Vol.1, No.3, 89-92 (2011)
opyright © 2011 SciRes. Openly accessible at http://www.scirp.org/journal/OJAS/
Open Journal of Animal Sciences
Effects of physiological age on residual feed intake of
growing heifers
Andrea N. Loyd, Charles R. Long, Andrew W. Lewis, Ronald D. Randel*
Texas AgriLife Research, Overton, TX, USA; *Corresponding Author: r-randel@tamu.edu
Received 22 July 2011; revised 6 September 2011; accepted 20 September 2011.
Using a retrospective evaluation of feed effi-
ciency, this study investigated the effects of
physiological age on residual feed intake (RFI)
in growing heifers. Data were collected during
1973 and 1974 at the McGregor location of the
Texas Agricultural Experiment Station. Heifers
(n = 77) were obtained from a large crossbreed-
ing program utilizing a five-breed diallel mating
scheme using Angus, Brahman, Hereford, Hol-
stein and Jersey breeds. At approximately 6
months of age, pre-pubertal heifers were indi-
vidually penned and received ad libitum access
to a balanced diet. Individual feed intake and
body weight data were collected at 28-day in-
tervals for 84 days prior to puberty and for 90
days after puberty. The diet was changed at
puberty to provide a low er energy density. Con-
sidering all females as cohorts, RFI was calcu-
lated for each heifer for each period using
separate models for the pre- and post-pubertal
periods. A moderate, positive Pearson correla-
tion (r = 0.48; P < 0.001) was detected between
pre- and post-pubertal RFI. Furthermore, heifer
RFI rank was compared between the pre- and
post-pubertal periods using Spearman rank or-
der correlat ion and a si mila r co rrel ation (r = 0.46;
P < 0.001) was revealed. This suggests that RFI
determined during the pre-pubertal period may
only be a moderate predictor of post-pubertal
RFI. As a result, physiological age should be
considered when evaluating cattle for feed effi-
ciency using RFI.
Keywords: Cattle; Puberty; Residual Feed Intake
Feed provided to cattle represents a significant portion
of the cost of producing beef [1]. Identifying and select-
ing cattle that are more efficient at utilizing feed re-
sources can potentially reduce feed usage. Selection
strategies for improving feed efficiency have been shift-
ing away from the conventionally used feed:gain ratio
(FG) and are incorporating residual feed intake (RFI) as
an indicator of feed efficiency in beef cattle. Many vari-
ables can influence the outcome of an RFI evaluation;
however, many of these factors can be controlled (i.e.
test duration, type and amount of diet provided, selection
of cohorts of animals, etc.). The Beef Improvement Fed-
eration (BIF) has developed guidelines to standardize the
methodology for RFI evaluation [2]. Specific to cohort
selection, it is recommended that cattle be at least 240
days of age and not differ by more than 60 days of age at
the beginning of an RFI feeding trial [2]. Although cattle
may be of similar chronological age, they may differ in
physiological maturity since an important biological phe-
nomenon (puberty) may occur during this time in bulls
and heifers.
Brody (1945) suggested that puberty coincides with
the inflection point of the growth curve, which corre-
sponds to a shift in body composition away from lean
accrual and towards fat deposition. Recent studies sug-
gest that differences in body composition may contribute
to differences in RFI [4-7]. Since gonadal steroids asso-
ciated with puberty alter body composition, pre-pubertal
RFI may differ from post-pubertal RFI. The objective of
this study was to investigate the potential consequences
of cohort selection, specifically physiological age, on the
outcomes of RFI evaluations. To accomplish our object-
tive, RFI was evaluated on heifers during two distinct
stages of development (pre- and post-puberty).
Since the expense associated with RFI research is
substantial, this study was conducted using feed intake
(FI) and body weight (BW) data previously collected
from heifers pre- and post-puberty. The FI and BW data
were initially reported as a component of an experiment
[8,9] that did not include RFI analysis. The current study
made use of these historical data to bring new informa-
tion to RFI research.
Furthermore, there are currently very few research
initiatives investigating feed efficiency of replacement
A. N. Loyd et al. / Open Journal of Animal Scienc es 1 (2011) 89-92
Copyright © 2011 SciRes. Openly accessible at http://www.scirp.org/journal/OJAS/
heifers. Most studies use steers and terminal heifers
when evaluating various facets of RFI and feed effi-
ciency [10-13]. Therefore, this study provides unique
and relevant RFI data pertaining specifically to heifers
that will be retained in the cowherd.
