EFFECTS OF FEEDER-CATTLE FRAME SIZE AND MUSCLE THICKNESS

ON SUBSEQUENT GROWTH AND CARCASS DEVELOPMENT. II.

ABSOLUTE GROWTH AND ASSOCIATED CHANGES
IN CARCASS COMPOSITIONI~

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J. D. T a t u m , H. G. D o l e z a l 3 , F. L. Williams, Jr. 4 ,
R. A. Bowling s and R. E. Taylor

Colorado State University 6,

Fort Collins 80523

ABSTRACT
Data from a serial slaughter of 324 steers were used to determine the effects of feeder-cattle
frame size (Large, L; Medium, M; Small, S) and muscle thickness (No. 1, No. 2, No. 3) on absolute
growth and carcass development. Yearling feeder steers representing nine frame-size • muscle-
thickness subclasses were backgrounded 28 d; subsamples of steers (n = 6) from each subclass were
slaughtered at 28~d intervals (O, 28, 56, 84, 112 or 140 d) during a 140-d finishing period. One side
from each carcass was dissected into muscle, bone and fat. Absolute growth during finishing
was linear. Allometric growth coefficients for muscle, bone and fat were similar for all nine sub-
classes. Frame size was related (P<.01) to differences in absolute growth rate (L>S), compositional
maturity at a common carcass weight (S>M>L) and slaughter weight at a constant carcass-fat
percentage (L>M>S). Muscle thickness did not influence absolute growth rate, but was associated
with differences (P<.01) in carcass muscle-to-bone ratio at a common bone weight and in muscle
percentage when carcass fatness was statistically standardized. The effects of muscle thickness on
carcass composition were most pronounced within the large-framed group. Relationships of these
results to the USDA feeder-carrie grading principles are discussed.
(Key Words: Beef Cattle, Carcass Grading, Frame Size, Body Measurements, Growth, Carcass
Composition.)

Introduction
The intrinsic value of a feeder animal is a
function of its genetic potential to grow and
develop rapidly and efficiently and to produce
a carcass composed of optimal proportions o f
muscle, bone and fat at a preferred market
l Sci. Set. Paper No. 2989 published with the ap-
proval of the Colorado State Univ. Agr. Exp. Sta. This
research was supported by the USDA Agr. Marketing
Serv., and the Colorado Dept. of Agr. under the
Federal-State Marketing Improvement Program.
=The authors express appreciation to Mr. W. E.
(Ned) Tyler for his contirbutions in design and direc-
tion of the experiment and to R. P. Clayton, D. A.
Daley, B. K. Klein, W. R. Lloyd, R. K. Miller, C. W.
Moran, M. D. Self, T. D. Stromberg, M. L. Wittier and
D. W. Woodburn for their assistance in data collection.
SPresent address: Dept. of Anita. Sci., Oklahoma
State Univ., Stillwater, OK 74078.
4 Livestock Division, Agr. Marketing Serv., USDA.
s Monfort of Colorado, inc. Greeley, CO.
s Dept. of Anita. Sci.
Received March 18, 1985.
Accepted August 23, 1985.
121
weight. Under conventional marketing circum-
stances, relative-value differences among feeder-
cattle are established using phenotypic indi-
cators of prospective feedlot performance and
carcass merit. The current USDA market
classification system for feeder cattle (USDA,
1979) employs two such i n d i c a t o r s - f r a m e size
and thickness.
The frame size-thickness grading concept is
based on the rationale that: (1) immature
skeletal size (frame size) is indicative of an
animal's potential mature size and associated
effects on growth rate and the weight at which
the animal will attain a specified level of fatness,
and (2) visually discernable differences among
feeder cattle in size, shape and thickness of the
musculature (muscle thickness) reflect inherent
variation in muscularity and its influence on
carcass yield grade and muscle-to-bone ratio.
Implicit in the USDA feeder-grading prin-
ciples is a connection between feeder-cattle size
and shape and the relative development of the
major carcass tissues (muscle, bone and fat) in
the finished slaughter animal. Previous research,
J. Anim. Sci. 1986. 62:121-131
122 TATUM ET AL.

