J. Anim Sci. 2006. 84:2515-2522. doi:10.2527/jas.2005-746
© 2006 American Society of Animal Science
Influence of genotype and diet on steer performance, manure odor, and carriage of pathogenic and other fecal bacteria. I. Animal performance1
C. L. Ferrell2,
E. D. Berry,
H. C. Freetly and
D. N. Miller3
USDA-ARS, US Meat Animal Research Center, Clay Center, NE 68933
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Abstract
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Although Brahman crosses constitute a large portion of US beef cattle, little information is available on their response to diverse feed resources compared with Bos taurus steers. Thus, the objectives were to evaluate genotype and diet effects on steer performance during the growing period and subsequent response to a high grain diet during the finishing period. Fifty-one steers [0 (15),
(20),
(7), and
Brahman (9), with the remaining proportion being MARC III] were allotted to 8 pens. Beginning on December 2, steers were individually fed chopped bromegrass hay (n = 26; DM = 85%, CP = 9.5%, ME = 2.19 Mcal/kg) or a corn silage-based diet (n = 25; DM = 51%, CP = 11.9%, ME = 2.75 Mcal/kg) for 119 d. All steers were then fed a high corn diet (DM = 79%, CP = 11.7%, ME = 3.08 Mcal/kg) to a target BW of 560 kg (176 d). Data were analyzed by ANOVA, with genotype, growing diet, and the 2-way interaction included. The interaction was not significant (P > 0.25). The MARC III and
Brahman steers weighed more (P < 0.01) than
or
Brahman steers initially and at the end of the growing period. Weight of bromegrass-fed (325 kg) steers was less than that of corn silage-fed (384 kg) steers at the end of the growing period. Steer ADG and intake of DM, CP, and ME were less (P = 0.087 to 0.001) for
and
Brahman than for 0 or
Brahman steers during growing, finishing, and total, but efficiency of gain did not differ (P > 0.10). Carcass weight, marbling score, quality grade (P < 0.05), and kidney fat (P = 0.06) differed among genotypes. Daily DMI (6.91 vs. 7.06 kg) was similar, but CP (0.66 vs. 0.84 kg) and ME (15.2 vs. 19.4 Mcal) intake of bromegrass fed was less (P = 0.001) than those of corn silage-fed steers. Values for DMI/gain (22.3 vs. 7.43 kg/kg), CP intake/gain (2.13 vs. 0.88 kg/kg), and ME intake/gain (48.8 vs. 20.4 Mcal/kg) were greater (P < 0.001) in bromegrass-fed than corn silage-fed steers. Over the total study, ADG was lower (0.96 vs. 1.01 kg), and DMI (7.82 vs. 7.19 kg), DMI/gain (8.21 vs. 7.10 kg/kg), and ME intake/gain (22.6 vs. 20.9 Mcal/kg) were greater (P < 0.05) in bromegrass-fed than in corn silage-fed steers. Carcass weight, dressing percent, adjusted backfat, and yield grade (P < 0.05) were greater for corn silage-fed than for bromegrass-fed steers. Feed intake and performance, but not efficiency, differed among these genotypes. Compensatory performance during finishing was insufficient to overcome reduced performance during the growing period.
Key Words: beef cattle Bos indicus compensation efficiency
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INTRODUCTION
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Several researchers have suggested that Bos indicus cattle utilize low-quality forage diets more efficiently than Bos taurus cattle (Ashton, 1962
; Kaurue et al., 1972
). Improved performance on low-quality forage diets may result from lower maintenance requirements of Bos indicus compared with Bos taurus in nutritionally restrictive environments (Frisch and Vercoe, 1977
) or greater intake relative to maintenance requirements, rather than improved utilization of low-quality diets per se. In contrast, when high-quality forage (Moran, 1976
) or forage-grain mixtures (Ledger et al., 1970
; ODonovan et al., 1978
) were fed, Bos taurus cattle consumed more feed relative to their maintenance requirements, gained weight faster, and were more efficient than Bos indicus cattle. Similarly, Beaver et al. (1989)
reported that Angus steers consumed more feed and gained faster, but had similar diet digestibility and feed efficiency compared with Brangus steers when fed a high concentrate diet. Krehbiel et al. (2000)
observed little effect of 50% Bos indicus influence on utilization of a high-grain diet. To our knowledge, compensatory gain responses of Bos indicus crossbred steers have not been directly compared with those of Bos taurus crossbred steers after moderate levels of dietary limitation of growth. Results from the cited studies have been used to suggest a diet by genotype interaction on feed utilization and animal performance, but are not conclusive.
