Published December 11, 2014
Review of Some Aspects of Growth and Development of Feedlot Cattle1
Fredric N. Owens*,2, Donald R. Gill*, David S. Secrist*, and S. W. Coleman†
*Oklahoma Agricultural Experiment Station, Animal Science Department, Stillwater 74078 and
†Grasslands Research Laboratory, ARS, El Reno, OK 73036
ABSTRACT: Growth in animals is defined as
accretion of protein, fat, and bone. Although growth
typically is measured as the change in live weight,
nutrient retention is estimated more precisely by
measuring empty body weight and composition,
whereas production economics are measured ideally
through carcass weights and quality. As a percentage
of live weight gain, carcass weight gain usually is a
much higher percentage during the feedlot phase than
during the growing phase of production because
dressing percentage (ratio of carcass:live weight)
increases with maturation and is greater with concen-
trate than with roughage diets. At a given fraction of
mature body size (maximum body protein mass),
body fat percentage seems to be a constant. Mature
size may be altered genetically and nutritionally.
Protein accretion declines to zero when cattle reach
their mature body size (approximately 36% fat in
empty body weight in modern cattle) even though
Key Words: Growth, Fat, Protein, Deposition, Feedlot Cattle
Introduction
Growth and development of ruminants have been
discussed and reviewed for many years. Discussions
and reviews are presented in both classic texts
(Brody, 1964) and more recent articles (Elsley, 1976;
Webster, 1977; Robelin and Geay, 1984; Rohr and
Daenicke, 1984; Oltjen and Garrett, 1988; Bergen and
Merkel, 1991; Lindsay et al., 1993). Various authors
have speculated or modeled that rate and composition
of tissue accretion may be controlled by chronological
age, physiological age (as related to sex, mature size,
maturity), energy intake, hormonal status, relative
1Presented at a symposium titled “Appraising Growth and
Development of Feedlot Cattle” at the ASAS 86th Annu. Mtg.,
Minneapolis, MN. Financial support was provided by Syntex Animal
Health. Approved for publication by the Director, Oklahoma Agric.
Exp. Sta. This research was supported under Project H-2123.
2To whom correspondence should be addressed.
Received November 28, 1994.
Accepted June 18, 1995.
3152
mature animals can continue to accrete fat. Although
fat accretion can be reduced by limiting the supply of
net energy, rate of fat accretion by finishing steers
given ad libitum access to high-concentrate diets
seems to reach a plateau at approximately 550 g daily.
Protein mass, in contrast, increases in proportion to
empty body weight. The protein:fat ratio of the carcass
can be increased through increasing mature size, by
administering hormones or hormonal modifiers, by
limiting energy intake during the growing period or
finishing period, or by slaughtering cattle at an earlier
stage of maturity. Energetically, efficiency of accretion
of fat is approximately 1.7 times that of protein. But
because more water is stored with deposited protein
than with deposited fat, lean tissue gain is four times
as efficient as accretion of fat tissue. Conversion of
protein to fat is very inefficient, suggesting that excess
protein is utilized inefficiently.
J. Anim. Sci. 1995. 73:3152–3172
turnover of tissues, and cell number (DNA) and
activity. This review will concentrate on methods of
performance measurement in feedlot cattle given ad
libitum access to concentrate diets (so that energy
accessibility should not limit growth rate), methods
by which composition of the body and products can be
altered, and theories about limits to growth rate.
Body Weight Measurements
Growth is routinely measured as the change in live
weight or mass. Weighing conditions vary. Full live
weight is measured without withholding feed or water;
thereby, it varies during a day due to patterns of feed
and water intake (Zinn, 1990). Precision of weight
measurement of individual animals is reduced by
animal movement in the scale and loss of digesta from
the tract. Weighing on consecutive days can reduce
daily variation slightly (Stock et al., 1983). Full live
weights are uncertain as an index of growth when feed
or water intakes and digestive tract fill are altered by
dietary roughage level, weather changes, or feeding
 GROWTH OF FEEDLOT CATTLE 3153

Figure 1. Relationship of empty body weight (EBW) to body weight, either shrunk (SBW) or unshrunk (BW), and
digestive tract fill (FILL). Equations for lines from top to bottom in the figure are from Jesse et al. (1976; EBW = .972
SBW − 13.6; so FILL = .028 SBW + 13.6); Williams et al. (1992; EBW = BW − FILL; where FILL = (.05354 + .329 NDF) ×
((BW/200)−.332) × (1.0 − .246 X − 1.481 X2 + 1.107 X−3) (1 + .35 X), where NDF is NDF content of the forage (decimal
fraction assumed to be .4), X is dietary concentrate level (decimal fraction assumed to be .9), and X is 0 for silage or
pasture and 1 for hay and assumed to be hay]; ARC (1980; EBW = BW / 1.09 − A, where A = 4 for concentrate diets,
14 for mixed diets, and 25 for long, dry roughage and was assumed to be 4); NRC (1984; EBW = .891 SBW; so FILL =
.109 SBW); Moulton et al. (1922; EBW = BW − FILL; where FILL = BW × .788 BW−.332); and this review where EBW =
.917 SBW − 11.39, so FILL = .083 SBW + 11.4.

patterns. In an attempt to avoid such problems,
weight can be measured “shrunk” (i.e., after a period
of time of feed and(or) water withdrawal). This
reduces the degree of variation in the amount of
digesta in the gastrointestinal tract. Primarily due to
loss from the digestive tract, shrinkage will vary with
time and diet. More digestive tract fill is lost when the
digesta are high in moisture content and when
animals are active. However, a period of shrink
disrupts feeding schedules and can complicate re-
alimentation even though ruminal outflow decreases
when feed is withheld.
A more precise index of energy and nutrient content
of the body is empty body weight ( EBW) . Here,
digesta are totally and laboriously removed from the
gastrointestinal tract after animals are slaughtered
and all remaining tissues are weighed. Various
regression equations have been developed to estimate
empty body weight from body weight (Figure 1).
Except for the NRC (1984) equation, all equations
predict that empty body weight increases as body
weight increases, being 85 to 95% of body weight. In
contrast to the other equations, that of Williams et al.
(1992) logically accounts for multiple factors by
including effects on fill by fiber content of the diet,
concentrate level, and animal maturation. Although
greater fill usually is detected with hay than with
other roughages, the suggestion that feeding hay
rather than silage or pasture will increase fill by 35%
is surprising and needs further study.
Empty body weight also can be estimated from
carcass weight (Figure 2). Here the relationship is
linear but with sizable differences in the intercept (30
to 61 kg) and the slope (1.36 to 1.45 times carcass
weight). Researchers often calculate shrunk live
weight from hot carcass weight and the mean dressing
percentage for a treatment or a slaughter group.
Dressing percentage for an individual animal is
defined as hot carcass weight divided by live weight.
Note that intercepts are involved with all equations
relating empty body weight to carcass weight (Figure
2 ) and that shrunk body weight is not directly
proportional to body weight (Figure 1). This suggests
that calculating either empty or shrunk body weight
simply by dividing carcass weight by dressing percent-
age for a group of animals may be biased if carcass
3154 OWENS ET AL.