reviewed by Berg and Butterfield (1976) and

Berg and Walters (1983), has shown that the
relative proportions of muscle, bone and fat in
the bovine carcass change continually during
the growing-finishing period and that these

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developmental changes are influenced by
various genetic and environmental effects. The
premise advanced in the feeder-cattle grading
standards is that evaluations of frame size and
muscle thickness identify two of the major
genetic sources of variation in carcass develop-
ment-mature size and muscularity.
Previous studies (Lush, 1932; Knox and
Koger, 1946; Stonaker et al., 1952; Kidwell and
McCormick, 1956; Kauffman et aL, 1970;
Brungardt, 1972; Crouse et al., 1974; Adams
and Smith, 1979; Butts et al., 1980a,b; Long et
al., 1980; Tatum et al., 1982) have characterized
differences in value-determining carcass traits
that result when cattle of diverse body types
(both within and among breeds) are compared
at various static slaugher endpoints. However,
there is a dearth of information concerning the
effects of feeder-cattle frame size and muscle
thickness on growth and development of the
constituent tissues of the carcass during fin-
ishing. Such information is essential for de-
veloping a comprehensive understanding of the
link between size and muscularity of the feeder
calf and value-determining characteristics of
the finished slaughter animal. The objective of
this study was to characterize the effects of
feeder-cattle frame size and muscle thickness on
absolute growth and associated changes in
carcass composition during a 140-d finishing
period.

Materials and Methods

Design, The experimental design is presented oo
Ox
in table 1. Data presented in this segment of the
study were obtained from 324 steers, rep-
Ox
resenting nine frame-size x muscle-thickness (F
Ox
x M) subclasses, slaughtered serially at 28-d v~

intervals (0, 28, 56, 84, 112 or 140 d) during a 0~

140-d finishing period. Two identical replicates

of the experiment were conducted in successive
years. u
Animals. A detailed description of the
sample of steers obtained for the experiment
was provided by Tatum et al. (1986). In brief,
yearling feeder steers were selected specifically
to represent nine F • M subclasses. The cattle
were similar in age, but were quite diverse with
regard to genotype. Twenty different breeds
. zzz: zzz zzz .
 FEEDER-CATTLE FRAME SIZE AND MUSCLE THICKNESS 123

(both beef and dairy) were represented in the

study. ~eq
The feeder steers were assigned to their
respective subclasses by a panel of five eval-
uators using visual assessments of (1) skeletal

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size in relation to age (USDA, 1979) and (2)
thickness of the musculature in relation to oo0
o0 O
a~
skeletal size (USDA, 1976). An objective basis
for the subjective methods used to allocate the E
cattle to their respective subclasses was pro-
vided in a companion report (Tatum et al., oo0
1986). In addition to classifying the cattle into
appropriate frame-size and muscle-thickness
categories, the evaluation panel also assigned a
condition score (1 = extremely thin; 9 = ex- eL
tremely fleshy) to each steer. oo0
Acclimation and Finishing. After the cattle
had been processed, each F • M subclass was
z
divided randomly into six, equally-sized slaugh-
ter groups. Steers from all nine subclasses with
z
the same slaughter-group designation were
penned together.
o
The steers were started on a 30% concentrate
diet, and a 28-d acclimation period was initiated
to facilitate correction of minor stress and
~o0
disease problems that normally occur following .o
transport and processing. The concentrate level Ox
of the diet was increased gradually from 30 to
60% during the acclimation period (table 2)..
On the day before the start of the finishing oo0

period, cattle assigned to the initial slaughter

group were segregated for slaughter. The
following day, remaining steers were weighed z o

(24-h shrunk weight) and placed on feed. At

the onset of the finishing period, the steers e~ eq g~
O
were receiving a 60% concentrate diet ad
libitum. The concentrate level of the diet was
increased to 85% during the first 28 d on feed
and remained at that level for the duration of
the experiment 9 Diet ingredients and the time t~

schedule for each diet are presented in table 2.

