Published December 8, 2014
Growth, carcass quality, and protein and energy metabolism in beef cattle
with different growth potentials and residual feed intakes1,2
F. C. P. Castro Bulle,* P. V. Paulino,*† A. C. Sanches,*‡ and R. D. Sainz*
*University of California, Davis 95616; †Universidade Federal de Viçosa, Viçosa-MG, Brazil;
and ‡Universidade Católica de Goiás, Goiânia-GO, Brazil
ABSTRACT: Twenty-four beef steers (predomi-
nantly Angus × Hereford, 14 to 18 mo of age, 403 ± 3
kg of BW), were housed and fed in individual pens for
about 122 d. Twelve steers came from a herd that had
been selected for growth (high growth; HG) and the
other 12 from a herd with no selection program (low
growth; LG). Another 6 steers (3 from each group) were
slaughtered at the beginning to obtain the initial com-
position. All steers were fed the same corn-based diet
(3.06 Mcal of ME/kg of DM, 13.6% CP) on an ad libitum
basis. Two weeks before slaughter, total urine was col-
lected for 5 d for estimation of 3-methylhistidine excre-
tion and myofibrillar protein breakdown rates. Com-
pared with LG steers, HG steers had less initial BW
but greater final BW, DMI (7.52 vs. 6.37 kg/d), ADG
(1.33 vs. 0.853 kg/d), G:F (0.176 vs. 0.133 kg/kg), ME
intake (0.233 vs. 0.201 Mcalⴢkg of BW0.75ⴢd−1), and re-
tained energy (RE; 0.0711 vs. 0.0558 Mcalⴢkg of
BW0.75ⴢd−1); gained more fat (676 vs. 475 g/d); and
tended to gain more whole body protein (100 vs. 72 g/
d), with no difference in residual feed intake (RFI).
Estimated net energetic efficiency of gain (kg) and ME
for maintenance (MEm) did not differ between the 2
Key words: beef steer, maintenance, protein turnover, residual feed intake
©2007 American Society of Animal Science. All rights reserved.
INTRODUCTION
Most genetic improvement programs for beef cattle
have emphasized outputs such as BW gain, fertility,
and carcass traits. Inputs must also be considered to
improve the efficiency of production and increase profit
(Herd and Bishop, 2000). Residual feed intake (RFI),
the difference between actual and expected intake, is
1
F. C. P. Castro Bulle acknowledges generous support from Con-
selho Nacional de Desenvolvimento Cientı́fico e Tecnológico (CNPq),
Brazil, during her PhD program.
2
Corresponding author: rdsainz@ucdavis.edu
Received June 8, 2006.
Accepted December 11, 2006.
928
groups, averaging 0.62 and 0.114, respectively. The HG
steers had greater HCW (350 vs. 329 kg), backfat (16.1
vs. 11.6 mm), and yield grades (3.53 vs. 2.80), with a
similar dressing percent, KPH fat, LM area, and mar-
bling score. Skeletal muscle protein gain (70.2 vs. 57.6
g/d) and fractional protein accretion rate (0.242 vs.
0.197 %/d) tended to be greater in HG than in LG steers.
Steers were classified into low (−0.367 kg/d) and high
(0.380 kg/d) RFI classes. Compared with the high RFI
steers, low RFI steers consumed less DM (6.61 vs. 7.52
kg/d) and ME (0.206 vs. 0.234 Mcalⴢkg of BW0.75ⴢd−1)
and tended to gain less fat (494 vs. 719 g/d), but were
similar for initial and final BW, ADG, G:F, protein gain,
HCW, dressing percent, backfat, KPH fat, LM area,
marbling score, and yield grade, as well as for all obser-
vations related to myofibrillar protein metabolism. Re-
sidual feed intake may be negatively correlated with
ME for maintenance. The maintenance energy require-
ment increased by 0.0166 Mcalⴢkg−0.75ⴢd−1 for each per-
centage increase in fractional protein degradation rate,
confirming the importance of this process in the energy
economy of the animal.
J. Anim. Sci. 2007. 85:928–936
doi:10.2527/jas.2006-373
an efficiency trait that is independent of BW and BW
gain (Koch et al., 1963). Like growth, RFI is moderately
heritable (Herd and Bishop, 2000; Arthur et al., 2001),
but little is known about the biology underlying differ-