Figure 2. Relationship of empty body weight (EBW) to carcass weight (CW). Equations for lines from top to bottom
in the figure are from Holzer and Levy (1969; EBW = 1.39 CW + 60.6); Holzer and Levy (1969; EBW = 1.44 CW +
41.2); Lofgreen et al. (1962; EBW = 1.45 CW + 31.7); Fox et al. (1976; EBW = 1.40 CW + 40.2); and Garrett and Hinman
(1969; EBW = 1.36 CW + 30.3).

weights differ greatly. Nevertheless, shrunk weight
calculated from carcass weight divided by mean
dressing percentage typically has approximately 20%
less variation than measured shrunk weights (Van
Koevering et al., 1995), indicating that precision is
improved when variation due to digestive tract fill is
removed. However, because dressing percentage and
carcass fat percentage typically but variably increase
with slaughter weight, empty body or carcass weight
alone is imprecise as an index of energy retention.
Classically, dressing percentage has been correlated
with fatness or degree of finish. In recent work by
Coleman et al. (1995), the correlation between
dressing percentage and weight of the digestive tract
contents was negative but slightly closer ( R 2 = .99)
than its correlation to fat percentage of the carcass
( R 2 = .93). Robelin and Geay (1984) observed that
digestive tract contents early in life represent under
5% of body weight but that this percentage increases
rapidly to reach approximately 20% of body weight in
cattle at 200 to 250 kg; thereafter, fill (the difference
between shrunk and empty body weight as shown in
Figure 1 ) declines to reach less than 12% when
shrunk body weight of growing animals reaches 350
kg. Additional factors altering fill were outlined by
Williams et al. (1992). These changes suggest that
the digestive tract reaches its mature size and weight
before other carcass components. This would cause fill
to be inversely correlated with fat content of the
carcass, another index of maturity. The fact that DMI
decreases as carcass fat increases has caused specula-
tion that accumulating fat reduces volume for the
digestive tract (Reid and Robb, 1971) and equations
predicting DMI often include carcass fat (Fox et al.,
1992). However, correlations alone do not prove cause
and effect. Furthermore, the effect of altered growth
rates on offal components is not fully clear. Murray et
al. (1977) indicated that weight of contents of the
reticulorumen was greater for steers with restricted
feed intake, and Carstens et al. (1991) has shown
that compensatory gain was attributable largely to
increased gut fill plus weight of the digestive tract and
other internal organs. Environmental factors such as
season (Pusillo et al., 1991) and day length (Tucker
et al., 1984) also may alter dressing percentage, but
whether these changes are due to altered body
composition or fill is not clear.
Because digestive tract fill changes with maturity
and diet, the proportionality among shrunk, empty
body, and carcass weight becomes suspect. Although
these three measurements usually parallel each other,
proportions can change markedly at different stages of
production. This means that the fraction of weight
gain that appears in the carcass can differ widely, as
 GROWTH OF FEEDLOT CATTLE
can be illustrated using data from Coleman et al.
(1995) in Figure 3. In this study, cattle were grown
for 145 d (fed either sorghum silage or a limited
amount of a high-concentrate diet). Subsequently,
these cattle were placed into the feedlot for 105 d.
Means for the growth phase (averaged across the two
systems) and the feedlot phase of production are
shown in Figure 3. As a proportion of live weight gain,
empty body weight gain averaged 77% during the
growth phase. In contrast, empty body weight gain
was 118% of live weight gain during the finishing
phase. Carcass gain, as a fraction of live weight gain,
was almost twice as great during the feedlot phase as
during the growth phase. Hence, the proportion of
shrunk or full weight gain that was carcass gain was
much greater during the feedlot than during the
growth phase. In some studies, with large, grazed
Holstein steers, carcass weight gain during the feedlot
phase has exceeded shrunk body weight gain! This can
happen only when dressing percentage increases
drastically. In the Coleman et al. (1995) study,
dressing percentage increased from approximately
57% to 64% during the 105-d finishing period. What
can explain this increase in dressing percentage?
From this same data set, proportional weights of
various body and digesta components are presented in
Figure 4. Weights of most organs, expressed as a
proportion of shrunk weight, remained quite constant;
Figure 3. Relative rates of live, empty body, carcass, and lean gain by cattle during growing (backgrounding on
either sorghum silage or limit-fed grain) and finishing (ad libitum access to grain) stages of production. Data from
Coleman et al. (1995).
3155
however, digestive tract fill decreased from 15.6 to
8.5% of shrunk weight as time on the feedlot diet
increased and calves were switched to ad libitum
intake of a concentrate diet. Both maturation and diet
may be involved in this change, because when higher
forage levels are fed throughout the finishing period,
percentage of weight as gastrointestinal fill may not
decrease (Buckley et al., 1990).
Body Composition Measurement
Ultimately, weights of closely trimmed retail prod-
uct are preferable to live or carcass weights as an
index of the commercial value of a carcass. Few
research trials have carried evaluation of growth to
this extreme, although genetic selection in poultry
already is evolving away from emphasis on decreasing
body fat content toward increasing breast muscle
weight. To date, most studies with beef cattle quan-
tifying tissue accretion have examined the quantities
of water, fat, protein and ash in either the empty body
or the carcass. Methods to assess body composition
were reviewed extensively by Blaxter (1989). Chemi-
cal composition of the ground empty body ideally
would be the standard. However, many studies rely on
other measurements such as chemical composition of
specified carcass parts or specific gravity of the total
3156 OWENS ET AL.
Figure 4. Components of empty body weight at various times in the feedlot. Data from Coleman et al. (1995).
carcass to predict empty body composition. For many
years, rib sections have been used to appraise total
body composition (Hopper, 1944; Hankins and Howe,
1946; Alhassan et al., 1975). Alhassan et al. (1975)
indicated that rib cuts improved accuracy of predicting
fat and energy content even though shrunk body
weight alone quite accurately predicted weights of
carcass protein and water.
Carcass specific gravity, based on the elegant and
extensive work by various workers (Lofgreen and
Garrett, 1968; Garrett and Hinman, 1969; Preston et
al., 1974), is used widely to estimate chemical
composition of the carcass and the empty body. The
regression equations were developed largely with
British cattle breeds weighing over 400 kg BW and
verification with other breeds and weights has not
been extensive. Some cattle breeds such as the
Holstein deposit a higher proportion of fat internally
in non-carcass components rather than in the carcass,
unlike the more traditional beef breeds (Dolezal et al.,
1993). Old and Garrett (1987) developed equations
for estimating body composition from carcass specific
gravity for Charolais cattle that differed from the
equations for British breeds. Alhassan et al. (1975)
suggested that specific gravity was less predictive of
body fat when body fat was under 20% of weight,
somewhat greater than the 12% minimum indicated
by Garrett and Hinman (1969). Nevertheless, most
estimates of body composition and tissue accretion are
based on this proven technique.
For estimating tissue accretion, repeated observa-
tions on a single animal would be preferable to the
posthumous measurements discussed above. Conse-
quently, extensive effort has been expended to quan-
tify body composition of live animals. Based on
Moulton’s (1923) concept of “chemical maturity,”
which states that composition of the fat-free body
reaches a constant when cattle reach about 3 mo of
age, regression equations have been developed relat-
ing body fat and body protein to body water (Kock and
Preston, 1979; Byers, 1980; Hammond et al., 1990).
Water, in turn, can be estimated by measuring
dilution of various pulse-dosed markers including
deuterium, tritium, and urea. Such techniques allow
body composition to be quantified at specific intervals
so that accretion during specific interim periods can be
calculated. Unfortunately, the accuracy of various
body composition prediction equations across various
weight classes from calves to finished steers is limited
(Rule et al., 1986), with slight changes in the ratio of
water:lean body mass (Blaxter, 1989) both across
breed types and frame sizes that differ in site of lipid
accretion (omental vs subcutaneous; Ferrell and
Jenkins, 1984b; Dolezal et al., 1993), and across
gender types (bulls and implanted steers have a
higher proportion of their weight in the forequarter).
Supporting Blaxter’s (1989) contention, regression of
protein against fat free empty body weight using the
data assembled for this review revealed that the
protein to fat-free empty body weight was directly
 GROWTH OF FEEDLOT CATTLE
related to EBW for cattle entering the feedlot phase
but inversely related to EBW for finished cattle.
Technologies to quantify muscle mass, fat thickness,
and marbling through magnetic resonance and ultra-
sound techniques have been developed and are finding
use at abattoirs (Gardner et al., 1995).
Impact of Weight on Body Composition
To examine effects of growth rate, body weight, and
energy retention specifically for cattle (steers and
bulls) given ad libitum access to high-concentrate
diets, we screened the published literature. In much of
the published work, equations rather than data are
provided and essential information relating to chemi-
cal composition, weight, cattle age, and feeding
duration often is lacking. The published information
that we located included literature cited below plus
papers by Preston et al. (1973), Eversole et al.
(1981), Schroeder et al. (1987), Anderson et al.
(1988a, b), Zinn (1989, 1992), Perry et al. (1991),
Zinn and DePeters (1991), Comerford et al. (1992),
Rumsey et al. (1992), and Ferrell and Jenkins
(1994). Values for rates of weight, fat, and protein
gained also were screened for apparent errors. When
the reported empty body weight gain fell more than
3157
10% below or 5% above that calculated from the
reported amounts of fat and protein retained (fat at
90% DM, fat-free lean tissue [protein] at 22% DM),
data for those cattle were excluded from the data set.
In each of these studies, steers or bulls had been given
ad libitum access to concentrate diets and tissue
accretion had been estimated by either carcass specific
gravity, rib section analysis, or complete empty body
grinding. This compilation included information from
16 different trials that had measured body composi-
tions for 1,174 cattle ( 4 to 72 animals per mean
value); initial weights ranged from 143 to 416 kg,
finished weights ranged from 356 to 571 kg, and daily
gains ranged from .67 to 1.87 kg.
The relationships between fat and protein mass to
empty body weight from this data set for feedlot cattle
are presented in Figure 5. The general shape of the
curve matches most of the relationships from the
literature indicating that mass of fat increased quad-
ratically with weight whereas protein mass increased
more linearly. This is the logical consequence of
having an accretion rate for fat that is greater than
that for protein. This change in body composition can
be ascribed to degree of maturity and should not be
extrapolated to suggest that rate of weight gain alters
composition of gain. When net energy intake is
restricted, rate of fat accretion often is reduced