Slaughter and Carcass Dissection. All animals O 0 0 ~ Ox r,~ O~

were weighed and slaughtered following a 24-h

fast. The cattle were slaughtered either a t the
Colorado State University abattoir or at a
small, commercial packing facility9 Slaughter
methods employed at the two locations essen-
tially were identical, Immediately following o,q.r
A c
slaughter, hot-carcass weight was recorded and
the carcasses were chilled at approximately 2 12.
Seventy-two hours postmortem, the left side dd
of each carcass was separated into primal and ~ex=
subprimal cuts and each cut was physically ~a ~0
separated into muscle, fat and bone (bone plus
cartilage and major tendons) using procedures t~
124 TATUM ET AL.
described by Abraham et al. (1980). Individual
weights were recorded for each cut and its
components. Carcass side weight was expressed
as the aggregate of all component weights.
Component weights corresponding to a par-
titular tissue-type (muscle, bone or fat) were
aggregated to provide the weights of muscle,
bone and fat in the entire side. For purposes of
analysis, the aggregates were doubled to corres-
pond to the weight of the entire animal.
Statistical Methods. Data collected in this
study were cross-sectional (Tanner, 1951). In
cross-sectional growth studies, each individual is
measured only once and the same measurements
are obtained for different groups of individuals
within each subclass (treatment group) at
successive sampling times to estimate a subclass
mean-growth response. This type of experi-
mental design is based on the assumption that
individuals within each sampling group are truly
representative of their respective subclass
(Fitzhugh, 1976). Frequently, however, varia-
tion in stage of development among animals
within a subclass, at the onset of the experi-
ment, causes some degree of overlap in size or
development of animals in adjacent sampling
groups. This results in an irregularly shaped
growth response, particularly when the number
of individuals measured at each sampling time is
small. To avoid this problem, the data were
adjusted to the mean initial weight and condition
score within each F x M subclass; previous
research has shown that among cattle of a
similar mature size, weight and condition are
closely associated with developmental status
(Elsley, 1976; Butts et al., 198On,b). Within-
subclass regression coefficients used for adjust-
ments were derived using a least-squares model
that included sampling time (days fed) and the
two covariates (initial weight and condition
score). Adjustment of the data on a within-
subclass basis reduced the variability among
individuals of the same frame size and muscle
thickness, but maintained among-subclass vaxi-
ation in initial size and development. Adjusted
data were used as input for all subsequent
analyses.
Preliminary plots of the data revealed that
the time curves of growth for all nine subclasses
were approximately linear. Due to the nature of
the weight-time plots and because the time
interval of experimental growth was relatively
short, polynomial equations were used to
represent absolute growth (Finney, 1978). Use
of polynomial equations not only provided
mathematical representation of growth, but
also allowed statistical estimation of instan-
taneous absolute growth rates (Brody, 1945).
In first- and second-degree polynomial equa-
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tions, the first derivative with respect to time
(dy/dx, where y = live weight and x = time) is
an estimate of the average absolute growth rate
(AGR) for a particular subclass at a given time
during the 140-d period.
Linear and quadratic regression equations
were calculated on a within-subclass basis.
Statistical tests for the quadratic term in the
equations revealed that the weight-time
relationships for the nine subclasses were
linear.
Once the nature of weight-time relationships
had been established, an analysis of variance
was performed using a least-squares model
(model I) that included fixed effects of year
(YR), frame size (F), muscle thickness, (M), the
frame-size • muscle-thickness interaction (F •
M) and a linear regression (b) on sampling time
(days fed). In addition, b x F, b • M and b x F
• M interactions were partitioned to test for
homogeneity of subclass regressions. In the
context of this analysis, heterogeneity (P<.05)
of subclass regression coefficients reflected
significant differences among the subclasses in
average AGR during the 140-d period.
Effects of frame size and muscle thickness
on carcass developmental changes during
finishing were examined using the bivariate
allometric equation (Y = ~x~3). The statistical
approach used for this purpose was consistent
