Heterosis and breed effects for beef traits of Brahman purebred and crossbred steers

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Heterosis and breed effects for beef traits of Brahman purebred and crossbred steers
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Riley, D. G.
Chase, C. C. Jr.
Coleman, S. W.
Phillips, W. A.
Miller, M. F.
Brooks, J. C.
Johnson, D. D.
Olson, T. A.
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University of Florida
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Gainesville, Fla.

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Heterosis and Breed Effects for Beef Traits of Brahman Purebred and
Crossbred Steers

D. G. Riley1, C. C. Chase, Jr.1, S. W. Coleman1, W. A. Phillips2, M. F. Miller3,
J. C. Brooks3, D. D. Johnson4, and T. A. Olson4

Brahman heterosis (with two Bos taurus breeds) for important carcass traits in the U.S. beef
industry was large and beneficial for most traits. Traits related to beef quality and palatability
still have room for improvement in Brahman straightbreds and crosses.


Summary
Heterosis was documented for traits that are
important in the determination of value for beef
carcasses in the United States. Brahman-Angus
carcasses had very high levels of heterosis for
live weight, hot carcass weight, adjusted 12th rib
fat thickness, longissimus area, and USDA yield
grade. Estimates of heterosis for the Criollo
breed Romosinuano were lower than Brahman-
Angus heterosis. Brahman direct effects were to
increase live weight, hot carcass weight,
adjusted 12th rib fat thickness, longissimus area,
and USDA yield grade. However, generally
consistent with previous reports, Brahman sire
and dam breed means for marbling score,
sensory panel overall tenderness, and %
carcasses grading Choice were lower than Bos
taurus breeds; Warner-Bratzler shear force
values for Brahman were higher. Angus-
Romosinuano means for live weight, hot carcass
weight, adjusted 12th rib fat thickness,
longissimus area, and USDA yield grade were
very similar to those raised on the Great Plains
(Wheeler et al., 2010). Neither Brahman-
Romosinuano nor Angus-Romosinuano steers
had any advantages for these traits as compared
to Brahman-Angus. Brahman-Angus crossbred
steers were projected to reach all end points
(hot carcass weight, 12th rib fat thickness, or
marbling score) sooner than most other breed
groups.

Introduction
Since Brahman crossbred cows are so widely
used by cow-calf producers in the Southern
United States, producer profitability is often
affected by the carcass value of the resulting
steers from matings to obtain crossbred
Brahman cows. The reputation for inadequate
carcass quality and tenderness for both Brahman
purebreds and crossbreds results in lower prices


paid at various places in the beef production
system for calves with visible Brahman
inheritance. Bos taurus Criollo breeds, such as
the Romosinuano of Colombia may offer an
alternative to Brahman, or a complement to
Brahman, and result in adapted females that
could be used in cow-calf production and in
steers that may perform adequately in U.S.
feeding and beef production traits.
Romosinuano cattle were available for a
crossbreeding project to investigate such issues
at the Subtropical Agricultural Research Station
in Brooksville. The objectives of this work were
to estimate heterosis and breed genetic effects
for traits affecting carcass value and beef
palatability traits from purebred and crossbred
steers produced by Brahman, Angus, and
Romosinuano cattle.

Procedure
Brahman, Angus, and Romosinuano bulls were
mated to cows of the same breeds in all
combinations. Straightbred (3 groups) and
crossbred (6 groups; reciprocal Fls were
considered to be distinct groups) steers were
spring-born in 2002, 2003, and 2004 (n = 137,
148, and 179 steers, respectively). Steers were
weaned at approximately 7 mo of age in
September of each year in three separate wk. A
total of 11 Brahman, 12 Angus, and 16
Romosinuano bulls sired steers with records in
these data (average of 13, 14, and 10 steers per
sire, respectively).

Approximately one mo after weaning, in mid-
October of each year, steers were transported by
semi-trailer to the Grazinglands Research
Laboratory (GRL) at El Reno, Oklahoma. After
a short receiving period, they were assigned to
two management groups and placed on winter


'Subtropical Agricultural Research Station, USDA, ARS, Brooksville, FL
2Grazinglands Research Laboratory, USDA, ARS, El Reno, OK
3Department of Animal and Food Sciences, Texas Tech University, Lubbock, TX
4 Department of Animal Sciences, University of Florida, Gainesville, FL










wheat pasture until the first week of May in the
subsequent year. Each year, the heaviest 10
steers from each breed group were assigned in
pairs to GRL feeding pens (n = 15; 6 steers per
pen). Assignments were made so that pairs of
steers from each breed group were penned at
least once with pairs of steers from every other
breed group. Those steers that then remained
were then assigned to larger feeding pens (4
pens in two of the project years, 6 pens in the
final project year; n = 12 to 16 steers per pen).