ences in growth potential or RFI.
Genetic selection for high growth rate produces cattle
with faster muscle protein accretion and greater effi-
ciency of gain (i.e., G:F), perhaps due to a lower mainte-
nance cost or a greater efficiency of gain (Herd et al.,
1991; Oddy et al., 1998). Because the rate of protein
accretion depends on the relative rates of protein syn-
thesis and degradation, rapidly growing animals must
have a greater ratio of protein synthesis to protein deg-
radation than slower growing animals.
The current study aimed to examine the relationships
among growth potential, feed intake, G:F, carcass com-
 Metabolism and residual feed intake in steers
position, energy metabolism, and myofibrillar protein
turnover. The main hypotheses were as follows:
1. Animals with a greater potential for growth have
greater DM intakes, rates of gain, G:F, and pro-
tein:fat in the gain.
2. Animals with a greater growth potential have lower
rates of protein turnover and lower ME require-
ments for maintenance (MEm).
3. Animals with lower RFI have lower DM and ME
intakes, with no change in ADG, G:F, or protein:fat
in the gain.
4. Animals with lower RFI have lower rates of protein
degradation in muscle and lower MEm.
5. The ME requirement for maintenance is positively
associated with the rate of protein turnover.
MATERIALS AND METHODS
Animals and Treatments
All procedures involving animals were approved by
the Animal Care and Use Committee of the University
of California, Davis.
Twenty-four crossbred beef steers, approximately 14
to 18 mo of age, with a mean weight of 403 (± 3) kg
were housed and fed in individual pens for 112 to 135
d (mean = 122 d). The pens were approximately 10 ×
2.5 m, and the steers had ad libitum access to feed
and water. Twelve steers came from the University of
California, Davis (UCD) herd, and the other 12 were
purchased from a local producer. Six additional steers
(3 from each group) were slaughtered at the beginning
of the experiment to estimate initial body composition.
Before the experiment, all steers underwent a growing
program for about 120 d at the same (UCD) feedlot to
minimize the effects of previous nutrition, so that the
steers were grown and finished between July 2003 and
March 2004. All animals were predominantly Angus ×
Hereford, with the purchased cattle having some Gelb-
vieh influence as well. The UCD herd has undergone
genetic selection for postweaning growth for over 15 yr,
including the purchase of semen only from Angus bulls,
with an EPD for yearling weight of >33.6 kg for the
previous 5 yr. By contrast, the purchased cattle came
from a herd with no selection or improvement program.
These 2 groups of animals were designated the high
growth (HG) and low growth (LG) groups, respectively.
All steers were fed the same diet on an ad libitum
basis. The diet was composed (as-fed basis) of flaked
corn (80%), alfalfa hay (5%), oat hay (5%), molasses
(4%), fat (2.5%), sodium bicarbonate (1.25%), urea (1%),
oyster shell (0.50%), trace mineralized salt (0.5%), am-
monium chloride (0.25%), monensin sodium (0.002%;
Rumensin 80, Elanco Animal Health, Greenfield, IN)
and tetrachlorvinphos (0.006%; Rabon 97.3, KMG-Ber-
nuth, Inc., Houston, TX). Estimated ME and CP con-
tents on a DM basis were 3.06 Mcal/kg and 13.6%,
respectively. Feed offered and refused were recorded
929
daily, and the ration levels were adjusted every week
on the basis of the average intake of the previous week
to maintain approximately 10% refusals. The steers
were weighed every 28 d, after being deprived of feed
(but not water) overnight, and ADG was estimated as
the slope of the regression of BW on time in days for
each steer. At each weighing, the steers were scanned
by real-time ultrasound using an Aloka 500V instru-
