RUP diets had similar
DMI (5.42 and 5.95 kg/d), suggesting that palatabil-
ity could have limited intake regardless of energy
concentration. Palatability might also explain why
heifers fed the low energy and low RUP diet had the
lowest DMI. The low energy and low RUP diet had
the highest proportion of orchardgrass hay, resulting
in a dry, unpalatable ration.
Body weight, ADG, WH, and WHI (kilograms of
BW/centimeter of WH) during phase 1 are presented
in Table 3. Body weights differed with respect to
energy but not with respect to percentage of RUP. To
achieve a recommended breeding weight of 340 to 385
kg at 13 mo (3, 12, 15), heifers must gain 0.8 kg/d.
Therefore, the low energy and low RUP diet stimu-
lated inadequate BW gains (0.62 kg/d), and the high
energy diets stimulated more than adequate BW
gains (1.0 kg/d). Body weights and ADG appear to be
directly related to DMI, regardless of RUP percentage
or energy concentration. As DMI increased, BW and
ADG also increased. In contrast, Tomlinson et al.
( 2 2 ) found that as RUP increased from 31 to 55% of
CP, ADG and BW increased, and DMI decreased.
That trial ( 2 2 ) lasted only 50 d; therefore, the
response could have changed over time. Wither
heights were not affected by RUP percentage but
increased as energy concentration increased. This
High energy RUP ×
high RUP SE RUP1 Energy2 Energy3
P
5.95 0.04 0.22 <0.01 <0.01
233.8 0.61 0.93 <0.01 <0.01
0.96 0.03 0.50 <0.01 0.08
109.2 0.09 0.74 <0.01 0.04
2.11 <0.01 0.89 <0.01 <0.01
240
 HEIFER NUTRITION AND GROWTH 2153

TABLE 4. Least squares means and standard errors for apparent feed efficiencies during phase 1.
Low energy High energy High energy High energy RUP ×
Item low RUP high RUP low RUP high RUP SE RUP1 Energy2 Energy3
P
DM, kg of intake/kg
of BW gain 8.12 7.58 7.44 6.43 0.31 0.02 0.01 0.45
CP, kg of intake/kg
of BW gain 1.08 1.08 1.02 0.89 0.04 0.11 0.01 0.11
TDN, kg of intake/kg
of BW gain 4.85 4.65 4.98 4.27 0.20 0.03 0.53 0.21
n 8 8 8 8
1RUP effect (tested by residual).
2Energy effect (tested by residual).
3Interaction between energy and RUP (tested by residual).

finding is supported by the results of Kertz et al. ( 8 ) ,
who found that WH increased when high energy diets
were fed compared with WH when low energy diets
were fed. Tomlinson et al. ( 2 2 ) reported high RUP
diets did not improve WH. The WHI was greater for
heifers fed high energy diets but was not affected by
RUP. Although body condition was not measured in
any other manner, the WHI data imply that heifers
fed high energy diets had greater body condition than
did heifers fed low energy diets.
Apparent feed efficiencies (kilograms of intake/
kilogram of BW gain) during phase 1 are presented
in Table 4. Dry matter and CP efficiencies improved
with high energy diets, and DM and TDM efficiencies
improved with high RUP diets. No significant inter-
actions occurred. Although BW gains did not improve
as RUP increased, feed efficiency increased, which
was the most significant advantage of the high RUP
diets in this trial. Tomlinson et al. (22), Zerbini and
Polan (25), and Swartz et al. ( 2 0 ) also found that
high RUP diets improved feed efficiency. In calves
that were < 13 wk of age, Swartz et al. ( 2 0 ) and
Heinrichs and Garman ( 6 ) reported that feed effi-
ciency did not improve as RUP percentage increased.
During the 1st 2 mo of phase 2, heifers fed all diets
except the low energy and low RUP diet lost BW
(Table 5). Body weight losses during this transition
period might be attributed to the observed increased
activity of heifers when first moved to pasture, which
might have reduced energy available for growth. This
result emphasizes the importance of breeding heifers
that are reared in a confinement facility prior to being
moved to pasture. Body weight losses likely were
detrimental to normal reproductive function (2, 24).
Services per conception were 1.1, 1.4, 1.6, and 2.6 for
heifers fed the low energy and low RUP, low energy
and high RUP, high energy and low RUP, and high
energy and high RUP diets, respectively, and age at
conception was 543, 588, 518, and 560 d, respectively.
Mean age at conception was high at 552 d (18.4 mo),