One third of the steers in each pen and breed
group was randomly assigned to each of three
harvest dates in August, September, and October
of each year, which corresponded to averages of
101, 129, and 157 d fed across the three project
years. Steers were transported approximately 5
h to Amarillo, TX, for commercial harvest.

Traits analyzed included those that affect USDA
yield and quality grades, and assessments of
palatability. Hot carcass weight was obtained
from harvest facility records, and was used in
conjunction with final live weight (shrunk
weight prior to transportation) to estimate
dressing percentage. After chilling for
approximately 18 to 24 h, trained personnel
measured fat thickness, longissimus area,
marbling score (100 to 199 = Devoid; 200 to
299 = Traces; 300 to 399 = Slight; 400 to 499 =
Small; 500 to 599 = Modest; 600 to 699 =
Moderate), and USDA quality grade at the 12th-
13th rib interface on each carcass. The left strip
loin from each carcass was obtained and
fabricated into 1-in thick steaks for palatability
analyses. The 3 most anterior steaks from the
strip loin were obtained and aged 7 d. They
were vacuum-packaged and transported frozen
to the University of Florida for shear force and
sensory panel evaluation. Steaks were randomly
designated for Wamer-Bratzler shear force and
sensory panel evaluations. Peak shear force was
measured on 6 to 8 cores per steak that were 0.5
inches in diameter using a Wamer-Bratzler
shearing device, and analyzed as the average of
those values. Cooked loin steaks were cut into
0.5-in2 samples and served warm to an 8- to 10-
member trained sensory panel, and were
evaluated for juiciness, flavor intensity, panel
tenderness, and detectable amount of connective


tissue on scales from 1 to 8 (1 = extremely dry,
extremely bland, extremely tough, abundant
amount; 8 = extremely juicy, extremely intense,
extremely tender, none detected). Off flavor of
steak samples was evaluated on a scale from 1 to
6 (1 = extreme off flavor; 6 = no off flavor
detected). Sensory traits were analyzed as the
average of panelists' responses.

Since serial harvest was conducted, average
values for hot carcass weight, adjusted 12th rib
fat thickness, and marbling score (712 lb, 0.5 in,
and Small06, respectively), the average days fed
(130), and the average regression of traits on
days fed across all breed groups for these traits
were used to estimate the number of feeding
days required for each breed group to reach the
end points above (carcass weight, adjusted 12th
rib fat thickness, and marbling score). These
methods have been described in detail and used
for over 30 years (Koch et al., 1976; Wheeler et
al., 1996; 2001; 2010).

Results
In general, crossbred steers were projected to
require fewer days on feed to the different end
points (Table 1). Angus steers were projected to
require fewer days on feed to achieve fat-based
endpoints than other breed groups. Brahman-
Angus steers were also projected to require
fewer days to reach 12th rib fat thickness and
marbling score end points. As expected,
Brahman steers were projected to require the
most days to achieve any of the three end points.
Brahman-Romosinuano crosses were projected
to require similar feeding time to reach a
marbling score end point, but there appeared to
be reciprocal differences within the Brahman-
Romosinuano crosses for the other end points.

When comparing crossbred steers, breed group
was not influential on marbling score,
percentage grading Choice or Standard, Wamer-
Bratzler shear force, or sensory panel
tenderness, and connective tissue amount, but
sire and dam breeds as main effects were
detected. Angus sire breed means (Table 2)
were greater than Brahman sire breed means for
marbling score, percentage grading Choice,
sensory panel tenderness (which indicated more
tender steaks), and connective tissue amount










(lower amounts). Angus sire breed means were
lower than Brahman for Wamer-Bratzler shear
force and percentage grading Standard.
Romosinuano had a lower marbling score sire
breed mean than Angus, but did not otherwise
differ from Angus for these traits. Romosinuano
sire breed means were greater than Brahman for
sensory panel tenderness and connective tissue
amount, and were lower than Brahman for
percentage grading Standard. Results and
differences of dam breed means were very
similar to those for sire breed means (Table 2).