Objectives of the current study were to: 1) evaluate diet x genotype interactions in terms of feed intake, feed utilization, and performance of steers fed ground bromegrass hay or corn silage-corn diet, and 2) evaluate effects of diet during the growing period on compensatory responses during the finishing period. Additional objectives are addressed in the companion papers (Berry et al., 2006
; Miller et al., 2006
).
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MATERIALS AND METHODS
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All animal procedures were reviewed and approved by the US Meat Animal Research Center Animal Care and Use Committee.
Fifty-one steers consisting of 15 MARC III (
Angus,
Hereford,
Red Poll, and
Pinzgauer), 20
Brahman x
MARC III (produced by artificial insemination of
Brahman x
MARC III cows to MARC III sires), 7
Brahman x
MARC III (produced by artificial insemination of MARC III cows to Brahman sires), and 9
Brahman x
MARC III (produced by artificial mating of
Brahman x
MARC III cows to Brahman sires) steers from the US Meat Animal Research Center (MARC) cattle populations were used for the study.
Upon weaning, the steers were transported to an intensive cattle confinement area followed by a 28-d adjustment period. The steers were then moved to 1 of 8 pens in a facility equipped with Calan-Broadbent electronic headgates (American Calan, Inc., Northwood, NH). The feeding facility consisted of pens that were approximately 9 x 9 m, with a concrete floor, and approximately one-third under a barn open to the south. Thus, the animals were partially protected from winter weather conditions. Approximately equal numbers of steers of each breed cross were represented in each pen. Steers were trained to use the individual headgates during the ensuing 28 d. During the 56 d postweaning period, the steers were fed a 50:50 blend of the chopped bromegrass hay and the corn silage-based diet (Table 1
) to appetite and were weighed at weaning and at 28 and 56 d postweaning.
At the end of the adaptation period (December 2), the steers were weighed, and 7, 11, 3, and 5 of the 0,
,
, and
Brahman steers, respectively, were fed brome-grass, and 8, 9, 4, and 4 of the 0,
,
, and
Brahman steers, respectively, were fed corn silage individually to appetite (4 pens of steers on each diet). The brome-grass diet was chosen as a moderate-quality forage that might be available as a backgrounding diet and was intended to nutritionally limit ADG to about 0.3 to 0.4 kg/d. The corn silage diet was designed as a growing ration and was intended to result in ADG of approximately 1.0 kg/d. Trace mineral salt was provided in each pen. The growing period lasted 119 d. Steers were weighed at the beginning, end, and at 14-d intervals during the growing period. All steers were then switched, over a 2-wk "step-up" interval, to the high-concentrate diet, and were individually fed ad libitum to achieve a final target weight of 560 kg. Steers were weighed at 14-d intervals until slaughtered.
Steers were fed once daily throughout the study. Samples of each diet were taken daily, frozen, and composited over each 14-d interval. Weight and samples of unconsumed feed were taken at weekly intervals and composited for each 14-d interval. Composite feed and ort samples were subsequently analyzed for DM (oven drying at 70°C to constant weight), carbon and nitrogen (model CN 2000; Leco Corp., St. Joseph, MI), and gross energy (adiabatic bomb calorimeter). Dietary ME and NE values, as well as Ca and P concentrations, were calculated from tabular values for dietary ingredients (NRC, 1996
).
All steers were slaughtered, upon achieving the target weight, at the MARC abattoir. Carcass data, including HCW, cold carcass weight, LM area, fat thickness at the 12th rib, adjusted fat thickness, percentage KPH, marbling score, quality grade, and yield grade were recorded by trained MARC personnel.
Steer weight or cumulative feed intake was regressed on days on feed for each animal for the growing and finishing periods. Those regressions were then used to calculate initial BW, BW at the end of the growing period, final BW, rate of gain, total DMI, and daily DMI of each steer during each period. Metabolizable energy intake was calculated by the use of DMI and tabular values for dietary ME concentration for each diet (Table 1
). Data were analyzed by ANOVA and individual steer was the experimental unit. The statistical model included steer breed cross (or genotype), grower diet, and the 2-way interaction (Snedecor and Cochran, 1973
). Means were compared by Students t-test if the F-test was significant. Residual ADG for the growing period, finishing period, and total study was derived from the regression of ADG on daily ME intake and beginning BW for the respective feeding periods with all animals included. Similarly, residual ME intake was derived as residuals from the regression of daily ME intake on ADG and beginning BW for the respective feeding periods, with all animals included. These approaches were similar to those reported by Koch et al. (1963)
and discussed by Pitchford (2004)
.