with methodology advocated by Berg et al.
(1978) for testing group differences in carcass
developmental patterns. Dependent (Y) and in-
dependent (X) variates were chosen specifically
to address two important aspects of carcass
development: (1) changes in percentages of
muscle, bone and fat in the carcass (Y = muscle,
bone or fat weight, X = carcass weight) and (2)
changes in the ratio of muscle and bone in the
carcass (Y = muscle weight, X = bone weight).
To facilitate analysis using least-squares tech-
niques, all variates were transformed into
logarithms, thereby transforming the aUometric
equation from a parabola into a linear function
(log Y = loga + 3log X). Additional terms (YR,
F , M , F • M, 3 x F, 3 • M, 3 x F • M) were
included in the least-squares model (model 2)
to permit statistical comparisons of subclass
means for Y at a constant value of X and to test
for homogeneity of subclass allometric coeffi-
cients (#).
 FEEDER-CATTLE FRAME SIZE AND MUSCLE THICKNESS
Least-squares analyses were performed using
the SPSS MANOVA program (Hull and Nie,
1981). Differences among means for static
comparisons were tested using Tukey's w
procedure (Steel and Torrie, 1960). Com-
parisons of subclass regression coefficients were
tested for significance using dummy variable
regression techniques described by Weisberg
(1980).
Results
Absolute Growth. Subclass regressions illus-
trating increases in live weight with respect to
time (absolute growth) are presented in figure
1. The analysis of variance corresponding to the
data displayed in the figure is provided in table
3.
Linear regression equations provided the
most effective mathematical representation of
absolute growth for all nine F X M subclasses
(figure 1). The apparent linearity of the weight-
b x F regceslg'(~ coefficients T~me.constant (70~m~S
Subclass b Su~ckss
Large t ,35 a L1
Medium 1,25 ab L2
1,13 b
L3
M1
M2
M3
51
$2
53
0 TO
Days
Figure 1. Weight-time relationships during the
140-d finishing period. Regression coefficients and
70-d means that do not have a common superscript
a r e different (P<.05).
125
time relationships implied that AGR remained
constant for the duration of the experiment.
Actually, the AGR for each subclass declined
very gradually during the finishing period.
However, due to the subtlety of the change in
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growth rate and the brevity of the experimental
growth interval, the slight curvilinearity of the
mean growth response for each subclass was
statistically obscure.
Significant mean squares for F, M and F • M
in table 3 were indicative of subclass differences
in live weight (i.e., position of the regression
line relative to the y axis; figure 1) at a constant
feeding time of 70 d. These differences resulted
from variation in body size and development
established prior to finishing, together with
weight variation arising from subclass differences
in growth rate during the first half of the
finishing period.
Tests for differences in AGR among the
subclasses were provided by the b • F, b • M
and b x F • M interaction terms in table 3.
Frame size was a highly significant source of
variation in AGR (table 3); b • F subclass
regressions are provided in figure 1. Large-
Mean.kQ
framed steers grew at the fastest rate (1.35
5483 a
513.7 b kg/d) during finishing; small-framed cattle grew
478.9 r slowest (1.13 kg/d). Growth rate for medium-
475,8 c
4~;',4 d
framed steers was intermediate (1.25 kg/d) and
428.7 9 did not differ (P>.05) from the AGR for steers
436.1 9 in the other two frame groups. Muscle thickness
432.4 e
388.0 ( did not influence AGR during finishing, as
evidenced by the lack of significance for the b
x M term in table 3.
Carcass Cornposition.'Developmental changes
in carcass composition during finishing were
characterized by the bivariate allometric
relationships in table 4. The allometric coeffi-
cients (/3) represent the ratio of the specific
(percentage) growth rates of the corresponding
dependent (Y) and independent (X) variates
(Seebeck, 1968). An allometric coefficient
equal to 1 indicated that the proportion of Y
relative to X remained unchanged during the
experimental period. Coefficients different
(P<.05) from 1 signified an increase (if/3> 1) or
decrease (if /3<1) in the proportion of Y as
the weight of X increased.
Examination of the allometric relationships
! in table 4 showed that fat grew at a much faster
140
rate, relative to carcass weight gain, than did
either muscle or bone (/3 = 2.03, .76 and .58 for
fat, muscle and bone, respectively). In addition,
the log-log relationship between muscle and
bone was isometric (/3 = 1), indicating that
126 TATUM ET AL.

