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A. L. Van Eenennaam, K. L. Weber and D. J. Drake
J ANIM SCI 2014, 92:2693-2701.
doi: 10.2527/jas.2013-7217 originally published online April 21, 2014
The online version of this article, along with updated information and services, is located on
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http://www.journalofanimalscience.org/content/92/6/2693
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per bull per calf crop. The peak Ncalf occurred at about 5
yr of age for bulls ranging from 2 to 11 yr of age. Weekly
conception rates as assessed by date of calving varied significantly and peaked at wk 3 of the calving season. The
distribution of calves born early in the calving season was
disproportionately skewed toward the highly prolific bulls.
The DNA paternity testing of the subset of those calves
born in wk 3 of the calving season was highly predictive
of overall bull prolificacy and may offer a reduced-cost
DNA-based option for assessing prolificacy. Prolificacy of
young bulls in their first breeding season was positively
linearly related (P < 0.05) to subsequent breeding seasons, explaining about 20% of the subsequent variation.
Prolificacy was also positively linearly related (P < 0.05)
to scrotal circumference (SC) EPD for Angus bulls that
had SC EPD Beef Improvement Federation accuracies
greater than 0.05. Varying prolificacy of herd bulls has
implications for the genetic composition of replacement
heifers, with the genetics of those bulls siring an increased
number of calves being disproportionately represented in
the early-born replacement heifer pool.
that regard, a herd bull has 2 qualities of value to commercial producers. One is his ability to impregnate as
many cows as possible, and the other is the ability to
pass genes for superior performance on to his offspring.
In the absence of the former, the latter is moot. Natural
service breeding is the predominant practice for beef
cattle operations in the United States, but few studies
have examined the variation in the number of calves
sired in multiple-sire breeding pastures and the consistency of an individual bulls performance over time.
Few genetic tools exist for selecting bulls with
superior breeding performance. Holroyd et al. (2002)
found that there were breed differences in a variety of
traits related to calf output (e.g., scrotal circumference,
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Table 1. Average bull age at the beginning of the breeding season and number of calves produced per natural service
bull in multisire breeding pastures on 3 commercial ranches (A, B, C) in northern California in 20092011
Ranch
A
A
A
A
A
A
B
B
B
B
B
B
C
C
C
A
B
C
A, B, C
A, B, C
A, B, C
A, B, C
A, B, C
A, B, C
1Where
Year
2009
2010
2011
2009
2010
2011
2009
2010
2011
20092011
20092011
20092011
2009
2010
2011
Spring
Fall
20092011
Calf crop
Spring
Fall
Spring
Fall
Spring
Fall
Spring
Fall
Spring
Fall
Spring
Fall
Fall
Fall
Fall
No. of
sires
18
19
22
19
17
19
8
10
8
12
4
12
30
27
38
114
54
95
85
88
90
77
186
263
Total
no. calves
353
346
435
328
402
286
141
214
142
247
110
266
642
567
573
2,150
1,120
1,782
1,696
1,719
1,637
1,583
3,469
5,052
Per bull
Minimum no. Maximum no. Mean no. calves
calves1
calves
SEM
3
47
19.9 3.8
1
47
19.6 18.2
3
45
19.8 3.8
1
48
18.4 22.1
4
53
24.2 4.7
1
33
16.8 14.5
1
45
16.7 10.0
10
50
21.8 9.3
3
30
16.5 7.4
4
44
20.2 12.9
18
42
26.5 14.4
3
51
22.8 6.2
2
54
20.3 3.0
1
52
19.9 3.8
1
64
14.4 5.7
1
53
18.6 6.0
1
51
19.9 3.8
1
64
21.1 13.2
1
54
19.9 3.8
1
52
20.1 1.9
1
64
19.7 7.7
1
53
20.5 12.2
1
64
19.2 5.0
1
64
18.9 13.1
bulls produced at least 1 calf. In 4.6% of the breeding seasons (12 out of 275) bulls produced no progeny.