Figure 5. Mass of fat (circles) and protein (squares) in empty body of feedlot cattle at various empty body weights
(EBW) in kilograms for cattle at the start of the feeding period (open symbols) and end of the feeding period (filled
symbols). Equations for the data set assembled were: Fat (kg) = −11.34 (± 15.2) + .0991 (± .095) EBW + .000494 (±
.00014) EBW2; R2 = .89; and Protein (kg) = −2.35 (± 3.64) + .2328 (± .0228) EBW − .000156 (± .000033) EBW2; R2 = .92.
3158 OWENS ET AL.

although protein growth may continue at nearly
normal rates if protein intake is adequate (Anderson
et al., 1988a).
Values in the equation did not match those
previously summarized by Simpfendorfer (1974) from
data sets involving 101 steers from several days to 57
mo of age from papers published before 1922. Those
equations based on historical data were: Fat ( k g ) =
−.61 + .037 EBW + .00054 EBW2 and Protein ( k g ) =
−2.418 + .235 EBW − .00013 EBW2. Similar equations
for this data set were: Fat ( k g ) = −11.34 + .0991 EBW
+ .000494 EBW2; R2 = .89; and Protein ( k g ) = −2.35 +
.2328 EBW − .000156 EBW2; R2 = .92.
Maturity and Composition
Mature body size or mature weight has received
several definitions. Several workers have employed
the definition of mature body weight proposed by
Taylor as discussed by Keele et al. (1992). By that
definition, mature weight is the point of body weight
equilibrium for cattle fed forage diets. At this point,
chemical fat content of the empty body is approxi-
mately 25%. In contrast, others including Brody
(1964) and Fox and Black (1984) have used the point
at which protein accretion ceases as an estimate of
mature body weight. We will use the latter definition.
The equations of Simpfendorfer (1974) and those we
devised above relating weight of protein to empty body
weight can be used to estimate the body weight at
which protein accretion reaches zero. Then, using the
equations that relate weight of fat to empty body
weight, one can calculate at what body fat percentage
protein accretion ceases. As presented in Figure 6,
Simpfendorfer’s equations predict that body protein
accretion ceases when empty body fat reaches 52%
(empty body weight of 904 kg). In contrast, equations
generated from our data set predict that protein
accretion ceases when empty body fat equals 36.2%
(at an empty body weight of 746 kg). This comparison
suggests that modern cattle have lower mature body
weights than those produced during the early part of
this century. For various classes of cattle, presumably
the proportional weight relationships developed by
Fox and Black (1984) would alter the weight but not
body composition at maturity. If one accepts that body
composition at a given fraction of mature weight is
fixed (and linearly proportional to maturity) and that
fat content of modern cattle at protein maturity is
about 36% ( a value of 34% was employed previously
in modeling lamb growth by Wang and Dickerson
[1991]), then one presumably can calculate mature
weight of an animal (maximum protein mass) from

