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Evaluation of bull prolificacy on commercial beef cattle ranches using DNA paternity

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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Evaluation of bull prolificacy on commercial

beef cattle ranches using DNA paternity analysis1,2
A. L. Van Eenennaam,*3 K. L. Weber,* and D. J. Drake
*Department of Animal Science, University of California, Davis 95616;
and University of California Cooperative Extension, Yreka 96097

Abstract: SNP-based DNA testing was used to assign

paternity to 5,052 calves conceived in natural service multisire breeding pastures from 3 commercial ranches in
northern California representing 15 calf crops over 3 yr.
Bulls present for 60 to 120 d at a 25:1 cow to bull ratio in
both fall and spring breeding seasons in ~40 ha or smaller fenced breeding pastures sired a highly variable (P <
0.001) number of calves (Ncalf), ranging from 0 (4.4% of
bulls present in any given breeding season) to 64 calves
per bull per breeding season, with an average of 18.9
13.1. There was little variation in Ncalf among ranches (P
= 0.90), years (P= 0.96), and seasons (P = 0.94). Bulls
varied widely (P < 0.01) in the average individual 205-d
adjusted weaning weight (I205) of progeny, and I205 varied between years (P< 0.01) and seasons (P < 0.01) but
not ranches (P = 0.29). The pattern for cumulative total
205-d adjusted weaning weight of all progeny sired by
a bull (T205) was highly correlated to Ncalf, with small
differences between ranches (P = 0.35), years (P = 0.66),
and seasons (P = 0.20) but large differences (P < 0.01)
between bulls, ranging from an average of 676 to 8,838kg

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.

Key words: beef cattle, bulls, DNA, markers, paternity, prolificacy

2014 American Society of Animal Science. All rights reserved.
J. Anim. Sci. 2014.92:26932701
In the commercial cow-calf sector, the principal determinants of income are the number of sale animals and
the value per sale animal (Garrick and Golden, 2009). In
1This work was supported by National Research Initiative Competitive
Grant No. 2009-55205-05057 (Integrating DNA information into
beef cattle production systems) from the USDA National Institute
of Food and Agriculture.
2The authors gratefully acknowledge the cooperation and labor
provided by the 3 collaborating ranches (Cowley Family Ranch, Kuck
Ranch, and Mole Richardson Farms).
3Corresponding author:
Received October 3, 2013.
Accepted March 22, 2014.

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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Van Eenennaam et al.

testicular tone, dominance, libido score, and semen

quality), but that those traits explained only 3557% of
the phenotypic variation in the number of progeny sired.
The objectives of this study were to use SNP-based
DNA testing to quantify variation in prolificacy of bulls
in multisire breeding groups using data from 3 large
commercial beef ranches in northern California, and to
investigate the potential uses of this information for management decisions in commercial beef cattle production
systems. Bull age and breed association EPD were examined to determine their relationship with bull prolificacy.
Additionally, various calf subsampling strategies were examined to determine their accuracy as as an approach to
avoid the costs associated with sampling DNA from the
entire calf crop.
Materials and Methods
Three commercial ranches, designated A, B, and C
and located in the Shasta Valley of northern California,
were evaluated for bull performance using DNA testing to determine parentage. Registered Angus sires
had been used extensively during the previous 10 yr on
these 3 ranches, making the cow herds primarily Angus.
Ranches raised their own replacement heifers.
Ranch A had a spring calving herd of 550 cows and
a fall calving herd of 350 cows. Breeding seasons were
60 d in length and included several breeding pastures
typically involving 2 to 5 bulls and a cow to bull ratio of
approximately 25:1. Breeding pastures were fenced and
generally less than 40 ha in size. Bulls used included
predominately Angus (n = 64) plus a small number of
South Devon (n = 2) and South Devon Angus cross
(n= 6) bulls bred and raised on the ranch.
Ranch B had a 200-cow spring calving herd and a
fall calving herd of about 300 cows. On this ranch breeding seasons were 90 d in length with several breeding
pastures made up of 2 to 5 bulls and a cow to bull ratio
of approximately 25:1. Breeding pastures were less than
40 ha in size. Bulls used include predominately Angus
(n = 19) and a small number of Horned Hereford (n = 2).
Ranch C had a fall calving herd of about 700 cows.
The breeding season was approximately 120 d. Breeding
pastures tended to be bigger than on the other 2 ranches,
with a larger number of bulls, 5 to 9, in each pasture,
but a cow to bull ratio of approximately 25:1 was maintained. Bulls used included predominately Angus (n =
37) and a small number of Red Angus (n = 3), Horned
Hereford (n = 1), and Polled Hereford (n = 1).
Before being joined with the cows, a breeding soundness examination (BSE) was conducted on all bulls, and
only bulls passing the exam were used. Breeding groups
consisted of replacement heifers as a single group and
mature (all other) cows. The cows were not assigned