TABLE 3. ANALYSIS OF VARIANCE FOR LIVE WEIGHT

Source of
vaz/~tion Degrees of freedom Mean squaresb

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Year 1 159
Frame size (F) 2 251,359"*
Muscle thickness (M) 2 86,062* *
F XM 4 2,791"*
Days on feed (b)t 1 1,142,015'*
b• F 2 2,921"*
b XM 2 703
bX FX M 4 1,064
Residual 305 501

aLinear regression of live weight on days fed.

bMean squares calculated using model 1.
**P<.01.

carcass muscle-to-bone ratio r e m a i n e d relatively Tests for h o m o g e n e i t y of subclass allometric

c o n s t a n t during finishing. These data suggested
t h a t at the onset o f t h e e x p e r i m e n t , the steers
already had attained m u c h o f their potential
skeletal and muscular d e v e l o p m e n t and were
progressing into t h e phase o f g r o w t h t h a t is
characterized by rapid d e p o s i t i o n o f carcass fat.
Due to the d i s p r o p o r t i o n a t e increase in the fat
c o m p o n e n t o f t h e carcass during the finishing
period, percentages o f muscle and b o n e de-
creased (~< 1) and t h e percentage of carcass fat
increased 03>1) as carcass weight increased
(table 4-).
coefficients s h o w e d that frame size and muscle
thickness had no effect on p r o p o r t i o n a t e
g r o w t h o f muscle, b o n e and fat. Individual
subclass coefficients were h o m o g e n e o u s for all
traits analyzed, indicating that the previously
described changes in carcass c o m p o s i t i o n
o c c u r r e d at similar rates for cattle in all nine F
X M subclasses.
T h e significant fixed effects in table r reflect
weight-constant differences in carcass composi-
tion. Means corresponding to these effects are
presented in table 5. Least-squares means

TABLE 4. ESTIMATES OF SELECTED ALLOMETRIC RELATIONSHIPSa

AMONG MUSCLE, BONE, FAT AND CARCASS WEIGHT

Significance of
Dependent Independent fixed effect* (model 2) c
variare, variate, Allometrlc Residual
log Y log X coefficient, a SEB t-test b F M F XM CV, %d

Muscle Carcass wt .76 .015 0<1 ** ** ** 1.95

Bone Carcass wt .58 .020 a<l ** ** * 6.33
Fat Carcass wt 2.03 .047 a>l ** ** NS e 2.72
Muscle Bone wt .98 .038 a=l NSe ** ** 3.46

aAllometric equation: Y = ~x~.

bResult* of a t-test, Ho:J$ = 1.
Clndividual subclass regressions were not different (P>.05). Significance of the F-test for each fixed effect
is presented: NS = not significant.
dResiduai coefficient of variation, corresponding to each dependent variate, is the square root of the error
mean square multiplied by 100.
eNot significant.
*P<.05.
**P<.01.
 FEEDER-CATTLE FRAME SIZE AND MUSCLE THICKNESS 127

TABLE 5. MEANSa FOR PHYSICAL CARCASS COMPOSITION AND MUSCLE-TO-BONERATIO

ADJUSTED TO THE GEOMETRIC MEANb OF CARCASS WEIGHT
AND BONE WEIGHT, RESPECTIVELY

Constant carcass wt Constant bone wt

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Separable Separable Separable Muscle-to-
Effect muscle, % bone, % fat, % bone ratio

Frame size (F)