The heterosis estimates for Brahman-Angus for
all 7 traits were of greater magnitude than
corresponding Romosinuano estimates (Table
3). Heterosis was detected for live weight, hot
carcass weight, dressing percentage, and
longissimus area for Brahman with
Romosinuano and Angus with Romosinuano.


Brahman-Romosinuano heterosis (9.5 to 15.6%)
was also detected for 12th rib fat thickness and
yield grade. It should be noted that heterosis for
yield grade was undesirable, that is, crossbreds
had greater numerical yield grades than
straightbreds.

The estimated Brahman direct effects were to
increase live weight, hot carcass weight,
dressing percentage, and yield grade (Table 3).
Angus direct effects were to increase live
weight, hot carcass weight, 12th rib fat thickness,
longissimus area, and yield grade. The
estimated Romosinuano breed direct effects
were to lower live weight, hot carcass weight,
dressing percentage, 12th rib fat thickness, and
yield grade. Maternal breed effects were
detected for Brahman and Romosinuano for
most of these traits; however, they were of
opposite numerical sign than direct effects.


Literature Cited
Kochetal. 1976. J. Anim. Sci. 43:48.
Wheeler etal. 1996. J. Anim. Sci. 74:1023.
Wheeler et al. 2001. J. Anim. Sci. 79:1209.
Wheeler et al. 2010. J. Anim. Sci. 88:3070.















Table 1. Numbers of steers and estimates of days of feeding needed for steers in breed groups to reach these
end points: Fat thickness (0.5 in), hot carcass weight (712 lb), and marbling score (406; Small06)
End point2


Breed Group' N Fat thickness Hot carcass weight Marbling score
BB 47 163 164 223
RR 72 170 155 148
AA 38 93 140 54
RB 36 170 163 180
BR 73 113 93 179
RA 41 129 137 100
AR 77 116 96 97
BA 34 76 83 115
AB 45 115 109 126
'Breed group designation: the first and second letters in a pair indicate the breed of sire and dam,
respectively for the steers in that group B = Brahman, A = Angus, and R = Romosinuano.
2For example, Romosinuano steers would require an estimated 170 days to reach an average of 0.5 in fat
thickness.


Table 2. Sire and dam breed means for beef quality traits and palatability traits of steaks
Breed Marbling2 Choice, Standard, Shear force, Tenderness3 Connective
% % lb Tissue3
Sire breed
Brahman 359.8b 31.0b 23.1b 9.7b 5.4b 6.1b
Angus 475.2a 75.0a 5.0a 8.6a 5.8a 6.5a
Romosinuano 393.1b 46.3ab 10.0a 9.3ab 5.8a 6.5a
SE1 11.1 4.9 2.7 0.3 0.08 0.07
Dam breed
Brahman 351.5c 30.9c 29.0b 9.7b 5.5b 6.2b
Angus 480.5a 73.6a 2.0a 8.8a 5.7a 6.4ab
Romosinuano 396.1b 47.8b 7.1 9.0ab 5.8a 6.5a
SE 9.4 4.3 2.9 0.2 0.07 0.06
'Standard errors were averaged across breed groups within traits.
2Marbling score: 300 to 399 = Slight; 400 to 499 = Small; 500 to 599 =Modest.
3Tenderness and detectable amount of connective tissue measured on scales from 1 to 8: 1 = extremely tough,
abundant amount; 8 = extremely tender, none detected.
abcWithin a column and breed group, means that without a common superscript differ (P<0.05).
















Table 3. Estimates of genetic effects for carcass traits
Live Hot carcass Dressing
weight, lb weight, lb percentage
Heterosis'


Fat
thickness, in


LMA5, Yield
in2 grade


BA' 135.8
12.4%
BR 81.4
7.6%
AR 71.2
6.4%
SE4 14.1
Direct effects
B 75.0
A 72.8
R -147.7
SE 36.2
Maternal effects


102.1
15.1%
57.3
8.6%
55.6
8.1%
10.1

63.5
42.3
106.0
23.4


1.7
2.7%
0.6
1.0%
1.1
1.7%
0.3

1.6

-1.4
0.6


0.1
19.9%
0.06
15.6%
2


0.02


0.15
-0.21
0.04


B -113.5 89.5 -1.6 -0.16
A 0.06
R 136.2 100.8 1.5 0.09
SE 21.2 13.5 0.4 0.03
1Heterosis estimates are presented in units of each trait, and as % of purebred average
2Empty cells indicate estimates that did not statistically differ from 0.
3B = Brahman; A = Angus; R = Romosinuano.
4Standard errors are averages within effects within traits.
5LMA = Loin muscle area (longissimus at the 12th and 13th rib).