TABLE 5. Average daily gains (ADG; kilograms per day) for Phase 1, Phase 2, postconfinement, and
overall.
Low energy Low energy High energy High energy
Item low RUP high RUP low RUP high RUP
ADG SE ADG SE ADG SE ADG SE
Phase 11,a 0.62 0.03 0.74 0.03 1.01 0.03 0.96 0.03
Phase 22,b 0.53 0.04 0.43 0.05 0.33 0.04 0.50 0.04
Postconfinement3,b 0.10 0.09 –0.51 0.09 –0.57 0.09 –0.37 0.09
Overall4 0.56 0.31 0.56 0.04 0.59 0.03 0.65 0.03
aEnergy effect ( P < 0.05).
bInteraction between energy and RUP ( P < 0.05).
1Six to 14 mo of age ( n = 960).
2Fourteen months of age until postcalving ( n = 26).
3First 2 mo of Phase 2 ( n = 32).
4Six months of age until postcalving ( n = 26).

Journal of Dairy Science Vol. 80, No. 9, 1997

2154
TABLE 6. Least squares means for calving BW (kilograms) and calving age (days).
Low energy
Item low RUP
X SE
Calving BW1,a 501 23
Calving agea 841 23
n 8
aNo significant effect of energy, RUP, or the interaction between RUP and energy.
1BW after calving.
but services per conception averaged only 1.68. Breed-
ing started after heifers left the confinement facility
or at 14 mo of age. Heifers apparently did not cycle or
displayed less visible signs of estrus for 1 to 2 mo
after leaving confinement. Estrus detection was con-
sistent throughout the trial (20-min daily observa-
tion); therefore, it was assumed not to be the cause of
delayed breeding. Only 4 of the 32 heifers conceived
prior to 15 mo of age.
Heifers fed the low energy and low RUP diet had
the lowest ADG during phase 1 (0.62 kg/d) but the
highest ADG during phase 2 (0.53 kg) compared with
heifers fed other diets. Conversely, heifers fed the
high energy and low RUP diet had the highest ADG
during phase 1 (1.01 kg/d) but the lowest ADG dur-
ing phase 2 (0.33 kg/d). Apparently, heifers that
gained poorly during phase 1 experienced compensa-
tory growth during phase 2. Park et al. ( 1 3 ) reported
that compensatory growth occurred when heifers that
were limited in nutrient intake [15% below NRC ( 1 0 )
recommendations] were offered a diet exceeding NRC
( 1 0 ) requirements by 40%. During the compensatory
period, heifers had higher and more efficient (kilo-
grams of DM/kilogram of BW gain) BW gains than
did heifers fed at 100% of NRC ( 1 0 ) requirements
throughout the trial of Park et al. (13). Although
diets in this study did not differ as drastically in
nutrient content from phase 1 to phase 2 as they did
in the study by Park et al. (13), it is probable that
heifers fed low energy diets experienced compensatory
growth during phase 2. As a result, overall ADG was
not different between treatments, suggesting that
high prepubertal growth rates did not affect calving
BW or age at conception and calving (Table 6).
CONCLUSIONS
Heifers receiving high energy diets supporting 0.9
kg of ADG from 6 to 13 mo of age had the highest
ADG, BW, DMI, and DM and CP efficiencies. Heifers
receiving high RUP diets from 6 to 13 mo of age were
more efficient per unit of DM and TDN consumed
than were heifers receiving low RUP diets. Dietary
Journal of Dairy Science Vol. 80, No. 9, 1997
BETHARD ET AL.
Low energy High energy High energy
high RUP low RUP high RUP
X SE X SE X SE
515 33 523 25 556 25
849 32 808 24 832 24
4 7 7
treatment from 6 to 13 mo of age did not influence
calving BW, calving age, or BW gain from birth until
calving. Apparently, high RUP diets improved feed
efficiency of growing Holstein heifers, particularly for
those fed high energy diets.
ACKNOWLEDGMENTS
The authors thank the Virginia Agriculture Foun-
dation for financial support and workers at the Vir-
ginia Tech Dairy Center.
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Journal of Dairy Science Vol. 80, No. 9, 1997