Large 64.4 c 18,0 c 17.6 e 3.68
Medium 61.0 d 16.3 d 22.7 d 3,73
Small 58.8 e 15.5 e 25.7 c 3.72
Muscle thickness (M)
No. I 63.4 c 16.4 d 20.2 d 3.93c
No. 2 60.8 d 16.3 d 22.9 c 3.73 d
No. 3 60.5 d 17.0 c 22.5 c 3.48 e
FXM
Large No. 1 66.7 c 17.5 d 3.99 c
Large No. 2 63.8 d 17.4 d 3,76 def
Large No. 3 62.5 d 19.0 c 3.32g
Medium No. 1 63.0 d 16.1efg 3.94 cd
Medium No. 2 59.9e 16.2 ef 3.70 ef
Medium No. 3 60.2 e 16.6 e 3.57 f
Small No, 1 60,0 e 15.4g 3.86cde
Small No. 2 58.2 e 15.3g 3.74 def
Small No. 3 58.2 e 15.6fg 3.57 f

aTabular values correspond to the least-squares means obtained from the allometric analyses presented in
table 4. The logarithms of each dependent varlate, adjusted by allometrie regression to the geometric mean of
the corresponding independent variate, have been converted into percentages or ratios for tabular presentation
only. All tests of significance were based on results of analyses using model 2 (table 4).
bGeometric means: carcass wt = 257 kg; bone wt = 42.4 kg.
c'd'e'f'gMeans, in the same column within an effect, that do not have a common superscript letter are differ-
ent (P<.05).

derived from the allometric analyses (i.e.,
logarithms of each dependent variate adjusted
via aUometric regression to the geometric mean
of the respective independent variate) were
transformed into percentages or ratios to
simplify interpretation and discussion. Group
differences in carcass fatness were examined
using means for percentage of carcass fat, while
contrasts of the relative proportions of muscle
and bone, independent of differences in fatness,
were evaluated by comparing means for muscle-
to-bone ratio (Berg and Butterfield, 1966).
Each frame-size group was statistically
distinct with regard to percentages of muscle,
bone and fat at a constant carcass weight (table
5). Percentages of muscle and bone in the
carcass were highest for large-framed cattle,
intermediate for medium-framed steers and
lowest for cattle in the small-framed group.
Frame groups ranked in the reverse order for
percentages of carcass fat. The rankings of the
frame groups for percentages of muscle, bone
and fat were particularly informative when the
relative maturing rates of the tissues (Berg and
Butterfield, 1968) were considered. Reverse
rankings for bone, the earliest-maturing tissue,
and fat, the latest-maturing tissue, were indica-
tive of group differences in compositional
maturity. The directional aspect of the rankings
suggested that frame size was inversely related
to compositional maturity ( S > M > L ) at a
constant carcass weight.
Adjustment of the data to a c o m m o n carcass
fatness (table 6) eliminated differences in
carcass composition among the frame groups,
demonstrating that the primary effect of frame
size on the relative proportions of muscle, bone
and fat was exerted via a relationship to the
fattening process. The essence o f this relation-
ship became more apparent when the fat-
constant live weights of the frame groups were
considered. Data in table 6 showed that large-,
medium- and small-framed steers attained the
mean percentage of carcass fat (approximately
22%) at markedly different weights (528,
451 and 407 kg, respectively). Since the relative
128 TATUM ET AL.

rates o f fat deposition for the three frame
groups were shown to be similar, variation in
fat-constant live weights r e f l e c t e d group differ-
ences in the weight at which the fattening
process was initiated ( M u k h o t y and Berg,
Within t h e large-framed group, No. 3 steers
had a higher (P<.05) percentage o f separable
b o n e than cattle in either of the o t h e r two
muscle-thickness classes (table 5). This differ-
ence in b o n e percentage corresponded to the