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RESULTS AND DISCUSSION
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The breed group (proportion Brahman) x diet interaction was not a significant (P > 0.25) source of variation for any trait evaluated; thus, least squares means are presented for the main effects of proportion Brahman and diet. Initial BW of
Brahman steers was greater (P < 0.05) than weights of other steers (Table 2
). At the end of the 119-d growing period,
Brahman steers were heaviest (P < 0.05), MARC III steers were intermediate and not different from
Brahman, and
Brahman were lightest and not different from the
Brahman. The ADG of MARC III and
Brahman steers was similar and greater (P < 0.05) than ADG of
and
Brahman steers. Daily DM, CP, and ME intake followed similar patterns among genotypes. However, neither DMI/gain (kg/kg), CP intake/gain (kg/kg), ME intake/gain (Mcal/kg), residual ADG, nor residual ME intake differed (P > 0.16) among steer breed groups. Because no breed cross by diet interaction was observed, we conclude that steers having 0,
,
, and
Brahman breeding responded to the different diets similarly. Thus, these results do not support previous suggestions that Bos indicus crosses utilize low-quality forages better than Bos taurus cattle (Ashton, 1962
; Kaurue et al., 1972
). These data are consistent with reports indicating that Bos indicus steers consume less feed (Beaver et al., 1989
; Huffman et al., 1990
), gain more slowly (Rogerson et al., 1968
; Adams et al., 1982
), but have similar feed conversion (Boyles and Riley, 1991
) to Bos taurus steers. Feed consumption and ADG of
Brahman steers was similar to that of MARC III steers, but greater than
and
Brahman steers, consistent with the expected greater level of heterosis for these traits in the
Brahman steers (Dickerson, 1973
; Koger et al., 1975
).
Initial BW was similar for steers fed bromegrass hay and corn silage during the growing period, but steers fed corn silage gained BW faster and weighed more (P < 0.0001) at the end of the growing period. Daily DMI was similar (P = 0.63) between the 2 diets, but CP and ME intake was greater (P < 0.0001) for corn silage-fed steers. The DMI/gain, CP intake/gain, ME intake/gain, and residual ADG values improved with (P < 0.001) corn silage compared with bromegrass hay, but no difference (P = 0.59) in residual ME intake was detected. Intercepts of linear relationships between ADG (kg/d) and ME intake (Mcal/d, Figure 1
) differed (0.136 ± 0.098 vs. 0.213 ± 0.125; P < 0.04) between steers fed brome-grass hay and those fed the corn silage diet, but slopes of the relationships were similar (0.032 ± 0.007 vs. 0.039 ± 0.006, P = 0.49). We interpret these relationships to suggest that ADG of corn silage-fed steers was greater than ADG of bromegrass hay-fed steers at the same ME intake. Conversely, steers fed bromegrass hay required 5.3 Mcal of ME/d more to achieve the same rate of gain as steers fed the corn silage diet. This observation suggests that either maintenance needs were lower for corn silage-fed steers or nutrients from the corn silage diet were utilized more efficiently than those from bromegrass hay, or both. The ratios of CP/ME intakes were similar for the bromegrass hay (43.4 g/Mcal) and corn silage (43.3 g/Mcal) diets suggesting the protein to energy ratio or protein availability was not causing the differences. Numerous other potential explanations exist. The data presented in Figure 1
also demonstrated that use of the pooled regression does not adequately describe the data. In this situation, use of residuals from the pooled regression is inappropriate. Results shown in Figure 1
also show why no differences (P > 0.40) were detected in residual ME intake (Table 1
) due to breed, diet, or the interactions. Similar problems with residual analysis were described in some detail by Darlington and Smulders (2001)
.

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Figure 1. Relationships between ADG and daily ME intake (DMEI) for steers fed bromegrass hay ( ; ADG = 0.14 ± 0.10 + [0.032 ± 0.007 x DMEI], R2 = 0.50) or corn silage ( ; ADG = 0.21 ± 0.13 + [0.039 ± 0.006 x DMEI], R2 = 0.61) during the growing period, as well as the pooled regression (; ADG = 0.57 ± 0.14 + [0.072 ± 0.008 x DMEI], R2 = 0.62).