Prolificacy
The overall mean Ncalf was 18.9 13.1 progeny,
with little variation among ranches, 18.6 6.0, 19.9
3.8, and 21.1 13.2 (P = 0.90) for Ranches A, B, and
C, respectively. Similarly, differences between years
(19.9 3.8, 20.1 1.9, and 19.7 7.7; P = 0.96) and seasons (spring, 20.5 12.2; fall, 19.2 5.0; P = 0.94) for
Ncalf were small. Additionally, Ncalf across the 15 calf
crops showed little variation (P = 0.51), ranging from
14.4 5.7 to 26.5 14.4 (Table 1). However, the mean
Ncalf per bull varied widely (P < 0.01), ranging from a
mean of 3.3 6.3 to 39.1 10.9 (Fig. 1). Repeatability
of Ncalf for bulls 3 or more years of age was 0.37, and
it was 0.33 when all bulls were included in the analysis.
I205. Individual calf 205-d weight did not vary significantly between ranches (232 10.1, 235 6.4, and 279
18.0 kg; P = 0.29) but showed significant year (225 2.4,
233 2.3, and 227 2.4 kg; P < 0.01) and season (spring,
237 2.5 kg; fall, 220 2.0 kg; P < 0.01) differences.
Again, bulls varied widely in mean progeny I205 (P <
0.01), ranging from means of 196 to 262 kg (Fig. 1).
T205. Total contribution of calf weight (sum of individual sex-adjusted 205-d wt) per bull showed a pattern
similar to that of Ncalf alone (Fig. 1). Total adjusted 205-
Figure 1. Mean number of calves (Ncalf; left axis), calf 205-d sexadjusted weight (I205), and total 205-d sex-adjusted weight (T205)/20 (right
axis) per natural service bull present in a multisire breeding pasture. Only
bulls that were present for the entire length of the breeding season and that
were in use for more than a single breeding season are included.
tion. The correlation between first breeding season prolificacy and mean subsequent prolificacy was 0.45 with
a rank order correlation of 0.47. Young bulls categorized
into 3 equal groups, HP, MP, and LP, based on their firstyear prolificacy were not related to subsequent categorization by chi-squared analysis (P = 0.20) or likelihood ratio chi-squared analysis (P = 0.20), although the sample
size of this young sire group was relatively small (n = 24).
Young bulls categorized by their first breeding season as
HP tended to remain HP (3/8 or 4/8), and only 12.5% (1/8)
fell to LP. Similarly, young bulls initially categorized as
LP mostly remained as LP (62.5%, 5/8), with 25% (2/8)
changing to MP but only 12.5% (1/8) improving to HP.
Prolificacy and EPD. Scrotal circumference (SC)
EPD was not significantly related (P = 0.16) to prolificacy all bulls were included in the data set. However, a
single outlier bull (from Ranch C) had a large Cooks distance (Di) and studentized residual. When that bull was
removed from the analysis, SC was related (P = 0.04) to
prolificacy (Fig. 3), where
Ncalf = 15.2 + 8.27 (3.0) SC (R2 = 0.13, SE = 8.7).
The equation for Ranches A and B combined (without
Ranch C) was within the confidence intervals for the
combined equation of Ranches A, B, and C (without
the single outlier bull).
Carcass weight (CW) was negatively related (P =
0.03) to prolificacy, where
Ncalf = 27.7 0.354 (0.158) CW (R2 = 0.09,
SE = 10.1).
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Figure 2. Calves per bull (Ncalf) per calf crop vs. age of the bull for
natural service bulls present in multisire breeding pastures on 3 northern
California commercial beef ranches (A, B, and C).
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Figure 4. Adjusted calves born per week of the calving season across
all 15 calf crops show that peak calving and implied peak conception occur
during wk 3 of the calving (and breeding) season. Means that differ (P < 0.05)
are noted with different letters. Error bars represent SEM.