ment (Corometrics Medical Systems, Wallingford, CT)
with a 3.5-MHz, 17.2-cm linear array transducer. Mea-
sures of subcutaneous fat over the 12th to 13th ribs
were used to determine the optimum slaughter end
point, aiming for an average of 12 mm.
Two weeks before being slaughtered, 12 animals (6
from each group) were taken to metabolic crates, where
they stayed for a period of 10 d (5 d of adaptation and
5 d of collection). Because of a limited number of meta-
bolic crates, collections and slaughters were staggered
over a 3-wk period. At the beginning of the collection
period, 20 mL of H2SO4 was added to the urine collection
container as a preservative. Every 6 h, the urine was
weighed and a 50-mL sample was obtained. Specific
gravity of a subsample was measured to calculate the
total volume of urine excreted in a 24-h period. The
remaining urine sample was frozen at −20°C until
analysis.
Just before slaughter, the animals were weighed
after an overnight feed deprivation. Carcass composi-
tion was estimated by carcass specific gravity according
to Garrett and Hinman (1969). Briefly, the right side
of the carcass was chilled for 48 h, and each quarter was
weighed under water using a digital balance suspended
over a cylindrical tank (diameter, 152 cm; depth, 168
cm). The carcasses were weighed as rapidly as possible,
and the water temperature was recorded so that the
carcass density could be corrected to 4°C. Cold carcass
weight was used in the equation to calculate carcass
density. Identical procedures were used for the 6 ani-
mals at the initial slaughter.
Energy Calculations
The procedures used to compute the energy retained
and the maintenance energy requirement were similar
to those of Lofgreen and Garrett (1968). Mean values
of compositions of the initial slaughter groups were
used to determine the initial empty body composition
of each animal, using different values for each group.
Gains of body components were calculated as the differ-
ence between initial and final weights of the respective
components. Empty body compositions were estimated
from carcass density, and heats of combustion of re-
tained fat and protein were assumed to be 9.385 and
5.539 Mcal/kg, respectively (Garrett and Hinman,
1969). Whole body energy gains (retained energy; RE)
were expressed as Mcalⴢkg of BW0.75ⴢd−1 to account for
differences in BW. Metabolizable energy intakes (MEI)
were estimated by multiplying the average DMI during
the experimental period by the formulated ME content
930 Castro Bulle et al.
(NRC, 2000) of the diet, and were also expressed as
Mcalⴢkg BW0.75ⴢd−1. The individual MEm was taken to
be the x-intercept of the linear regression between RE
and MEI, after calculation of the slope of this equation
(i.e., the net energetic efficiency of gain, kg) from each
steer’s gains of protein and fat (Williams and Jen-
kins, 2003).
Residual Feed Intake
For calculation of RFI, DMI was regressed against
the average metabolic BW and ADG (Archer et al.,
1997) as follows:
DMI, kg/d = β0 + (β1 × BW0.75) + (β2 × ADG) + ε,
where ε represents the RFI (actual DMI minus the
expected DMI). Because there was no significant differ-
ence between equations for the HG and LG groups (data
not shown), a single regression equation was fitted to
all of the data.
The animals were classified into 2 RFI classes ac-
cording to their individual RFI. Animals with individ-
ual RFI ≥ 0.5 SD above or below the mean were classi-
fied as high or low RFI (n = 8/class), respectively. How-
ever, only 4 steers per RFI class were used for the
metabolism studies.
Myofibrillar Protein Metabolism
The myofibrillar protein breakdown rate was esti-
mated from the urinary excretion of 3-methylhistidine
(3MH; Harris and Milne, 1981). The breakdown rate
of all myofibrillar proteins was assumed to be similar,