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1971).
Muscle-thickness classification was associated
with significant differences in muscle and b o n e
percentages at a constant carcass weight and in
muscle-to-bone ratio at a c o m m o n b o n e weight
(table 4). F o r these traits, however, the effects
of muscle thickness were d e p e n d e n t u p o n
frame-size classification (the interaction be-
t w e e n F and M was significant; table 4).
Within each frame-size group, greater muscle
thickness was associated with higher muscle-
t o - b o n e ratios, and contrasts b e t w e e n e x t r e m e
muscle-thickness classes (No. 1 vs No. 3) were
significant (table 5). However, consistent
differences in muscle-to-bone ratio b e t w e e n
adjacent muscle-thickness classes (No. 1 vs No.
2, No. 2 vs No. 3) were observed o n l y a m o n g
large-framed cattle.
distinct inferiority of the large No. 3 steers in
carcass muscle-to-bone ratio. Additionally,
within the large- and m e d i u m - f r a m e d groups,
steers with No. 1 muscle thickness had a higher
(P<.05) percentage of muscle in their carcasses
than did either No. 2 or No. 3 cattle. Un-
questionably, this was due, in part, to the lower
percentage o f carcass fat for the No. 1 cattle
(table 5); however, since No. 1 steers in these
frame groups also had superior muscle-to-bone
ratios, a p o r t i o n o f their advantage in muscle
yield was attributable to superior muscularity.
The m a g n i t u d e o f t h e differences in muscle
yields corresponding to muscle-thickness classi-
fication, and exclusive o f variation in fatness,
was e x a m i n e d directly by comparing the
fat-corrected means provided in table 6. Ranges
for muscle percentage (% for No. 1 minus % for

TABLE 6. MEANS FOR LIVE WEIGHT AND PERCENTAGES a OF SEPARABLE MUSCLE

AND BONE ADJ USTED TO A CONSTANT CARCASS-FAT PERCENTAGE b

Fat-constant means c
Live Separable Separable
Effect weight, kg muscle, % bone, %

Frame size (F) ** NS NS

Large 528.0 d 61.5 16.5
Medium 451.0 e 61.5 16.5
Small 406.7 f 61.5 16.5
Muscle thickness (M) ** ** **
NO. 1 485.1 d 62.2 d 15.8 f
No. 2 454.8 e 61.5 e 16.5 e
No. 3 445.8 e 60.7 f 17.3 d
F•
Large No. 1 62.4 d 15.6 h
Large No. 2 61.7efg 16.3efg
Large No. 3 60.1 h 17.9 d
Medium No. 1 62.2 de 15.8g h
Medium No. 2 61.3fg 16.7 ef
Medium No. 3 61.og 17.0 e
Small No. 1 61.9 def 16.1fg h
Small No. 2 61.sefg 16.sefg
Small No. 3 61.0g 17.0 e

aSeparable muscle and bone are expressed as percentages of carcass weight.