Figure 6. Fat and protein content of weight gain calculated from equations of Simpfendorfer (1974) and literature
review of this paper. From the Simpfendorfer equations, protein content is maximum when empty body weight is
903.8 kg and fat content of empty body is 52.44%. Regression of values from this literature summary indicate that
protein content is maximum when empty body weight is 745.6 kg and empty body fat is 36.29%.
 GROWTH OF FEEDLOT CATTLE
fat content and empty body weight. For example,
expected protein mass at maturity would be double
the protein mass measured when empty body fat
percentage equals 18%. Compared with the difficulty
in determining when empty body protein mass reaches
a plateau, it is relatively simple to quantify fat content
that in turn can be used to predict degree of maturity.
Whether the variation among individual animals of a
given genotype in empty body fat content can be fully
attributed to different mature body weights or to
differences in timing of lean and fat deposition
remains unclear.
How can mature size be altered? Certainly mature
size can be altered by genetic or phenotypic selection,
as demonstrated by Jenkins et al. (1981) and Ferrell
and Jenkins (1984a). Furthermore, placing cattle on
ad libitum concentrate diets early in life seems to
reduce mature size (Coleman et al., 1993; Gill et al.,
1993b). For early-maturing cattle, moderate growth
rates are recommended through 75% of slaughter
weight (Coleman et al., 1993) to avoid a reduced
carcass weight as animals reach their mature size.
Nevertheless, as animals increase in maturity, the
percentage of fat in the empty body will increase
(Jesse et al., 1976; Byers 1980; Fox et al., 1992).
Ever since Brody (1964) developed his growth
curves for animals relating body weight to three
factors (i.e., post-conception age, mature weight, and
a deceleration factor) a debate has raged between two
schools of thought about chemical composition of the
body during growth. The concept supported by much of
the research data is that body composition at a given
empty body weight is fixed (Reid and Robb, 1971;
Jesse et al., 1976). Hence, altering the rate of growth
should have no impact on composition at a specified
body weight. In contrast, cattle producers for years
have fed “growing rations” to cattle of smaller frame
size to increase carcass weight at slaughter, the time
when animals had accumulated an adequate amount
of intramuscular fat. Note that this practice of
growing cattle would be ineffective if the former
concept were valid. Whenever two intelligent groups
disagree on a topic, both usually are correct, they
simply are not fully defining their position.
In elegant studies by Preston (1978), Angus and
Charolais steers were fed for 486 d; half were not
implanted but the other half were implanted repeat-
edly with estrogenic implants. Weight at 486 d, which
may be proportional to mature weight, was increased
by implants by 13 and 18% for the Angus and the
Charolais steers, respectively; this mature weight
difference is similar to that expected between bulls
and steers. Subsequently, body composition is shifted
by estrogenic implants in a fashion similar to that
expected from an increased mature size, as has been
reported previously by Byers (1980), Loy et al.
(1988), and Lemieux et al. (1990). These findings
suggest one obvious solution to the body composition
3159
dilemma discussed above: body composition when
expressed as a percentage of mature weight or size
may be fixed, but mature size, rather than being fixed,
may be elastic. Here, mature size is defined as the
point at which protein mass reaches a plateau. Beyond
this point, animals can deposit fat and gain weight,
but protein accretion presumably would be nil. The
point at which protein accretion reaches zero is not
fully clear because it is very difficult to determine the
point at which a plateau begins. Indeed, Wright and
Russell (1984) indicated that for every kilogram of
increase in empty body weight, 600-kg mature cows
deposited 808 g of fat and only 40 g of protein (4%).
Ferrell and Jenkins (1984b) fed cull cows three levels
of energy for 140 d. When ME intake was increased
from a low to a medium level, empty body weight
increased 81 kg and empty body fat and empty body
fat-free dry matter (lean mass that should be
proportional to protein) were both increased (37 and
8 kg, respectively). A further increase in ME intake of
these cull cows, from medium to high, also increased
empty body weight 36 kg, but this was primarily due
to increased empty body fat, not empty body lean mass
(34 and .4 kg, respectively). At the medium ME level,
cows had attained an empty body fat content of 32.3%.
This suggests that deposition of fat-free dry matter
ceases before deposition of empty body fat. Presence of
a specific empty body fat concentration above which
lean deposition ceases remains debatable, however,
because regressions of fat-free dry matter against
empty body weight show little curvature (Charles
Williams, personal communication). Certainly, previ-
ous body condition and hormonal status or implants
could alter propensity for protein accretion by mature
animals.
Impact of Nutritional History on
Protein and Fat Accretion
Effects of various production systems and back-
grounding conditions were examined in several
reports by Gill et al. (1993a, b). Angus-crossbred
steers of a narrow genetic base were placed in the
feedlot at five different ages. The first group was
placed in the feedlot after being weaned at an early
age (105 d). Other groups entered the feedlot after
weaning at the normal time (240 d), after such calves
had grazed wheat pasture for an additional 112 d, or
after having grazed native winter range followed by
an additional period of grazing summer grass for
either a short season (68 d ) or the full summer (122
d). Representative animals were slaughtered at each
of these points as well as following feeding to a
constant degree of finish (1.2 cm thickness of fat over
the longissimus muscle as measured ultrasonically).
In Figure 7 are shown the composition at the end of
these various background periods (the lower points on
3160 OWENS ET AL.
Figure 7. Fat and protein accretion vs empty body weight during the feedlot phase of cattle subjected to different
growing or backgrounding systems. Solid line is regression line from Figure 6. Data from Gill et al. (1993a,b).
each line) and at the end of the feedlot period (upper
points on each line) for the various groups as
compared to the overall regression lines for all cattle
from Figure 5. First, compare the body compositions of
the various groups at the start of the feedlot period
that corresponds to the end of the different back-
ground periods (lower points on each line). Protein
mass had increased with age, matching the expected
relationship from the overall regression line. Fat mass
had increased with age or weight, but fat mass
accumulated during the background period varied
with energy intake during the background period; it
was lower for calves that had grazed the full summer
season and had heavier initial weights than for those
that had grazed a shorter time. Despite wide differ-
ences in the relationship of fat to body weight, body
protein mass had closely matched body weight.
Another method of illustrating these same data is
to substitute a time scale for a weight scale. This helps
illustrate the effects of time or age on body composi-
tion. Lower and upper points on each line for these
five sets of cattle (Figure 8 ) represent the mass of fat
or of protein at the animal ages at the start and the
end of the feedlot phase. On the average, each
additional background day had increased empty body
protein by 60 g and had increased empty body fat by
240 g. Slopes of lines connecting the starting and
ending points (change in mass per unit of time) for
each group of cattle reflect rates of fat and protein
accretion. Surprisingly, rates of fat accretion were
virtually equal regardless of the starting point.
Unfortunately, intermediate points are not available
to detect the degree of curvature of these lines.
In contrast to fat, protein accretion rates at the
different ages differed with background. For the three
younger sets of cattle, the higher the initial protein
mass, the steeper the line or the more rapid the
accumulation of protein. With longer grazing, and
perhaps a protein shortage during backgrounding,
protein growth rate during finishing tended to be
slower despite a larger initial protein mass. Whether
this represents stunting or simply lack of appropriate
hormonal stimulation later in life is not clear. These
cattle all received estrogenic implants but not trenbo-
lone. From Figure 7, it seems that protein accretion
closely paralleled body weight but total accretion of fat
varied with initial body fat content. Indeed, Figure 8
suggests that rate of fat accretion during the finishing
period was more closely related to time rather than
body weight. To examine this effect of time further, we
examined the impact of performance on body composi-
tion.
Impact of Performance on Body Composition
The impact of rate of gain on body composition has
been examined by Byers (1980), Old and Garrett
 GROWTH OF FEEDLOT CATTLE
Figure 8. Fat and protein accretion vs animal age during the feedlot phase of cattle subjected to different growing
or backgrounding systems. Data from Gill et al. (1993a,b).
(1987), and Slabbert et al. (1992). These workers all
tested the impact of reducing rate of gain on rates of
accretion of fat and protein by restricting feed intake.
In most of these studies (Figure 9), reducing rate of
gain by restricting feed intake (moving to the left on
the graph) reduced fat content of gain when compared
with the overall regression line noted for cattle given
ad libitum access to feed to be discussed later.
Restricting feed intake reduced protein accretion in
several studies. In all but one study, fat accretion was
reduced. Often, however, final body composition was
not altered enough for differences to be significant. If
accretion of fat alone is reduced, restricting energy
intake can increase the protein:fat ratio of the empty
body. Based on the concept of constancy of body
composition, these data also could be interpreted to
suggest that restricting feed intake increases mature
body size. Nevertheless, results of limit-feeding
studies have been extrapolated directly to cattle given
ad libitum access to feed and, combined with the fact
that maturation increases fat content of the body,
have been interpreted to indicate that animals with
faster rates of gain will accrete more fat and less
protein. Such extrapolation is like concluding that
stride length can fully explain differences in perfor-
mance of race horses because hobbling a horse will
decrease its racing performance.
By stepwise regression, we evaluated the relation-
ship between empty body weight gain and rates of fat
3161
and protein accretion from the literature data set
discussed above (Figure 10). Although other factors
including maturity, gender, genotype, and age may
alter the relationship of rate of weight gain to
composition of gain, sufficient data are lacking to
subdivide this data set by most of these factors.
Consequently, relationships were developed across
these factors with cattle given ad libitum access to
feed. Differences observed may be attributable to
either genetic or environmental factors. Although
diversity of the literature decreases statistical preci-
sion and the relationships detected may be casual
rather than causal, including diverse data should
increase the range of extrapolation and applicability of
findings across these factors.
Regression lines indicate that as empty body gain
( EBADG) increased up to approximately 1.3 kg/d,
rate of fat accretion increased. But with gains higher
than 1.3 kg/d, similar to gains expected for feedlot
cattle, fat accretion reached a plateau ( P < .01)
whether data from feedlot cattle with all rates of gain
were included (Fat accretion [kg/d] = .591 EBADG −
.1474 EBADG2 − .0123; n = 100; R2 = .52) or cattle
with rates of gain below .8 kg/d were excluded (Fat
accretion [kg/d] = .766 EBADG − .2105 EBADG2 −
.123; n = 75; R2 = .28). The upslope at slower rates of
gain suggests that fat accretion was limited by energy
intake. Unfortunately, diet compositions and net
3162 OWENS ET AL.