or at least similarly affected by the phenotype groups.
The fractional degradation rate (FDR) of myofibrillar
protein was calculated from total 3MH excretion and
estimates of the 3MH pool size based on carcass compo-
sition, as follows:
FDR, %/d = ([3MH]urine, ␮mol/L × urine volume, L/d)
× (3MHmuscle, ␮mol)−1,
where [3MH]urine represents the concentration of 3MH
in the urine, and 3MHmuscle is the total muscle 3MH
pool.
The size of the 3MH pool in myofibrillar protein was
calculated by multiplying the estimated skeletal muscle
protein mass by the 3MH content of mixed muscle pro-
tein (3.5106 ␮mol of 3MH/g of muscle protein; Nishi-
zawa et al., 1979). Initial and final skeletal muscle pro-
tein contents (ISMP and FSMP, respectively) were esti-
mated by using the following equations derived using
these cattle (Sainz, unpublished observations):
ISMP, kg = (pISM × BW, kg) × 0.18, and
FSMP, kg = (pFSM × HCW, kg) × 0.18,
where pISM is the proportion of skeletal muscle in the
body of the initial groups (0.3675 and 0.3891 for HG and
LG steers, respectively), and pFSM is the proportion of
skeletal muscle in the carcass of the final slaughter
groups, given by
pFSM = [76.6 − (0.702 × carcass fat, %)]/100.
In both equations, the skeletal muscle was assumed to
be 18% protein. This assumption was based on previous
experience as well as published muscle protein contents
that ranged from 15 to 23% (Atwater and Woods, 1896;
Gopinath and Kitts, 1984; Brackebusch et al., 1991).
The rate of skeletal muscle protein gain was calculated
as the difference between the final and initial skeletal
muscle protein contents divided by the number of days
the animals were on trial. The fractional accretion rate
(FAR) of protein was calculated as the rate of skeletal
muscle protein gain divided by the average total skele-
tal muscle myofibrillar protein pool. The fractional syn-
thesis rate (FSR) of the myofibrillar protein pool was
calculated as the sum of FDR and FAR.
Analysis of Urine
Urinary 3MH analysis was performed with a Bio-
Chrom 30 amino acid analyzer (Biochrom Ltd., Cam-
bridge, United Kingdom). Urine samples were deprotei-
nized by mixing 400 ␮L of urine with 400 ␮L of 10%
sulfosalicylic acid solution. The mixture was centri-
fuged for 25 min at 24,000 × g, the pH was adjusted
to 2 with 0.4 N lithium hydroxide, and 50 ␮L of the
deproteinized supernatant was loaded on the cation-
exchange column. A standard elution with 1.65 M lith-
ium citrate buffer (pH = 3.55) was conducted using
an 8-␮m cation-exchange resin (Blom and Huijmans,
1992). The AA were detected by absorbance at 540 nm
after postcolumn derivatization with ninhydrin reagent
using a reaction coil set at 135°C. For calibration, nor-
leucine (Sigma, St. Louis, MO) was used as an external
standard. Data analysis was performed using the
EZChrome software (Scientific Software, San Ramon,
CA). The creatinine concentration in urine was mea-
sured using the Metra Creatinine assay kit (Quidel
Corp., San Diego, CA).
Statistical Analyses
All calculations, except carcass composition and skel-
etal muscle protein degradation, were made from the
first to the last day of the experimental period. The
growth, composition, and protein degradation data
were analyzed using ANOVA, with phenotype (i.e.,
growth potential group or RFI class) as the main effect.
Simple Pearson correlation analyses were conducted to
study the associations among and between ADG, DMI,
protein and fat gains, G:F, MEm, RFI, and FDR. All
statistical analyses were conducted using Minitab Re-
lease 13 (Minitab Inc., State College, PA). The results
 Metabolism and residual feed intake in steers 931
Table 1. Performance characteristics of beef steers with high growth (HG) or low growth
(LG) potential
Growth potential