0.82
7.1%
0.39
3.3%
0.56
4.5%
0.186


0.67

0.39



0.50
0.217


0.4
13.6%
0.3
9.5%


0.08

0.6
0.3
-0.9
0.13

-0.7

0.5
0.09




Full Text

PAGE 1

Summary Heterosis was documented for traits that are important in the determination of value for beef carcasses in the United States. Brahman Angus carcasses had very high levels of heterosis for live weight, hot carcass weight, adjusted 12 th rib fat thickness, l ongissimus area, and USDA yield grade. Estimates of heterosis for the Criollo breed Romosinuano were lower than Brahman Angus heterosis. Brahman direct effects were to increase live weight, hot carcass weight, adjusted 12 th rib fat thickness, longissimus area, and USDA yield grade. However, generally consistent with previous reports, Brahman sire and dam breed means for marbling score, sensory panel overall tenderness, and % carcasses grading Choice were lower than Bos taurus breeds; Warner Bratzler shea r force values for Brahman were higher. Angus Romosinuano means for live weight, hot carcass weight, adjusted 12 th rib fat thickness, longissimus area, and USDA yield grade were very similar to those raised on the Great Plains (Wheeler et al., 2010). Nei ther Brahman Romosinuano nor Angus Romosinuano steers had any advantages for these traits as compared to Brahman Angus. Brahman Angus crossbred steers were projected to reach all end points (hot carcass weight, 12 th rib fat thickness, or marbling score) s ooner than most other breed groups. Introduction Since Brahman crossbred cows are so widely used by cow calf producers in the Southern United States, producer profitability is often affected by the carcass value of the resulting steers from matings to obtain crossbred Brahman cows. The reputation for in adequate carcass quality and tenderness for both Brahman purebreds and crossbreds results in lower prices paid at various places in the beef production system for calves with visible Brahman inheritance. Bos taurus Criollo breeds, such as the Romosinua no of Colombia may offer an alternative to Brahman, or a complement to Brahman, and result in adapted females that could be used in cow calf production and in steers that may perform adequately in U.S. feeding and beef production traits. Romosinuano cattl e were available for a crossbreeding project to investigate such issues at the Subtropical Agricultural Research Station in Brooksville. The objectives of this work were to estimate heterosis and breed genetic effects for traits affecting carcass value an d beef palatability traits from purebred and crossbred steers produced by Brahman, Angus, and Romosinuano cattle. Procedure Brahman, Angus, and Romosinuano bulls were mated to cows of the same breeds in all combinations. Straightbred (3 groups) and cro ssbred (6 groups; reciprocal F 1 s were considered to be distinct groups) steers were spring born in 2002, 2003, and 2004 (n = 137, 148, and 179 steers, respectively). Steers were weaned at approximately 7 mo of age in September of each year in three separa te wk. A total of 11 Brahman, 12 Angus, and 16 Romosinuano bulls sired steers with records in these data (average of 13, 14, and 10 steers per sire, respectively). Approximately one mo after weaning, in mid October of each year, steers were transported by semi trailer to the Grazinglands Research Laboratory (GRL) at El Reno, Oklahoma. After a short receiving period, they were assigned to two management groups and placed on winter Heterosis and Breed Effects for Beef Traits of Brahman Purebred and Crossbred Steers D. G. Riley 1 C. C. Chase, Jr. 1 S. W. Coleman 1 W. A. Phillips 2 M. F. Miller 3 J. C. Brooks 3 D. D. Johnson 4 and T. A. Olson 4 1 Subtropical Agricultural Research Station, USDA, ARS, Brooksville FL 2 Grazinglands Research Laboratory, USDA, ARS, El Reno, OK 3 Department of Animal and Food Sciences, Texas Tech University Lubbock, TX 4 Department of Animal Sciences, University of Florida Gainesville, FL Brahman heterosis (with two Bos taurus breeds) for important carcass traits in the U.S. beef industry was large and beneficial for most traits. Traits related to beef quality and palatability still have room for improvement in Brahman straightbreds and crosses.