Figure 9. Fat (heavy upper lines) and protein (heavy lower lines) as a percentage of empty body weight gain for
steers restricted in growth by limit feeding. Lines connect treatment means within individual trials. Regression lines
across all data points from ad libitum studies are shown in lighter lines. Data from Preston et al. (1973), Old and
Garrett (1987), and Delfino and Mathison (1991).

energy intake are not available from most of the
experiments in this data set. However, presence of a
significant quadratic factor and the plateau questions
the validity of the concept that rate of fat accretion is
more rapid for faster-gaining cattle given ad libitum
access to feed. Note that these values are all from
cattle given ad libitum access to feed, not from
individual animals with identical genetics. Conse-
quently, these data probably reflect differences in
genotype in addition to ADG. If one assumes that 1 )
protein accretion is weight-dependent, 2 ) body compo-
sition at any fraction of mature weight is constant,
and 3 ) fat accretion reaches a plateau, this means
simply that cattle that are capable of faster rates of
gain, like cattle with larger mature weight, deposit
more protein per unit of fat deposited.
Protein accretion did not reach a plateau at higher
rates of gain but continued to increase at higher rates
of empty body weight gain. The regression based on
data from feedlot cattle indicated that protein accre-
tion rate was proportional to daily weight gain
(Protein accretion [kg/d] = .1550 EBADG − .0228; n =
100; R2 = .82); when cattle gaining less than .8 kg
empty body weight daily were excluded, protein
accretion rate increased in proportion to daily weight
gain squared (Protein accretion [kg/d] = .05736
EBADG2 − .07505; n = 75; R2 = .73). These regression
values indicate that rate of empty body weight gain
was related more closely to rate of protein accretion
than to rate of fat accretion, a logical conclusion if fat
accretion reaches a plateau. In view of the scatter of
points, it is not surprising that many workers in the
past have failed to detect significant differences in
body composition with only slight differences in rates
of gain and, based on lack of a significant effect,
concluded that rate of gain did not alter body
composition.
These plots could be interpreted to suggest that fat
accretion rate may reach a ceiling whereas rate of
protein accretion does not, but instead remains
dependent on animal maturity. The maximum fat
accretion rate of approximately 550 g/d for steers and
bulls given ad libitum access to feed with daily empty
body weight gains higher than 3 kg conflicts directly
with the classic concept that fat accretion at a given
physiological age for a specified genotype is greater
with more rapid rates of gain. Nevertheless, this
regression is across rather than within genotypes. In
most of the studies that support the classic concept
that faster gains increase fat deposition (Preston et
al., 1973; Byers, 1980; Old and Garrett, 1987; Delfino
and Mathison, 1991), the supply of feed available to
one genetic group of cattle was intentionally limited.
As discussed previously, fat accretion rate is decreased
 GROWTH OF FEEDLOT CATTLE 3163

Figure 10. Rates of fat and protein accretion vs empty body weight gain (kilograms/day) of finishing steers and
bulls given ad libitum access to concentrate diets. Fat accretion (kg/d) = .543 EBADG − .1426 EBADG2 + .0492; R2 =
.28 and protein accretion (kg/d) = .1718 EBADG − .0482; R2 = .89.