Item HG LG Pooled SE1 P-value2

Initial BW, kg 394 409 3.20 0.003

Final BW, kg 546 516 9.51 0.037
ADG, kg/d 1.33 0.853 0.066 <0.001
DMI, kg/d 7.52 6.37 0.150 <0.001
G:F 0.176 0.133 0.0076 0.001
Residual feed intake, kg/d 0.059 −0.064 0.106 0.442
Body protein gain, g/d 100 72 10.5 0.068
Body fat gain, g/d 676 475 63.4 0.037
Retained energy, Mcalⴢkg−0.75ⴢd−1 0.0711 0.0558 0.00451 0.026
ME intake, Mcalⴢkg−0.75ⴢd−1 0.233 0.201 0.004 <0.001
Net energetic efficiency of gain3 (kg) 0.604 0.633 0.0165 0.218
ME requirement for maintenance,3 Mcalⴢkg−0.75ⴢd−1 0.114 0.114 0.005 0.976
1
n = 12/group.
2
Probability of a type I error.
3
Net energetic efficiency of gain was estimated from the relative gains of protein and fat of each animal
according to Williams and Jenkins (2003) and was then used to calculate the ME requirements for mainte-
nance.

were interpreted as statistically significant with a 5% atinine excretion in urine, FDR, and FSR were similar
probability level, but actual P-values are also given. between the HG and LG steers. The ratio of 3MH to
creatinine excretion can be regarded as an alternative
RESULTS indicator of the rate of fractional muscle protein degra-
dation. These values all agree, showing that these
The performance characteristics of beef cattle highly groups did not differ in their muscle masses or in the
selected for growth (HG) and nonselected animals (LG) rates of myofibrillar protein degradation.
are shown in Table 1. Animals in the HG group had As shown in Table 1, there was no difference between
lower initial BW (P = 0.003) but greater final BW (P = HG and LG steers in RFI. This was expected, because
0.037) than animals in the LG group. Steers in the HG RFI is meant to be independent of the size and rate of
group showed greater (P = 0.001) DMI (7.52 vs. 6.37 gain (Kennedy et al., 1993). To examine hypothesis 3,
kg/d), ADG (1.33 vs. 0.853 kg/d), and G:F (0.176 vs. steers were classified according to their RFI. Compared
0.133 kg/kg) compared with LG animals. Steers in the with the high RFI steers, low RFI steers consumed less
HG group tended to gain more whole body protein (100 DM (6.61 vs. 7.52 kg/d; P = 0.029) and ME (0.206 vs.
vs. 72 g/d; P = 0.068) and gained more fat (676 vs. 475 0.234 Mcalⴢkg BW0.75ⴢd−1; P = 0.009), with no differences
g/d; P = 0.037) than LG steers. There was no difference between RFI classes in initial or final BW, ADG, or G:F
between the groups in RFI. As seen for DMI, HG steers (Table 4). Rates of whole body protein gain were similar
had greater MEI (0.233 vs. 0.201 Mcalⴢkg BW0.75ⴢd−1;
P < 0.001) and RE (0.0711 vs. 0.0558 Mcalⴢkg BW0.75ⴢd−
1
; P = 0.026) compared with LG animals. Estimated kg Table 2. Carcass traits of beef steers with high growth
and MEm (Mcalⴢkg BW0.75ⴢd−1) did not differ between (HG) or low growth (LG) potential
these 2 groups, averaging 0.62 and 0.114, respectively. Growth potential
Carcass weights were greater (P = 0.018) in the HG
steers (350 kg) than in the LG group (329 kg; Table 2). Item HG LG Pooled SE1 P-value2
This was a function of increased BW, because there was Final HCW, kg 350 329 5.82 0.018
no difference in dressing percent. Steers in the HG Dressing percent 64.1 63.7 0.383 0.467
group had greater backfat (16.1 vs. 11.6 mm; P = 0.008) Carcass backfat, mm 16.1 11.6 1.08 0.008
and higher yield grades (3.53 vs. 2.80; P = 0.004) com- KPH,3 % 2.50 2.45 0.147 0.829
LM area, cm2 80.3 82.3 1.66 0.388
pared with the LG group. There were no differences
Marbling score4 710 667 38.6 0.443
between groups in KPH fat, LM area, or marbling score. Yield grade 3.53 2.80 0.158 0.004
Myofibrillar protein metabolism did not differ be- Quality grade High Average
tween HG and LG steers, although initial skeletal mus- choice choice
cle protein was greater (25.5 vs. 24.6 kg; P = 0.003) in 1
n = 12/group.
LG steers and skeletal muscle protein gain (70.2 vs. 2
Probability of a type I error.
3
57.6 g/d; P = 0.095) and FAR (0.242 vs. 0.197%/d; P = Kidney, pelvic, and heart fat were estimated visually as a percent-
age of the carcass weight.
0.059) tended to be greater in HG animals (Table 3). 4
Marbling score: 500, Small0; 600, Modest0; 700, Moderate0; 800,
Values for final skeletal muscle protein, 3MH, and cre- Slightly abundant0; 900, Moderately abundant0.
932 Castro Bulle et al.

Table 3. Skeletal muscle protein metabolism of beef steers with high growth (HG) or low
growth (LG) potential
Growth potential
No. per
Item group HG LG Pooled SE1 P-value2

Initial skeletal muscle protein, kg 12 24.6 25.5 0.200 0.003

Final skeletal muscle protein, kg 12 33.2 32.6 0.600 0.480
Skeletal muscle protein gain, g/d 12 70.2 57.6 5.08 0.095
Total 3MH3 in muscle, mmol 12 117 114 2.11 0.486
Total 3MH excretion in urine, mmol/d 6 1.64 1.69 0.259 0.903
Fractional degradation rate (FDR), %/d 6 1.41 1.52 0.227 0.744
Fractional synthesis rate (FSR), %/d 6 1.65 1.71 0.227 0.886
Fractional accretion rate, %/d 12 0.242 0.197 0.0157 0.059
FSR:FDR 6 1.19 1.15 0.032 0.323
Creatinine excretion in urine, g/d 6 8.28 8.91 1.09 0.693
3MH:creatinine, mmol/g 6 0.193 0.191 0.0131 0.924
1
n = 6 or 12/group.
2
Probability of a type I error.
3
3MH = 3-methylhistidine.