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All steers were fed the same high-concentrate diet to appetite during the finishing phase of the study. Beginning weight for the finishing period (ending weight for the growing period) differed due to proportion Brahman and diet during the growing period (Table 3
), but as defined by the experimental protocol, final BW was not different due to breed group, previous diet, or the 2-way interaction. However, there were fewer (P = 0.002) days on feed during the finishing period for MARC III (155 d) and
Brahman (134 d) steers than for
(196 d) or
(199 d) Brahman steers, reflecting differences in beginning weight and a trend (P = 0.096) for differences in rate of gain. Daily DM, CP, and ME intakes by MARC III and
Brahman steers were greater than daily intakes by
or
Brahman steers. However, because of the greater time required to achieve the target final weight, total ME intakes during the finishing period were 3,928, 4,530, 3,503, and 4,127 Mcal for 0,
,
, and
Brahman steers (P = 0.016), respectively. Neither DMI/gain, CP intake/gain, ME intake/gain, nor residual ADG differed (P > 0.12) among breed groups indicating that even with the substantial differences in feed intake and days on feed, estimates of feed conversion did not differ substantially among the breed groups. Residual ME intake tended (P = 0.11) to differ among breed groups, primarily because
Brahman had lower residual ME intake than the other groups. This result is consistent with other measures of efficiency in that the
Brahman had lower feed intake, grew slower, and required more days on feed to reach slaughter weight, and yet had numerically lower DM, CP, or ME consumption per kilogram of gain than other groups of steers.
Steers that had been fed bromegrass hay during the growing period weighed less at the beginning of the finishing period than those fed corn silage, but final weights (by design; P = 0.66) and days on feed (P = 0.99) were similar. Rate of gain by steers that had been fed bromegrass hay was nearly 40% greater (P < 0.0001) during the finishing period than rate of gain of steers that had been fed corn silage. The greater rate of gain reflected, in part, greater (P < 0.01) daily intake of DM, CP, and ME. Measures of feed conversion (DMI/gain, CP intake/gain, ME intake/gain, and residual ADG) were improved (P < 0.01) for steers that had been fed bromegrass hay during the growing period compared with those that had received the corn silage diet. No effects of previous diet or breed cross (P = 0.11) on residual ME intake was detected.
Relationships between rate of gain during the growing and finishing periods (Figure 2
) show that when all animals are included, greater rate of gain during the growing period (ADG1) was associated (P < 0.001) with lower rate of gain (ADG2) during the finishing period (ADG2 = 1.45 0.36 x ADG1, R2 = 0.26). These results are consistent with conventional thinking (Carstens, 1995
), in that greater rates of gain during the growing period are associated with lower subsequent performance. However, when evaluated within dietary treatment group, rate of gain during the finishing period was positively associated (P = 0.02) with rate of gain during the growing period. These observations suggest that animals that had greater rates of gain compared with contemporaries fed the same diet continued to perform at a greater level after being switched to the high-concentrate diet.

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Figure 2. Relationships between rate of gain during the finishing period (ADG2) and rate of gain during the growing period (ADG1) for steers fed bromegrass hay ( ; ADG2 = 1.05 ± 0.11 + [1.00 ± 0.31 x ADG1], R2 = 0.31) or corn silage ( ; ADG2 = 0.87 ± 0.15 + [0.19 ± 0.15 x ADG1], R2 = 0.06) during the growing period, as well as the pooled regression (; ADG2 = 1.46 ± 0.06 [0.36 ± 0.09 x ADG2], R2 = 0.26).
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Relationships between ADG and ME intake during the finishing period for steers that had received brome-grass hay or corn silage during the growing period, as well as the pooled regression, are presented in Figure 3
. The regressions for steers previously fed corn silage and the pooled regression did not deviate from linearity, but the relationship between ADG and daily ME intake for steers fed bromegrass hay during the growing period was nonlinear (quadratic, P < 0.005). The nonlinearity appeared to be largely attributable to 2 steers that had very high intakes (11.9 and 11.7 kg of DMI daily) with approximately average rates of gain (1.32 and 1.44 kg/d). However, removal of the 2 "outliers" resulted in a very similar regression (ADG = 3.42 + 0.336 x DMEI 0.0056 x DMEI2, R2 = 0.64), where DMEI = daily ME intake. The ADG of steers previously fed bromegrass hay (1.32 ± 0.03) or corn silage (1.08 ± 0.03) adjusted to mean daily ME intake differed (P < 0.0001). Furthermore, maximum gain (1.63 kg/d) was achieved at 30.3 Mcal of ME intake per day in steers previously fed bromegrass. At the same ME intake, steers that had been fed corn silage gained 1.21 kg/d. These results indicate that ME consumed was used more efficiently for live weight gain by steers previously fed bromegrass hay. These observations are consistent with previous observations (Carstens, 1995
) and show that part of the compensatory gain response is due to improved efficiency of feed utilization.