by restricting energy intake as shown in Figure 9.
Restricting feed intake also could affect body composi-
tion either by altering the hormonal status or cascade
of growth hormone to IGF or by changing the quantity
of specific nutrients available for metabolism. The fact
that restricting feed intake enhances energetic effi-
ciency more than expected from net energy relation-
ships (Old and Garrett, 1987; Hicks et al., 1990a)
supports the concept that limiting energy intake alters
composition and thereby may increase mature size. If
one accepts the concept of constancy of composition at
a given fraction of mature size, then animals with
higher mature size automatically would deposit more
protein at a given rate of fat accretion.
The relationships of rates of fat and protein
accretion to empty body energy gain (based on NRC
[1984] calculated assuming that depot fat contains 8
kcal/g and fat-free tissue, being 73% water, 22%
protein and 5% minerals, has 1.2 kcal/g) are
presented in Figure 11. As indicated by Lapierre et al.
(1992) and as could be extrapolated from these lines,
at very low rates of energy retention, protein accretion
can exceed fat accretion, but fat accretion increased
when energy retention increased. Each slope was
curved slightly. These relationships are based on
regression of trials with steers and bulls fed high-
concentrate diets. Effects of gender and breed type,
estrogenic and androgenic implants, somatotropin and
beta-agonists on protein and fat accretion rates have
been discussed by Bergen and Merkel (1991) and
Lindsay et al. (1993). For heifers, fat accretion rates
more than 600 g/d have been reported and, surpris-
ingly, fat as well as protein accretion rates were
increased by 200-mg implants of trenbolone acetate
(Bergen and Merkel, 1991). In contrast, somatotropin
enhanced protein accretion while retarding fat accre-
tion (Dalke et al., 1992; Lapierre et al., 1992). These
findings indicate that ceilings to growth might be
modified by gender and hormonal status. Such is the
case with swine in which gender and somatotropin
administration both alter fat and protein accretion
rate (Campbell et al., 1991).
Are Faster-Gaining Cattle Fatter?
Curves developed by Byers (1980) indicate that as
rate of gain increases, both protein and fat accretion
increase, with fat accretion increasing at an increasing
rate and protein accretion possibly decelerating. In
contrast, no deceleration in protein accretion is
evident in values cited by Fox and Black (1984),
Lemieux et al. (1990), or Slabbert et al. (1992).
Unfortunately, correlations between rate of weight
gain and accretion of protein and fat are confounded
by such factors as maturity, genetics, age, and weight.
On a theoretical basis, the question above can have
two different, but reasonable, answers, First, faster-
3164 OWENS ET AL.
Figure 11. Relationship of fat and protein accretion to energy accretion (ER) in megcalories/day for steers and
bulls given ad libitum access to concentrate diets. Fat accretion (kg/d) = .15 ER − .0057 ER2 − .162; R2 = .94; Protein
accretion (kg/d) = .267 − .072 ER + .0094 ER2; R2 = .34.
gaining animals will be fatter if they have capacity to
dispose of more calories by fat accretion. This, in turn,
should reduce mature size. Second, if animals that
gain less rapidly have lower energy intakes, they may
be physiologically younger and thereby leaner than
rapidly gaining cohorts. However, if faster-gaining
animals have a larger mature size, a higher initial
protein mass, or altered hormonal status that en-
hances protein accretion, then faster-gaining animals
should be no fatter than their contemporaries. Indeed,
in results from Gill et al. (1993a, b), it was the calves
that were early-weaned that had the slowest feedlot
gains but more empty body fat at a lighter slaughter
weight, not cattle that had grazed for several months
before being placed in the feedlot. They also had the
lowest feed intakes. This suggests that degree of
maturity and body composition may be predicted from
voluntary feed intake.
Composition of gain (fat and protein accretion as a
percentage of empty body weight gain) for steers and
bulls given ad libitum access to high-concentrate diets
and gaining weight at different rates from the
compiled literature data set discussed above is
presented in Figure 12. Percentage of fat in empty
body weight gain declined whereas percentage of
protein increased as rate of gain increased (Fat, % of
EBADG = 61.5 − 16.7 EBADG; R2 = .38; Protein, % of
EBADG = 11.7 + .86 EBADG; R2 = .19). The directions
of change in protein and fat with rate of gain in these
equations conflicts with the classical observation that
severely restricting feed intake reduces fat accretion
(Figure 9). Perhaps the plateau of the regression
simply reflects the fact that cattle capable of more
rapid gains may be leaner (larger mature size).
Alternatively, the relationship of fat accretion to
energy intake may be curvilinear rather than linear.
Perhaps the fat accretion ceiling is only reached with
ad libitum energy intake.
The fact that fat accretion rate for cattle gaining at
rapid rates tended to reach a plateau for feedlot cattle
gaining at moderately rapid rates suggests that fat
accretion rate may be a ceiling that in turn could limit
energy and feed intake. If one combines the concepts
of maximum fat accretion with the constancy of body
composition at a specific proportion of mature size, one
might estimate daily accretion rates for fat and
protein at specific proportions of mature size. This in
turn would allow one to predict rate of gain and NEg
intake. Rate of protein accretion declines markedly as
animals approach mature size, becoming nil at mature
size. However, most feedlot cattle are marketed before
they reach mature weight.
Limits to Growth
Many models of growth have been proposed. Unfor-
tunately, none of them accurately predicts fat and
 GROWTH OF FEEDLOT CATTLE
Figure 12. Fat and protein in accreted tissue as a percentage of empty body weight gain (EBADG) at various rates
of gain. Fat, % of EBADG = 61.57 − 16.7 EBADG; R2 = .38; protein, % of EBADG = 11.7 +.86 EBADG; R2 = .19.
protein accretion (Arnold and Bennett, 1989). These
workers proposed that some new concepts are needed
to explain the variation in rates of fat and protein
accretion. In contrast, when feed intake is known and
is used with the net energy equations, weight gain for
groups of animals are well predicted. For groups of
animals of similar mature weight, this should make
fat and protein accretion predictable. This suggests
that one major limitation in modeling relates to
inaccuracy in predicting intake of energy. The specific
factors limiting intake of feedlot diets or energy-rich
diets have been proposed variously to relate to
ruminal conditions (pressures, acid concentrations,
osmolality, absorption), nutrient absorption (specific
acids), and nutrient utilization (heat increment).
Feed intake has been reviewed in several symposia
(Owens, 1986, 1995).
Despite wide differences within and among trials in
rates of water and protein (lean) accretion, rate of fat
accretion by rapidly growing steers and bulls given ad
libitum access to high-concentrate diets may reach a
plateau at approximately 550 g/d, as discussed previ-
ously. Certainly fat accumulation can be reduced by
restricting intake of NEg, but with cattle given ad
libitum access to concentrate diets, few accumulated
more than approximately 630 g/d, although in one
study yearling steers (groups with both a moderate