between RFI classes, but low RFI steers tended (P =
0.078) to gain less fat (494 vs. 719 g/d) than high RFI
steers. The low RFI steers had a mean RFI of 0.747 kg/
d less than the animals classified as high RFI (−0.367
vs. 0.380 kg/d, P < 0.001). However, estimated kg and
MEm did not differ (P > 0.10) between the RFI classes.
These observations are in agreement with those of
Nkrumah et al. (2006), who reported positive correla-
tions between RFI and the production of methane
and heat.
Dressing percent, HCW, backfat, KPH fat, LM area,
marbling score, and yield grade were similar between
the RFI classes (P > 0.20, Table 5). Likewise, all obser-
vations related to myofibrillar protein metabolism were
similar between the low and high RFI groups (P > 0.10,
Table 6).
Table 7 shows the simple Pearson correlations among
ADG, DMI, protein and fat gains, G:F, MEm, RFI, and
FDR. For all animals in the experiment, DMI, ADG,
and G:F were all positively and significantly correlated
with each other. Fat gain was correlated (P < 0.05)
positively with DMI, ADG, G:F and FDR, and nega-
tively with MEm, and tended (P < 0.10) to be correlated
positively with RFI. Dry matter intake was also corre-
lated (P < 0.05) with RFI. Metabolizable energy for
maintenance was positively correlated with FDR (P <
0.05) and tended to be positively correlated with RFI
(P < 0.10).
On the basis of the positive correlation between MEm
and FDR, we examined this relationship further using
linear regression (Figure 1). The equation for this line
indicates that MEm would increase by 0.0166 Mcalⴢkg
BW0.75ⴢd−1 for each percentage unit increase in the frac-
tional rate of myofibrillar protein turnover.
DISCUSSION
The differences between beef cattle selected for
growth (HG) and nonselected animals (LG) reported
here include greater DM and ME intakes, ADG, fat
gain, RE, and G:F for the HG line animals. Estimated
kg and MEm, however, did not differ between groups,
indicating that the increased gains by the HG group
were associated with the greater intakes. Owens et al.
(1995) discussed the relationship between growth po-
tential and intake, and concluded that the greater in-
take of faster growing animals was caused by their
faster growth. The data from this experiment cannot
confirm or refute that concept, but they do support the
implicit association among these variables. Because of
greater fat accretion, steers in the HG group had heav-
ier and fatter carcasses than their LG counterparts, as
shown by the greater back fat and yield grade.
Results from the growth experiment showed a clear
tendency for greater protein deposition in HG steers,
which could occur only through increased protein syn-
thesis, decreased protein degradation, or both. Animals
selected for growth tended to have greater absolute and
fractional accretion rates of muscle protein than the
LG steers; however FDR and FSR did not differ between
these 2 groups. In this experiment, animals chosen for
the protein metabolism measurements were halter-bro-
ken and trained to the collection equipment months in
advance of the RFI measurement. Therefore, only 4
steers from the high and low RFI classes were used for
protein metabolism measurements, and the range of
RFI between the high and low RFI groups was quite
narrow (0.728 kg/d). Because of the small number of
replicates in the protein metabolism study, these re-
sults must be viewed with caution.
The results of the current study can only partly con-
firm hypothesis 3: As expected, there were no differ-
ences between RFI classes in initial and final BW or
ADG, and DM and ME intakes were associated with
RFI. On the other hand, G:F, estimated kg, and MEm
were similar between RFI classes, as were all of the
muscle protein metabolism measurements. Therefore,
hypothesis 4 could not be confirmed. Although not sta-
 Metabolism and residual feed intake in steers 933
Table 4. Performance traits of beef steers with low or high residual feed intake (RFI)
RFI group

Item Low High Pooled SE1 P-value2

Initial BW, kg 405 402 4.24 0.598

Final BW, kg 533 541 12.7 0.652
ADG, kg/d 1.111 1.182 0.130 0.705
DMI, kg/d 6.61 7.52 0.263 0.029
G:F 0.164 0.155 0.0133 0.630
Residual feed intake, kg/d −0.367 0.380 0.0672 <0.001
Body protein gain, g/d 94.0 82.9 14.8 0.605
Body fat gain, g/d 494 719 83.5 0.078
Retained energy, Mcalⴢkg−0.75ⴢd−1 0.0585 0.0773 0.00705 0.080
ME intake, Mcalⴢkg−0.75ⴢd−1 0.206 0.234 0.0065 0.009
Net energetic efficiency of gain3 (kg) 0.600 0.640 0.0201 0.185
ME requirement for maintenance,3 Mcalⴢkg−0.75ⴢd−1 0.108 0.119 0.059 0.219
1
n = 8/group.
2
Probability of a type I error.
3
Net energetic efficiency of gain was estimated from the relative gains of protein and fat of each animal
according to Williams and Jenkins (2003) and was then used to calculate the ME requirements for mainte-
nance.