2.1. Animals and Experimental Design
Data used for this study were collected during 1973
and 1974 from the McGregor location of the Texas Ag-
ricultural Experiment Station as previously described
[8,9]. Heifers (n = 77) were obtained from a large cross-
breeding program that utilized a five-breed diallel mat-
ing scheme. Heifer breeds included straightbred Angus
(n = 7), Brahman (n = 5), Hereford (n = 5), Holstein (n =
6) and Jersey (n = 5) and F1 Angus × Brahman (n = 3),
Angus × Hereford (n = 4), Angus × Holstein (n = 4),
Angus × Jersey (n = 5), Brahman × Hereford (n = 6),
Brahman × Holstein (n = 6), Brahman × Jersey (n = 6),
Hereford × Holstein (n = 5), Hereford × Jersey (n = 5)
and Holstein × Jersey (n = 5) crosses (reciprocals
pooled). Pre-pubertal heifers were individually penned at
approximately 6 months of age in 3 m by 10 m open,
dirt-floored pens and were allowed ad libitum access to a
balanced diet. After reaching puberty, the diet was
changed for each heifer to reduce energy density (Table
1). Feed intake and BW data were recorded at 28-day
intervals for 84 ± 6 days prior to puberty and for 90 ± 4
days after puberty for each heifer following procedures
described by Long et al. (1979). Puberty was defined as
the first ovulatory estrus. Heifers were exposed to
marker bulls during overnight exercise periods to aid in
estrus detection. Heifers were also examined by rectal
palpation every 3 weeks and when marked by a bull to
determine ovarian activity [9].
2.2. RFI Calculation
In order to compare RFI during two distinct physio-
logical periods, the pre-pubertal and post-pubertal peri-
ods were considered separately for RFI calculation. As
Table 1. Pre- and post-pubertal diet compositions.
Ingredient (as fed) Pre-pubertal Post-pubertal
Sorghum, % 48.5 33.0
Cottonseed meal, % 20.0 10.0
Cottonseed hulls, % 25.0 50.0
Vegetable fat, % 4.0 4.0
Vitamin/mineral supplement, % 2.5 3.0
such, a separate model was fitted for each period to de-
termine pre- and post-pubertal RFI. All heifers were
considered as cohorts for each of the pre- and post-pu-
bertal RFI calculations. Initial BW and average daily
gain (ADG) of each period were computed from linear
regression of BW on day of test using the PROC REG
function of SAS (2002). Mid-test BW was estimated for
each period using the intercept and slope from the re-
gression and adjusting for 3% shrink. Metabolic mid-test
BW (MMBW) was calculated as mid-test BW0.75 for
each period. Expected daily FI was predicted for each
period by linear regression of average daily feed intake
(ADFI) on MMBW and ADG using the GLM procedure
of SAS (2002). The model fitted for each period was
Yi = β0 + β1MMBWi + β2ADGi, (1)
where Yi = expected daily feed intake, β0 = the regres-
sion intercept, β1 = the partial regression coefficient of
FI on MMBW, and β2 = the partial regression coefficient
of FI on ADG. Residual feed intake was computed for
each heifer for each period by subtracting actual feed
intake from expected feed intake.
2.3. Statistical Analysis
Pearson correlation coefficients were determined us-
ing the PROC CORR function of SAS (2002) to corre-
late pre- and post-pubertal RFI. Spearman rank order
correlation was used to evaluate changes in heifer RFI
rank from the pre- to post-pubertal periods [15].
Age, BW, ADG, ADFI, and RFI summary statistics are
presented in Table 2. As expected age, BW and ADFI
were greater for the post-pubertal period than for the
pre-pubertal period. Average daily gain was reduced in
the post-pubertal period, likely as a result of the de-
creased energy density of the diet. By definition, mean
RFI was 0.00 kg/day for both the pre- and post-pubertal
periods. The range between the most efficient and least
efficient heifers was 2.79 kg/day for the pre-pubertal
period and 6.04 kg/day for the post-pubertal period. De-
spite these heifers being individually fed, this range is
consistent with previous reports of RFI for growing cat-
tle receiving ad libitum access to feed in groups using
GrowSafe® [12] or Insentec [13,16] electronic feeding
equipment. Current BIF guidelines suggest that feed
intake data obtained from individually-penned cattle are
inadequate [2]. However, the initial concept of RFI was
published using data from cattle fed in individual pens
[17]. Furthermore, recent results suggest that feeding
cattle in individual pens yields similar results to group-
feeding cattle as long as the cattle are stimulated to come
to the bunk more than once daily [18]. In our study,
A. N. Loyd et al. / Open Journal of Animal Scienc es 1 (2011) 89-92
Copyright © 2011 SciRes. http://www.scirp.org/journal/OJAS/
Table 2. Pre- and post-pubertal summary statistics for growing heifers (n = 77).