and a high genetic potential for growth) deposited
approximately 850 g of empty body fat daily (Short et
3165
al., 1994). Might 550 g/d reflect a general ceiling?
Certainly lactating animals can secrete 1.5 kg of fat
each day, but over half of this is produced in the
mammary gland and secreted directly. Do lactating
animals consume more feed because the mammary
gland serves as a sink for energy disposal? Flatt
(1978) proposed that the rate of fat synthesis is
limited when the ATP:ADP ratio becomes excessive.
Lipogenesis requires reducing equivalents (NADH).
Metabolic cycles producing NADH generate ATP
during the process. More ATP is generated during
formation of NADH by some pathways than from
others. For example, ATP generation is less in the
malate/citrate cleavage cycle than from the pentose
phosphate cycle, but ruminant liver and adipose
tissues do not employ citrate cleavage unless glucose
supply is enhanced. If an increased supply of glucose
from intestinal digestion of starch would permit
lipogenesis to occur using the citrate cleavage path-
way, lipogenesis in the total body could be enhanced.
Site of fat synthesis is altered by nutritional status in
rats (Kim and Freake, 1993), and synthesis is greater
for intestinal tissue than for liver in the glucose-
deprived rat, a condition that may be similar to that in
the gluconeogenic ruminant. Intestinal fat synthesis
could explain why an increased supply of intestinal
starch often enhances efficiency by finishing cattle
(Owens et al., 1986) and lactating cows (Nocek,
1990) even though additional glucose is not recovered
3166 OWENS ET AL.
in the portal blood stream (Huntington and Reynolds,
1986).
If accumulation of excess ATP limits production of
reducing equivalents for fat synthesis, then altering
lipid turnover, modifying the VFA ratio (glucose being
synthesized from propionate vs only being spared by
acetate), and increasing the number of sites of fat
synthesis also might alter rate of fat accretion by
altering the ATP surplus. Furthermore, direct deposi-
tion of absorbed fatty acids, circumventing the need
for fatty acid synthesis, should increase the fat
accretion ceiling. The concept of a ceiling on fat
synthesis and the rate of fat accretion, as modified by
insulin/glucagon ratio, raises some fascinating ques-
tions for biochemists and physiologists.
As with fat, protein accretion represents the bal-
ance between synthesis and degradation. Protein
synthesis rate for growing ruminants is at least five
times greater than accretion rate (Bergen and Merkel,
1991). Accretion is dependent on hormonal status and
is enhanced by larger mature size. Certain hormonal
and partitioning agents can enhance accretion either
by enhancing synthesis or retarding turnover. In
addition, as mass of lipid accumulates in the empty
body with maturation, protein accretion is decreased.
As an overview, one might model fat accretion as a
constant of 550 g/d for feedlot cattle gaining more than
1.3 kg/d unless previous nutritional history permits
this value to be exceeded. Indeed, this could serve as
the maximum amount of excess energy that can be
accreted, which in turn would place a ceiling on
energy intake. Rate of protein accretion, in contrast,
depends on the initial protein mass, with greater
accretion with a larger initial protein mass in the
empty body. Working with this premise, one could
examine various methods of changing the protein:fat
ratio in the empty body and, presumably, in the
carcass.
First, growing cattle for a period of time before
placing them in a feedlot should increase mature size,
which in turn implies an increase in the initial protein
mass, the protein:fat ratio at a specific slaughter
weight, and the weight at a specific empty body fat
percentage. Increasing mature body size either geneti-
cally or by hormonal implants should act similarly,
although certain hormones also may decrease rate of
fat synthesis (Mersmann, 1991). Second, retarding
fat accumulation by limiting intake of NEg during
growth or finishing also will increase the protein:fat
ratio providing that protein accretion is not retarded.
Animals that have lost body fat and exhibit compensa-
tory gain also should have a higher protein to fat ratio.
And finally, slaughter at a lighter weight also will
increase the ratio.
Some of the differences between the classic and this
novel theory for the control of growth and body
composition deserve discussion. Classically, energy
intake is presumed to control both the rate and the
composition of accreted tissue. With faster gains, the
NRC (1984) net energy equations indicate that
accretion of both fat and protein increases (Figure
13). Similarly, as animals reach a higher percentage
of their mature weight, fat accretion rate is projected
to increase. The net energy equations predict that rate
of fat accretion can reach 1,150 g/d. That value is
much higher than indicated from regression analysis
of data discussed above from cattle given ad libitum
access to high-concentrate diets. Those data indicated
that although fat accretion rate varies with proportion
of mature weight (mean during the feeding period), it
did not increase as rate of empty body gain increased
(Figure 13). This would suggest that the height of the
plateau in rate of fat accretion varied only with degree
of maturity during the feeding period. Note that the
rate of fat gain and thereby of energy retention from
this data set was markedly lower than that projected
from NRC (1984) equations. With both the equations,
rate of protein gain increased with rate of empty body
weight gain but was lower for animals that were at a
higher percentage of mature weight during the feeding
period. Protein deposition values also tended to be
greater from the literature summary than NRC
(1984) would predict.
Derivations of these two different relationships may
be partly responsible for their differences. The NRC
(1984) values are largely extrapolations across groups
of cattle that had been fed different amounts of
energy, with energy intake limited for half of the
cattle. In contrast, only data from cattle given ad
libitum access to concentrate diets were used in our
literature data summary. Such cattle may differ in
their genetic potential for growth and(or) mature
size. Further, it may be erroneous to extrapolate these
literature data to conditions in which growth rate is
limited by programmed feeding. Comparison of the
two relationships suggests that increasing mature
weight would have much greater impact on body
composition following the literature data summary
equations than predicted by the traditional equations.
Total energy retention is the sum of accretion of
energy in the form of fat and protein. If rates of fat
and protein accretion are set by genetics and maturity,
then the rate of energy accretion, when combined with
energy expended for maintenance, could limit and
thereby control energy intake. By the classical ap-
proach, growth rate and composition are driven by
energy intake. In contrast, the plateau fat deposition
theory would suggest that energy intake is a response
to growth and body composition instead of the driving
force.
If fat accretion rate has a ceiling, then feed
(energy) intake by feedlot cattle would depend on
energy retention in the form of fat, which would be
fixed, plus protein, which in turn would depend on sex,
hormonal status, and mature size, and decrease with
maturity plus maintenance requirements. Whether
 GROWTH OF FEEDLOT CATTLE 3167