tistically significant, the numerical differences between body fat (Szasz et al., 2004), usually those correlations
low and high RFI steers in MEm (0.108 vs. 0.119 Mcalⴢkg were low and the SE relatively high (Basarab et al.,
BW0.75ⴢd−1), estimated kg (0.60 vs. 0.64), and fat:protein 2003), suggesting that the data should be interpreted
in the gain (5.26 vs. 8.67) may have biological relevance, cautiously.
because in combination they were able to explain the Three factors should be considered when evaluating
differences in RFI between groups. these conclusions. First, at the age at which these stud-
Although the fat:protein ratio in the gain was less in ies were conducted (14 to 18 mo), both HG and LG
low RFI steers than in high RFI steers, none of the steers were not in the steep part of their growth curve,
measurements of carcass traits differed between RFI where the phenotypic differences would have been more
classes. This is in agreement with Arthur et al. (2001) apparent. The metabolic rate tends to slow down as the
and Basarab et al. (2003), who reported a low or nonex- animal ages; adult animals usually show lower levels
istent phenotypic correlation between RFI and ultra- of skeletal muscle protein synthesis, degradation, and
sound backfat thickness. However, the relationship be- accretion. Thus, it should be borne in mind that the
tween RFI and carcass characteristics, mainly fat results observed at the age of the animals in the current
traits, has been the most contentious since the introduc- study may not be representative of those of other ages
tion of the RFI concept (Basarab et al., 2003). Although or stages of maturity. Second, different from the animal
some researchers have reported some phenotypic or ge- model used by Oddy et al. (1998), in which the selection
netic correlations or both between RFI and carcass and for HG and LG rate involved only pure Angus cattle
from a common herd, animals in this study came from
different herds and the selection for HG and LG was
Table 5. Carcass traits of beef steers with low or high done within 2 types of crossbred animals (Angus × Here-
residual feed intake (RFI) ford vs. Angus × Hereford × Gelbvieh). Although the
comparison between these 2 crossbred cattle might
RFI group
have confounded the difference that would have been
Item Low High Pooled SE1 P-value2 found between the HG and LG lines if both were from
HCW, kg 340 343 8.48 0.827
the same pure breed, the groups studied reflect a very
Dressing % 63.8 63.4 0.49 0.511 realistic situation, because crossbred cattle are more
Carcass backfat, mm 14.3 12.5 1.52 0.430 commonly raised than purebreds. Finally, it is im-
KPH,3 % 2.38 2.56 0.16 0.781 portant to consider the problems involved with estima-
LM area, cm2 83.2 79.7 2.31 0.297
tion of the skeletal muscle protein pool size. Overesti-
Marbling score4 729 671 46.7 0.399
Yield grade 2.92 3.33 0.237 0.243 mation of this pool would lead to underestimation of
Quality grade High Average FDR in the animal. Some researchers have used an
choice choice approximation (Munro, 1969), considering that most
1
n = 8/group. mammals have between 40 and 50% muscle in their
2
Probability of a type I error. bodies, irrespective of the species or sex of the animals.
3
Kidney, pelvic, and heart fat were estimated visually as a percent- This may be true in nonruminants; however, in cattle
age of the carcass weight.
4
Marbling score: 500, Small0; 600, Modest0; 700, Moderate0; 800 most reports would indicate that approximately 31 to
Slightly abundant0; 900, Moderately abundant0. 35% of the BW is muscle (Callow, 1961; Henrickson et
934 Castro Bulle et al.

Table 6. Skeletal muscle protein metabolism of beef cattle with low or high residual feed
intake (RFI)
RFI group
No. per
Item group Low High Pooled SE1 P-value2

Initial skeletal muscle protein, kg 8 25.3 25.1 0.265 0.600

Final skeletal muscle protein, kg 8 33.7 32.1 0.734 0.140
Skeletal muscle protein gain, g/d 8 67.7 60.2 7.40 0.483
Total 3MH3 in muscle, mmol 8 118 113 2.58 0.140
Total 3MH excretion in urine, mmol/d 4 2.03 1.72 0.308 0.513
Fractional degradation rate (FDR), %/d 4 1.74 1.56 0.292 0.685
Fractional synthesis rate (FSR), %/d 4 1.96 1.75 0.275 0.617
Fractional accretion rate, %/d 8 0.229 0.208 0.0231 0.526
FSR:FDR 4 1.16 1.14 0.0489 0.803
Creatinine excretion in urine, g/d 4 10.5 8.75 1.200 0.347
3MH:creatinine, mmol/g 4 0.189 0.197 0.0159 0.720
1
n = 4 or 8/group.
2
Probability of a type I error.
3
3MH = 3-methylhistidine.