Figure 3. Mean calves per bull (Ncalf) vs. scrotal circumference (SC)
EPD of Angus natural service bulls present in multisire breeding pastures on
3 northern California commercial beef ranches (A, B, and C). Bull 648 was
classified as an outlier on the basis of a high Cooks distance (Di) and studentized residual (P < 0.05).
Calving Distribution
Calving distribution showed the preponderance of
calves being born early in the calving season (Fig. 4).
The largest number of calves born in a single week occurred in wk 3 in 12 of the 15 (80%) calving seasons
evaluated. Pooled across ranches and adjusted for ranch,
year, and season, peak calving occurred in wk 3. In the 3
seasons where peak calving was not during wk 3, it occurred in wk 2 twice and wk 1 once.
The HP bulls sired more calf births per week during
the early part of the calving season than the MP or LP
bulls (P < 0.01; Fig. 5). Bulls siring more progeny (HP)
had a disproportionately higher percentage of calves
born early in the calving season. Low prolificacy bulls
tended to have a consistently low number of calves born
throughout the calving season. These data suggest that
high prolificacy is associated with the breeding of a
greater than expected number of cows early in the breeding season, ultimately leading to a larger total number of
progeny for the calf crop.
Alternative Prolificacy Assessment Methods
As an alternative to assessing bull prolificacy by determining parentage of all calves, progeny from only a
single week (wk 3) or a subset of weeks (wk 2, 3, and 4)
were modeled and compared to DNA sampling and testing all calves. Prolificacy assessments based on either a
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Table 2. Percent concordance between alternative prolificacy assessment methods comparing assignments of
bulls into equal-sized top (high), middle (medium), and
bottom (low) third prolificacy groups based on sampling
only those calves born in a 3-wk period (wk 2, 3, and 4)
or a single week (wk 3) of the calving season to prolificacy assessments based on sampling the entire calf crop
Reassessed using only
Original
prolificacy calves born in wk 2, 3, and 4, %
assessment1 High
Medium
Low
High
83
17
0
Medium
14
63
23
Low
0
17
83
1Based
circumference EPD provide an early indication of potential sire prolificacy and avoid environmental influences
associated with actual SC measurements. Tropical cattle
selected for high or low pregnancy rate breeding value
resulted in bull progeny with larger scrotal circumference at 18 mo of age for the higher pregnancy rate group
(Mackinnon et al., 1987). Bamualim et al. (1984) found
actual SC was correlated to pregnancy rate (r = 0.15) in
the year of measurement but was negatively correlated to
lifetime pregnancy rate (r = -0.10) and that these correlations were much lower than those for breeding soundness
scores, which were r = 0.36 and 0.47, respectively.
Although the relationship between SC EPD and bull
prolificacy was not strong, SC EPD may serve as an indicator trait for the economically relevant traits of heifer
pregnancy and stayability. Others have reported moderate
positive genetic correlations between SC EPD and these
traits in Nellore cattle (Eler et al., 2006; Van Melis et al.,
2010), although the correlation was not found to be as
strong in Bos taurus breeds (Evans et al., 1999; MartnezVelzquez et al., 2003). The relationship between SC EPD
and male reproduction as measured by prolificacy during
a natural service breeding season reported in this study
does not appear to be among those published elsewhere.
In a post hoc sense bulls appear to be somewhat
consistent in their success or lack of success in breeding
large numbers of cows. Repeatability of DNA paternity
determined prolificacy was 0.33 to 0.37 over 3 yr under
our intensive conditions with Bos taurus bulls on different ranches with different breeding seasons and 0.43 to
0.69 under extensive Australian conditions with mixed
Bos indicus bulls (Holroyd et al., 2002), suggesting applicability to wide conditions. The variation in prolificacy can partly be explained by moderate repeatability,
but other underlying reasons remain elusive.
By paternity testing only calves born in wk 2, 3,
and 4 or only from wk 3, sampling costs would be reduced to approximately 50% and 20%, respectively, of
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References
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