PAGE 2

wheat pasture until the first week of May in the subsequent year. Each ye ar, the heaviest 10 steers from each breed group were assigned in pairs to GRL feeding pens (n = 15; 6 steers per pen). Assignments were made so that pairs of steers from each breed group were penned at least once with pairs of steers from every other bre ed group. Those steers that then remained were then assigned to larger feeding pens (4 pens in two of the project years, 6 pens in the final project year; n = 12 to 16 steers per pen). One third of the steers in each pen and breed group was randomly assig ned to each of three harvest dates in August, September, and October of each year, which corresponded to averages of 101, 129, and 157 d fed across the three project years. Steers were transported approximately 5 h to Amarillo, TX, for commercial harvest. Traits analyzed included those that affect USDA yield and quality grades, and assessments of palatability. Hot carcass weight was obtained from harvest facility records, and was used in conjunction with final live weight (shrunk weight prior to transpor tation) to estimate dressing percentage. After chilling for approximately 18 to 24 h, trained personnel measured fat thickness, longissimus area, marbling score (100 to 199 = Devoid; 200 to 299 = Traces; 300 to 399 = Slight; 400 to 499 = Small; 500 to 599 = Modest; 600 to 699 = Moderate), and USDA quality grade at the 12 th 13 th rib interface on each carcass. The left strip loin from each carcass was obtained and fabricated into 1 in thick steaks for palatability analyses. The 3 most anterior steaks from the strip loin were obtained and aged 7 d. They were vacuum packaged and transported frozen to the University of Florida for shear force and sensory panel evaluation. Steaks were randomly designated for Warner Bratzler shear force and sensory panel evalu ations. Peak shear force was measured on 6 to 8 cores per steak that were 0.5 inches in diameter using a Warner Bratzler shearing device, and analyzed as the average of those values. Cooked loin steaks were cut into 0.5 in 2 samples and served warm to an 8 to 10 member trained sensory panel, and were evaluated for juiciness, flavor intensity, panel tenderness, and detectable amount of connective tissue on scales from 1 to 8 (1 = extremely dry, extremely bland, extremely tough, abundant amount; 8 = extreme ly juicy, extremely intense, extremely tender, none detected). Off flavor of steak samples was evaluated on a scale from 1 to 6 (1 = extreme off flavor; 6 = no off flavor detected). Sensory traits were analyzed as the Si nce serial harvest was conducted, average values for hot carcass weight, adjusted 12 th rib fat thickness, and marbling score (712 lb, 0.5 in, and Small 06 respectively), the average days fed (130), and the average regression of traits on days fed across all breed groups for these traits were used to estimate the number of feeding days required for each breed group to reach the end points above (carcass weight, adjusted 12 th rib fat thickness, and marbling score). These methods have been described in deta il and used for over 30 years (Koch et al., 1976; Wheeler et al., 1996; 2001; 2010). Results In general, crossbred steers were projected to require fewer days on feed to the different end points (Table 1). Angus steers were projected to require fewer day s on feed to achieve fat based endpoints than other breed groups. Brahman Angus steers were also projected to require fewer days to reach 12 th rib fat thickness and marbling score end points. As expected, Brahman steers were projected to require the most days to achieve any of the three end points. Brahman Romosinuano crosses were projected to require similar feeding time to reach a marbling score end point, but there appeared to be reciprocal differences within the Brahman Romosinuano crosses for the oth er end points. When comparing crossbred steers, breed group was not influential on marbling score, percentage grading Choice or Standard, Warner Bratzler shear force, or sensory panel tenderness, and connective tissue amount, but sire and dam breeds as main effects were detected. Angus sire breed means (Table 2) were greater than Brahman sire breed means for marbling score, percentage grading Choice, sensory panel tenderness (which indicated more tender steaks), and connective tissue amount

PAGE 3

(lower amoun ts). Angus sire breed means were lower than Brahman for Warner Bratzler shear force and percentage grading Standard. Romosinuano had a lower marbling score sire breed mean than Angus, but did not otherwise differ from Angus for these traits. Romosinuano sire breed means were greater than Brahman for sensory panel tenderness and connective tissue amount, and were lower than Brahman for percentage grading Standard. Results and differences of dam breed means were very similar to those for sire breed means ( Table 2). The heterosis estimates for Brahman Angus for all 7 traits were of greater magnitude than corresponding Romosinuano estimates (Table 3). Heterosis was detected for live weight, hot carcass weight, dressing percentage, and longissimus area for Brahman with Romosinuano and Angus with Romosinuano. Brahman Romosinuano heterosis (9.5 to 15.6%) was also detected for 12 th rib fat thickness and yield grade. It should be noted that heterosis for yield grade was undesirable, that is, crossbreds had greater numeri cal yield grades than straightbreds. The estimated Brahman direct effects were to increase live weight, hot carcass weight, dressing percentage, and yield grade (Table 3). Angus direct effects were to increase live weight, hot carcass weight, 12 th rib fa t thickness, longissimus area, and yield grade. The estimated Romosinuano breed direct effects were to lower live weight, hot carcass weight, dressing percentage, 12 th rib fat thickness, and yield grade. Maternal breed effects were detected for Brahman a nd Romosinuano for most of these traits; however, they were of opposite numerical sign than direct effects. Literature Cited Koch et al. 1976. J. Anim. Sci. 43:48. Wheeler et al. 1996. J. Anim. Sci. 74:1023. Wheeler et al. 2001. J. Anim. Sci. 79:1209. Wheeler et al. 2010. J. Anim. Sci. 88:3070.