Table 7. Simple correlation coefficients among performance traits, maintenance energy

requirements, and myofibrillar protein degradation in beef steers1
Item ADG DMI Protein gain Fat gain G:F MEm RFI

DMI 0.884***
Protein gain 0.417* 0.267
Fat gain 0.637*** 0.727*** −0.096
G:F 0.966*** 0.744*** 0.447* 0.534**
MEm −0.298 −0.075 −0.015 −0.740*** −0.382†
RFI −0.025 0.440* −0.287 0.375† −0.225 0.421†
FDR −0.432 −0.383 0.067 −0.698* −0.474 0.760** −0.139
1
MEm = metabolizable energy requirement for maintenance; RFI = residual feed intake; FDR = fractional
degradation rate of myofibrillar proteins.
†P < 0.10, *P < 0.05, **P < 0.01, and ***P < 0.001.

Figure 1. Regression between the ME requirement for maintenance (MEm) and fractional degradation rate (FDR)
of myofibrillar proteins: (䊏) high residual feed intake (RFI) steers; (䊐), low RFI steers; (▲), steers with an RFI within
0.5 SD of the mean (zero).
 Metabolism and residual feed intake in steers
al., 1965; Nishizawa et al., 1979) or that approximately
52 to 57% of empty body protein is associated with
skeletal muscle. The use of an incorrect estimate of
muscle in the animal body could lead to as much as a
25% difference in estimated FDR. In this study, skeletal
muscle was estimated to comprise between 29 and 39%
of the whole body.
In balancing energy inputs and outputs, heat produc-
tion represents a substantial component of the energy
budget of ruminant livestock. The fraction of total main-
tenance energy lost as heat generally ranges from 100%
at maintenance (by definition) to about 70% in growing
heifers (Reynolds et al., 1991). The metabolic sources
of this expenditure at both the cellular and organ level
have been reviewed (McBride and Kelly, 1990; Baldwin
and Sainz, 1995) and include the maintenance of cell
structure and functional integrity and processes such
as ion transport and protein turnover. Although varia-
tions in maintenance and G:F are sometimes more
highly associated with weight and metabolic activity of
the visceral organs, such as the gut and liver, than with
body proteins, body fat, or composition of gain (Ferrell
and Jenkins, 1985), our results showed a Pearson corre-
lation coefficient between FDR and MEm of 0.76 and
a positive linear association between MEm and FDR.
Reduced rates of protein degradation allow an increase
in lean body mass without a proportionate increase in
maintenance energy needs. Tomas et al. (1991, 1998)
and Oddy et al. (1998) reported that efficiency of feed
utilization in selected lines of rats, broilers, and Angus
cattle, respectively, was inversely related to rates of
skeletal muscle protein breakdown. This study extends
those observations to show a clear positive relationship
between the rate of muscle protein breakdown and the
maintenance energy requirement.
Frequently, observed differences in efficiency of
growth have been attributed to differences in body com-
position. More specifically, differences in rates of water
and protein accretion relative to the rate of fat accretion
are thought to have a major influence on the rate and
efficiency of BW gain, primarily because of the lower
energy content of water and protein than of fat. Con-
versely, greater maintenance costs have been associ-
ated more closely with body protein than with body fat
(Pullar and Webster, 1977; Ferrell et al., 1979). Our
results show a positive association between MEm and
FDR of myofibrillar protein. Therefore, it is reasonable
to conclude at this point that the animal with a greater
skeletal muscle protein turnover rate, as reflected by
its degradation rate, will have a greater energy require-
ment for maintenance and consequently less gross G:F.
Selection for G:F has been shown to increase the final
body size (Basarab et al., 2003) because of the high
correlation between these variables. This could have a
negative impact on the beef production system, espe-
cially when considering the high cost of animal feed.
On the other hand, improved feed efficiency can be
achieved by selecting animals using RFI without a cor-
related response in mature size. The animals classified
935
as low RFI had BW gains similar to those classified as
high RFI but ate 12% less feed, making them more
efficient in terms of feed utilization. In other words,
animals with lower RFI used less energy in the physio-
logical processes involved in maintenance, resulting in
more net energy available for tissue accretion.
This study demonstrated that RFI may be negatively
correlated with the maintenance energy requirement.
The maintenance energy requirement was positively
related to the rate of myofibrillar protein turnover, con-
firming the importance of this physiological process in
the energy economy of the animal.
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