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Figure 3. Relationships between ADG and daily ME intake (DMEI) during the finishing period for steers that had received bromegrass hay ( ; ADG = 3.32 ± 0.84 + [0.327 ± 0.062 x DMEI] [0.0054 ± 0.0011 x DMEI2], R2 = 0.64) or corn silage ( ; ADG = 0.57 ± 0.15 + [0.021 ± 0.007 x DMEI], R2 = 0.32) during the growing period, as well as the pooled regression (; ADG = 0.42 ± 0.17 + [0.033 ± 0.007 x DMEI], R2 = 0.33).
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Over the entire study, days on feed were less (P < 0.01) for MARC III and
Brahman steers than for
or
Brahman steers (Table 4
), and rate of gain was greatest for MARC III, intermediate for
Brahman, and least for
and
Brahman steers. Although daily feed intake (DM, CP, or ME) differed (P < 0.01) among breed groups, total ME intake was not different (P = 0.41). Consistent with the latter observation, none of the measures of efficiency of feed utilization differed (P > 0.10) among breed groups.
Initial BW, final BW, and days on feed for the entire study of steers fed bromegrass hay during the growing period were similar to those of steers fed corn silage, but rate of gain tended (P = 0.07) to be lower and daily DMI tended (P = 0.06) to be greater. The daily CP and ME intakes and total ME intake during the entire study of the 2 groups were similar (P > 0.50), but DMI/gain (P < 0.001), ME intake/gain (P = 0.05), and residual ADG (P = 0.03) favored steers fed corn silage during the growing period. No differences in CP intake/gain (P = 0.17) or residual ME intake (P = 0.36) were detected.
Even though final live BW did not differ (P > 0.10) due to breed group, growing period diet, or the interaction, carcass weight differed (Table 5
) due to both breed group (P = 0.02) and growing period diet (P = 0.01). Dressing percent did not differ among breed groups (P = 0.13), but was greater (P = 0.002) for steers fed corn silage during the growing period than for steers fed bromegrass hay reflecting, in part, differences in DMI and presumably, gut fill, during the finishing period. Marbling score and quality grade was lower (P < 0.01) for
and
Brahman than for MARC III or
Brahman steers. Percentage KPH tended to differ (P = 0.06) among breed groups, with
Brahman having the least,
Brahman intermediate, and
Brahman and MARC III having the greatest amount. Neither fat thickness at the 12th rib, LM area, nor yield grade differed (P > 0.17) among breed groups. Influence of proportion Brahman on carcass characteristics was consistent with observations reported by Crouse et al. (1989)
. Growing period diet did not have significant affects (P > 0.20) on marbling score, quality grade, fat thickness, LM area, or KPH, although adjusted fat thickness (P = 0.04) and yield grade (P = 0.05) were greater in steers that had consumed the corn silage diet during the growing period.
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Table 5. Influence of breed composition and diet during the growing period on carcass characteristics of steers slaughtered at similar final weights
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IMPLICATIONS
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Differences in numerous traits among breed crosses were observed in this study. Those differences reflected, in part, lower feed intake, lower performance, and reduced carcass quality characteristics of Brahman compared with MARC III steers. Many of these genotypic differences appeared to be negated by the high level of heterosis of the
Brahman steers. No evidence was observed to support suggestions that Brahman cross-bred steers respond to varying diet quality differently than Bos taurus steers. Performance and feed utilization was reduced in steers grown on bromegrass hay, but compensation was observed when they were switched to a high-concentrate diet, compared with those that were grown on a corn silage diet. However, when evaluated over the total system, compensatory responses were insufficient to totally overcome reduced performance during the growing period.
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Footnotes
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1 Mention of a trade name, proprietary product, or specific equipment does not constitute a guarantee or warranty by the USDA and does not imply approval to the exclusion of other products that may be suitable. 
3 Present address: USDA-ARS, Soil & Water Conservation Research Unit, 121 Keim Hall, East Campus, University of Nebraska, Lincoln 68583-0937. 
2 Corresponding author: ferrell{at}email.marc.usda.gov
Received for publication December 21, 2005.
Accepted for publication April 21, 2006.
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Miller, D. N., E. D. Berry, J. E. Wells, C. L. Ferrell, and H. C. Freetly. 2006. Influence of genotype and diet on steer performance, manure odor, and carriage of pathogenic and other fecal bacteria. III. Odorous compound production. J. Anim. Sci. 84:25332545.[Abstract/Free Full Text]
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