PAGE 4

Table 1. Numbers of steers and estimates of days of feeding needed for steers in breed groups to reach these end points: Fat thickness (0.5 in), hot carcass weight (712 lb), an d marbling score (406; Small 06 ) End point 2 Breed Group 1 N Fat thickness Hot carcass weight Marbling score BB 47 163 164 223 RR 72 170 155 148 AA 38 93 140 54 RB 36 170 163 180 BR 73 113 93 179 RA 41 129 137 100 AR 77 116 96 97 BA 34 76 83 115 AB 45 115 109 126 1 Breed group designation: the first and second letters in a pair indicate the breed of sire and dam, respectively for the steers in that group B = Brahman, A = Angus, and R = Romosinuano. 2 For example, Romosinuano steers would require an estimated 170 days to reach an average of 0.5 in fat thickness. Table 2. Sire and dam breed means for beef quality traits and palatability traits of steaks Breed Marbling 2 Choice, % Standard, % Shear force, lb Tenderness 3 Connective Tissue 3 Sire breed Brahman 359.8 b 31.0 b 23.1 b 9.7 b 5.4 b 6.1 b Angus 475.2 a 75.0 a 5.0 a 8.6 a 5.8 a 6.5 a Romosinuano 393.1 b 46.3 ab 10.0 a 9.3 ab 5.8 a 6.5 a SE 1 11.1 4.9 2.7 0.3 0.08 0.07 Dam breed Brahman 351.5 c 30.9 c 29.0 b 9.7 b 5.5 b 6.2 b Angus 480.5 a 73.6 a 2.0 a 8.8 a 5.7 a 6.4 ab Romosinuano 396.1 b 47.8 b 7.1 a 9.0 ab 5.8 a 6.5 a SE 9.4 4.3 2.9 0.2 0.07 0.06 1 Standard errors were averaged across breed groups within traits. 2 Marbling score: 300 to 399 = Slight; 400 to 499 = Small; 500 to 599 = Modest. 3 Tenderness and detectable amount of connective tissue measured on scales from 1 to 8: 1 = extremely tough, abundant amount; 8 = extremely tender, none detected. a,b,c Within a column and breed group, means that without a common superscript differ ( P <0.05).

PAGE 5

Table 3. Estimates of genetic effects for carcass traits Live Hot carcass Dressing Fat LMA 5 Yield weight, lb weight, lb percentage thickness, in in 2 grade Heterosis 1 BA 3 135.8 12.4% 102.1 15.1% 1.7 2.7% 0.1 19.9% 0.82 7.1% 0.4 13.6% BR 81.4 7.6% 57.3 8.6% 0.6 1.0% 0.06 15.6% 0.39 3.3% 0.3 9.5% AR 71.2 6.4% 55.6 8.1% 1.1 1.7% 2 0.56 4.5% SE 4 14.1 10.1 0.3 0.02 0.186 0.08 Direct effects B 75.0 63.5 1.6 0.6 A 72.8 42.3 0.15 0.67 0.3 R 147.7 106.0 1.4 0.21 0.9 SE 36.2 23.4 0.6 0.04 0.39 0.13 Maternal effects B 113.5 89.5 1.6 0.16 0.7 A 0.06 R 136.2 100.8 1.5 0.09 0.50 0.5 SE 21.2 13.5 0.4 0.03 0.217 0.09 1 Heterosis estimates are presented in units of each trait, and as % of purebred average 2 Empty cells indicate estimates that did not statistically differ from 0. 3 B = Brahman; A = Angus; R = Romosinuano. 4 Standard errors are averages within effects within traits. 5 LMA = Loin muscle area (longissimus at the 12 th and 13 th rib).