Figure 13. Fat and protein accretion as altered by maturity and rate of gain as projected by the traditional (NRC,
1984) and literature survey results from steers and bulls given ad libitum access to concentrate diets. Traditional
values were calculated from NRC (1984) equations for energy retention for large-frame steers assuming that fat tissue
contained 8 kcal/g, lean contained 1.2 kcal/g, and mature weight was 625 kg. Non-fat tissue empty body gain was
assumed to be 22% protein. Stepwise regression equations for the literature survey data set, ignoring factors not
significant at P < .15, were: Empty body fat gain (g/d) = 227.8 + 527.8 Maturity; R2 = .327; Empty body protein gain
(g/d) = 87.7 + 72.5 EBADG2 − 92 Maturity2; R2 = .913, where EBADG is kilograms/day and Maturity is empty body
fat percentage divided by 36.

maintenance requirements should be calculated from
metabolic size or protein mass, as suggested by results
from Ferrell and Jenkins (1984a) and Baker et al.
(1991), is not clear. In either case, voluntary energy
intake would be expected to decrease as cattle reach a
higher proportion of their mature weight due to
decreased energy retention as protein. This decrease
would be juxtaposed chronologically with an increased
energy need for maintenance. Such curves would
match the field observations (Hicks et al., 1990b) that
feed intake by feedlot cattle reaches a plateau, in
contrast to the NRC (1984) projection that feed
intake remains directly proportional to metabolic size
as animals mature.
Efficiencies of Fat and Protein Accretion
Despite extensive study, the relative efficiencies of
protein and fat synthesis and accretion remain ob-
scure, partly because efficiency can be expressed in
several ways (wet tissue vs calories; synthesis vs
accretion), partly because protein and fat accretion
rates are strongly correlated and have proven difficult
to quantify separately, and partly because main-
tenance requirements cannot be readily separated
from accretion requirements. Many researchers dis-
cuss caloric efficiency; in contrast, most livestock
producers measure efficiency in terms of live weight,
not calories. Most of the research with metabolic
efficiency comes from studies with laboratory animals
or nonruminants (Table 1). Efficiencies of fat accre-
tion as measured in animal trials are reasonably close
to the theoretical efficiency of fat synthesis. In
contrast, efficiency of protein accretion is about half
the theoretical efficiency of protein synthesis. In some
of these studies, efficiencies of synthesis were meas-
ured; in others, efficiency values are for accretion. As
nicely described by Bergen and Merkel (1991),
continual and rapid turnover of protein consumes
energy and decreases efficiency. Based on averages
from Table 1, the amount of energy required to accrete
1 g (9.2 kcal) of dry fat, at 12.1 kcal, is surprisingly
close to the amount needed to accrete 1 g (5.7 kcal) of
dry protein. On a caloric basis, fat accretion had an
average efficiency of 76% (heat loss of 24%), whereas
efficiency of protein accretion was only 47% (more
than 53% heat loss), as illustrated in Figure 14.
3168 OWENS ET AL.

Table 1. Energetic efficiency of fat and protein synthesis and accretion

Fat Fat Protein Protein

Species synthesis accretion synthesis accretion Source
%
Theoretical 70 — 94 — Blaxter, 1962
Theoretical — — 90 — Schiemann, 1963
Theoretical, ruminants 72 — 82 — Baldwin, 1968
Theoretical — — 86 38 to 55 Bergen and Merkel, 1991
Several — — — 24 Miller and Payne, 1963
Chickens — 78 — 51 Petersen, 1970
Rats — 70 — 46 Schiemann et al., 1969
Rats — 65 — 43 Pullar and Webster, 1974
Mice — — — 58 to 67 Bernier et al., 1987
Pigs — — — 45 Kotarbinska and Kielanowski, 1963
Pigs — 63 — 77 Kielanowski, 1965
Pigs — 77 — 43 Thorbek, 1970
Pigs — 70 — 54 Close et al., 1974
Pigs — 81 — 84 Close et al., 1979
Lambs — 81 — 82 Kielanowski, 1965
Lambs — 82 — 33 Orskov and McDonald, 1970
Lambs — 79 to 92 — 10 to 20 Rattray and Joyce, 1976
Lambs — — — 13 Matras and Preston, 1989
Calves — 85 — 45 Kirchgessner et al., 1976
Cattle — 69 — 34 Reid et al., 1980
Cattle — — — 46 to 55 Hayden, 1988
Mean ± SE 71 ± 1 76 ± 8 88 ± 5 47 ± 20

Figure 14. Energetics of accretion of fat and protein from various sources. Values are means across species from
Table 1.
 GROWTH OF FEEDLOT CATTLE
Thereby, on a caloric basis, fat accretion is approxi-
mately 1.6 times as efficient as protein accretion.
Accretion of energy is less efficient for protein than for
fat because of faster and less efficient turnover of
depot protein than of depot fat (Bergen and Merkel,
1991). Protein synthesis rate typically ranges from 5
to 10 times that of accretion in young growing
ruminants; accretion comprises a larger proportion of
total synthesis with faster accretion rates. Presuma-
bly, the large variation in efficiency of protein
accretion is due to variations in turnover rate. With
ratios of synthesis to accretion of 1, 5 (their estimate
for swine), and 10 (their estimate for cattle),
efficiency of protein accretion theoretically drops from
80% to 55 and 38% (Bergen and Merkel, 1991). Less
than maximum growth rates, by reducing protein
synthesis rate more than protein degradation rate,
will reduce efficiency of protein accretion. Most factors
that increase the synthesis rate for protein also
increase protein degradation rate (Bergen, 1974).
Faster protein turnover at greater accretion rates will
increase heat production and decrease gross energetic
efficiency. The energy expenditure associated with
protein turnover may explain why level of feeding
alters “maintenance” cost. Although the energetic
inefficiency of protein turnover is truly a “main-
tenance” cost, it generally is included as part of the
cost of protein accretion. Chemical compounds that
may decrease turnover without decreasing synthesis,
such as clenbuterol, cimaterol, or trenbolone acetate,
will enhance efficiency of protein accretion (Morgan et
al., 1989; Baldwin et al., 1991; Bergen and Merkel,
1991; Lindsay et al., 1993).
Growing animals do not store fat and protein
without retaining tissue fluids. Depot fat contains
approximately 10% water, whereas fat-free lean tissue
(protein) has approximately 78% water. So, when
calculated on a wet-tissue basis, kilocalories required
per unit of protein is less than one-fourth that of fat
(1.2 vs 8.3 kcal/g) independent of animal or diet
composition, as shown in Figure 14. Consequently,
when discussing efficiency of animal production, one
must distinguish between efficiency of conversion of
weight (ratio of weight gained to weight of feed
consumed) or of energy (ratio of retained to consumed
calories). In most cases, livestock are fed diets rich in
carbohydrate. And source of calories also may alter
efficiency of synthesis. Millward and Garlick (1976)
projected that synthesis of fat from carbohydrate
results in loss of approximately 15% of the energy as
heat. Whether losses are similar from acetate or
glucose are not certain. Protein synthesis from amino
acids seemed reasonably efficient at 85%. Body
synthesis of fat from absorbed fatty acids, at 98%,
resulted in less heat loss than synthesis of fat from
carbohydrate (Figure 14), making fat supplementa-
tion energetically advantageous. If heat dispersal
capacity is limited, this difference becomes relevant
3169
for livestock producers. Of additional concern is the
poor efficiency (35%) with which any excess protein is
converted to fat. If protein is inefficiently converted to
fat, an excess of either escape or microbial protein may
have less value than either carbohydrate or protein
that is degraded within the rumen. This low efficiency
might explain partly the adverse effects of excess
dietary protein on energetics reported by Tyrrell et al.
(1970).
Implications
Change in live weight is imprecise as an indicator
of growth (accretion of protein and fat tissue). When
cattle reach their mature body size (maximum body
protein mass), body fat content is approximately 36%
of empty body weight regardless of sex and back-
ground, even though fat content of mature animals
can exceed 36%. Mature body size depends on several
factors, including genetics and nutritional and hor-
monal status. For finishing steers and bulls gaining
more than 1.3 kg daily, rate of fat accretion seems to
reach a plateau at approximately 550 g daily. In
contrast, protein accretion rate depends on age and
mature body size and increases as rate of empty body
weight gain increases. The protein:fat ratio of the
carcass can be increased by increasing mature size;
this in turn can be achieved through administering
hormonal implants, limiting energy intake during the
growing or finishing period, or slaughtering finished
cattle at an earlier stage of maturity.
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