bMeans were adjusted using analysis of covariance techniques to the overall mean percentage of carcass fat
(21.98%).
CSignificance of the F-test for each fixed effect is presented; NS = not significant.
d'e'f'g'hMeans, in the same column within an effect, that do not have a common superscript letter are differ-
ent (P<.05).
**P<.01.
 FEEDER-CATTLE FRAME SIZE AND MUSCLE THICKNESS
No. 3) were 2.3, 1.2 and .9 within the large-,
medium- and small-framed groups, respectively.
Means for live weight in table 6 show that
No. 1 cattle attained the mean percentage of
carcass fat at significantly heavier slaughter
weights than the other two thickness groups,
which explains the lower percentage of fat for
the No. 1 steers at a constant carcass weight
(table 5). Slaughter weights for No. 2 and No. 3
cattle at a constant fat percentage were similar.
As discussed in the first report in this series
(Tatum et al., 1986), muscling differences in
the present study tended to reflect breed
variation in muscularity. Cattle in the No. 1
muscle-thickness class were steers of beef
breeding which were perceived to be superior in
muscularity. Steers classified as No. 2 also
exhibited a predominance of beef breeding but
were considered average in muscle expression.
The No. 3 class consisted of steers exhibiting
strong dairy or Longhorn-breed influence plus a
few steers of beef breeding which were below
average in muscling. Significant differences in
carcass muscle-to-bone ratio generally con-
trasted "muscular," beef-type steers with cattle
exhibiting a predominance of dairy or Long-
horn breeding.
Discussion
Since 1979, USDA grades for thrifty feeder
cattle have been determined using separate
subjective evaluations of frame size and muscle
thickness (USDA, 1979). The conceptual basis
for this grading system is predicated upon
fundamental principles of bovine growth and
development. Existing evidence suggests that
mature size and muscularity are two major
sources of genetic variation in growth and
carcass composition (Berg and Butterfield,
1966; Berg, 1968; Berg and Butterfield, 1976;
Elsley, 1976; Berg and Waiters, 1983).
Previous research has shown that mature size
exerts an influence on bovine growth and
carcass composition via genetic relationships to
growth rate, maturing rate and weight at the
onset of fattening (Fitzhugh and Taylor, 1971;
Berg and Butterfield, 1976; Brown et al., 1983;
Mclnerney, 1984). In that connection, mature
size bears a strong, positive relationship to
absolute growth rate (Brinks et al., 1964;
Cartwright, 1970), and is negatively correlated
with maturing rate (Taylor, 1965). Further-
more, mature size appears to be positively
associated with the weight at which the fattening
129
phase of growth is initiated (Berg and Butter-
field, 1976). As a result, cattle of large potential
mature size (both among and within breeds)
normally grow faster, attain a given degree of
maturity at older ages and begin to fatten at
Downloaded from https://academic.oup.com/jas/article-abstract/62/1/121/4658455 by Goteborgs Universitet user on 26 November 2019
heavier weights than their smaller contem-
poraries. Thus, when cattle of diverse mature
sizes (large vs small) are compared at-similar
slaughter weights (Koch et al., 1976, 1979,
1982), larger genotypes tend to be younger
(because of their faster growth rates), less
mature (because of their slower maturing rates
and younger ages) and leaner (because of
their tendency to fatten at relatively heavy
weights). Conversely, when similar comparisons
are made at comparable levels of fatness
(Stonaker et al., 1952; Koch et al., 1976, 1979,
1982; Butts et al., 1980a,b), differences in
carcass composition are reduced, but larger
cattle tend to be older (because of their slower
maturing rates) and heavier (due to their
older ages, faster growth rates and tendency to
fatten at heavy weights).
Apart from compositional differences which
are linked to genetic variation in mature size
and its influence on carcass fatness, there are
economically important differences in yield of
muscle which are attributable to inherent
variation in muscle to bone ratio (Berg, 1968;
Kauffman et al., 1970; Berg and Butterfield,
1976). Previous studies have documented
that genetic differences in muscle-to-bone ratio
are expressed early postnatally (Berg et al.,
1978) and that such differences are associated
with visually discernible variation in live-animal
shape (Kauffman et al., 1970, 1973). In general,
existing evidence suggests that the largest
differences in muscle-to-bone ratio are found
between the conventional beef cattle breeds
and the dairy cattle breeds (Hankins et al.,
1943; Carroll et al., 1964; Cole et al., 1964;
Berg, 1968; Broadbent et al., 1976).
Collectively, the foregoing concepts form
the basis for using frame size and muscle
thickness as criteria for market classification of
feeder cattle. In the official U.S. standards for
grades of feeder cattle, evaluations of frame size
are used to indicate differences in potential
mature size, while muscle-thickness assessments
are used to reflect inherent variation in muscu-
larity. Results of the present study support the
USDA feeder-grading principles to the extent
that: (1) feeder cattle frame-size classification
was indicative of differences in absolute growth
rate and slaughter weight at a specified level of
130 TATUM ET AL.
fatness, and (2) m u s c l e - t h i c k n e s s classification
was associated w i t h d i f f e r e n c e s in carcass
m u s c l e - t o - b o n e ratio.
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