Pre-pubertal Post-pubertal
Tr a it1 Mean SD2 Min3 Max4 Mean SD2 Min3 Max4
Initial age, months 5.6 0.6 3.9 7.6 11.8 2.0 8.6 15.9
Age at puberty, months 10.9 2.0 7.7 15.4
Test duration, days 84 6 78 119 90 4 84 94
Initial BW, kg 116.1 25.7 51.3 191.1 273.2 54.6 175.9 377.0
Final BW, kg 196.0 33.7 108.6 285.2 333.1 60.9 223.3 454.2
MMBW, kg 43.0 6.1 26.1 59.3 70.8 10.1 52.3 88.8
ADG, kg/day 0.95 0.17 0.41 1.45 0.67 0.17 0.21 1.17
ADFI, kg/day 6.0 1.0 3.9 8.5 9.0 1.6 5.9 13.5
RFI, kg/day 0.00 0.57 –1.47 1.32 0.00 1.11 –2.08 3.96
1BW = body weight; MMBW = metabolic mid-test body weight; ADG = average daily gain; ADFI = average daily feed intake; RFI = residual feed intake; 2SD
= standard deviation; 3Min = minimum; 4Max = maximum.
feed was transferred to the troughs several times daily
such that the cattle were frequently stimulated to con-
sume feed [8].
Openly accessible at
A Pearson correlation coefficient of 0.48 (P < 0.001)
was detected between pre-pubertal and post-pubertal RFI.
Spearman rank order correlation revealed a similar cor-
relation (r = 0.46; P < 0.001) between pre-pubertal and
post-pubertal RFI. These moderate correlation coeffi-
cients suggest that RFI determined prior to puberty may
be only a moderate predictor of post-pubertal RFI. These
results are consistent with other studies that evaluated
RFI on the same cattle at different ages. Archer et al.
(2002) reported a moderate phenotypic correlation of
0.40 between RFI measured in heifers during the post-
weaning period and again as non-gestating, non-lactating
cows after two parities. Arthur et al. (2001) compared
RFI of weanling and yearling bulls and observed a phe-
notypic correlation of 0.43. A genetic correlation of 0.55
was reported between steers evaluated for RFI during the
growing and finishing phases [11]. A repeatability esti-
mate for RFI of 0.62 was reported between the growing
and finishing phases of Limousin x Friesian heifers [13].
One of the proposed benefits of using RFI as a meas-
ure of feed efficiency in beef cattle is that it accounts for
between-animal variation in maintenance and growth
[20]. Furthermore, RFI is phenotypically independent of
its component traits (ADG and MMBW) and should
allow for comparisons between animals at different
stages of production during the RFI evaluation [21].
Under this principle, the evaluation of RFI using animals
of different ages should be valid. However, physiologi-
cal maturity has been implicated as a source of variation
when determining RFI [16] as feed efficiency measure-
ments are dependent on the stage of maturity of the ani-
mals at evaluation [22].
As cattle grow, composition of their gain shifts from
protein accretion to fat deposition [23]. Since the ener-
getic expense associated with protein accrual is less than
for fat deposition [24], the efficiency with which cattle
convert feed into BW gain is reduced as they mature. A
substantial shift from protein accretion towards adipose
deposition occurs around the time of puberty [3]. These
changes in body composition associated with advancing
physiological maturity could partially explain the mod-
erate correlations between pre- and post-pubertal RFI
reported in this study. This suggests that RFI determined
during the pre-pubertal period may be only a moderate
predictor of feed efficiency during the post-pubertal pe-
Admittedly, it is not possible to separate the effects of
puberty status from the effects of the diet change from
the pre-pubertal to the post-pubertal feeding periods in
the current study. To definitively attribute differences in
pre-pubertal and post-pubertal RFI, the same diet would
have to have been fed throughout the experiment. As this
is a retrospective analysis of previously collected data,
this could not be controlled. Nonetheless, previous ex-
periments that have evaluated RFI on cattle fed the same
diet at different ages observed similar correlations as
reported here [10,19]. As a result, physiological maturity
should be considered when selecting cohorts of cattle for
RFI evaluation.
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