to the same breeding group each year, but rather were

assigned on the basis of practical considerations (e.g.,
amount of feed available in different pastures) at the
judgment of the ranch manager. Cows in the various
fall or spring herds generally stayed with those breeding
herds. Bulls were observed on a daily basis or several
times per week during the breeding season and were
removed from the breeding pasture for injury or poor
body condition. When bulls were removed, replacement
bulls were most frequently obtained from other breeding groups as idle substitute bulls were not typically
available. The replacement bull selection decision was
based on the judgment of the rancher when considering
a variety of practical factors, including reassigning bulls
from breeding pastures that had a slightly lower cow to
bull ratio, selecting bulls that were observed to be very
actively breeding cows, and making selections to avoid
known bull dominance issues such as a history of observed aggressive behavior between specific individuals
and avoiding the comingling of older bulls with young
inexperienced bulls. Replacement heifers were bred to
younger (generally less than 3 yr of age), lighter-weight
bulls that had typically been purchased for calving ease.
These bulls were shifted from replacement heifer to mature cow breeding pastures as they became older and
heavier after a couple of years.
Birth dates and dam identification were obtained
at calving, and calves were individually identified (n =
5,052). Birth weights were taken only on Ranch B. At
marking time, electronic ear tags (Destron Fearing, St.
Paul, MN) were applied to calves, and hair samples were
obtained for DNA testing. Calf genotypes (~100 SNP)
were obtained as part of the Bovine SeekSire genotyping service (GeneSeek Inc., Lincoln, NE). Parentage
SNP genotypes for bulls were extracted from Bovine
SNP50 BeadChip genotypes (Illumina, San Diego, CA),
which had been acquired for these bulls in the course of
a previous project. Bull genotypes were compared with
calf genotypes using the SireMatch program (J. Pollak,
U.S. Meat Animal Research Center, Clay Center, NE).
Bull prolificacy was defined as the number of calves assigned to each bull by DNA paternity testing (Ncalf) for
an entire calf crop.
Individual calf weights were obtained at approximately 205 d of age. Weights were adjusted using Beef
Improvement Federation (BIF) linear adjustments for
cow age and calf age (BIF, 2010), although some of the
calves fell outside the recommended age range on weigh
day because of practical constraints associated with
calves going to summer pastures such that they were not
accessible for weighing during the BIF prescribed age
range. Weights for each calving group were obtained
on consecutive days when it was not possible to weigh
the entire group on 1 d. Individual 205-d weights (I205)

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Commercial beef bull prolificacy

were also adjusted for sex differences by least squares

using a model that included fixed effects of ranch, year,
season, and calf sex. Total adjusted 205-d weight of
progeny (T205) for each bull was defined as the sum of
the I205 of all his progeny for each calf crop. Prolificacy
of young bulls (age less than 3 yr) was defined as that
observed for their first breeding season and was compared to the average of their prolificacy recorded in subsequent breeding seasons
Prolificacy and EPD Relationships
To determine the relationships between American
Angus Association EPD and bull means for Ncalf, I205,
or T205, these variables were evaluated using scatterplots for each EPD. Because of the use of some young
low-accuracy bulls, all analyses were restricted to bulls
with BIF EPD accuracies greater than 0.05 for the trait
being evaluated. Average phenotypic values that showed
a probable linear relationship with EPD (P < 0.20) using
linear regression were selected for further evaluation.
Prolificacy and Calving Distribution
The beginning of the calving season (d 1) for each
calf crop was assigned as the date the fifth calf was
born on a given ranch. Birthdates were grouped into
7-d intervals representing weeks of the calving season.
Bulls were categorized into 3 equal-sized prolificacy
groups, high (HP), middle (MP), and low (LP) prolificacy, based on the total number of progeny produced in
each calf crop. Additionally, the prolificacy grouping of
young bulls (age less than 3 yr) for their first breeding
season was compared to their average prolificacy grouping in subsequent breeding seasons.
Alternative Prolificacy Assessment Methods
As a less costly alternative to parentage-testing all
calves for assessment of prolificacy, smaller subsets of
calves were used to sort bulls into prolificacy groups HP,
MP, and LP. Alternative subsets included calves from a single week (wk 3) of the calving season or the sum of calves
from 3 wk (wk 2, 3, and 4). The original prolificacy assessments based on the entire calf crop were compared to the
assessments based on these subsets using a chi-squared test.
Statistical Analyses
Analyses of Ncalf were restricted to bulls present for
the full duration of the breeding season and were analyzed using a model that included fixed effects of ranch
(R), year (Yr), and season (Sn). The contributions of
Ncalf and I205 to T205 were estimated by comparing R2

from regressions with T205 as the dependent variable

and inclusion of Ncalf and I205 together or singularly
(Systat). Correlations were estimated with Pearson correlations. Ncalf repeatability for bulls that were present
for more than two seasons was calculated for all bulls,
and for mature bulls older than 3 years of age using the
method of Lessells and Boag (1987), which accounts
for among-group and within-group variance for unequal
group size. Effect of bull age on Ncalf was evaluated using the linear mixed effects model (LMER) function of
R and a model that included fixed effects R, Yr, Sn, and
bull age and random effect of individual bull.
Linear regression of selected breed association EPD
and average prolificacy were evaluated for P levels.
Individual bulls were examined in scatterplots, and suspect outliers based on a very high Cooks distance (Di)
and studentized residual (P < 0.05) were removed from
the analyses if their inclusion alone obfuscated a statistically significant linear relationship that was present in
the absence of that data point.
The mean number of calves born each week for bulls
with 2 or more calf crops was calculated by least squares
using a model that included fixed effects of R, Yr, Sn, and
week (W), and mean separation significance was corrected by Bonferroni adjustments. Mean weekly calving rate
for each group was calculated by least squares using a
model that included fixed effects of R, Yr, Sn, W, prolificacy group, and prolificacy group by week.
Prolificacy for the first breeding season for young
bulls was compared to the average value in subsequent
breeding seasons by regression. First season prolificacy
grouping (HP, MP, or LP) was compared to the average
grouping in subsequent breeding seasons by Pearson,
rank order correlation, and chi-squared analyses.
Because more than 20% of the cells in the chi-squared
analysis were below 5, a likelihood ratio chi-squared
analysis was also conducted.
Paternity Assignments
A total of 5,052 calves were assigned paternity on the
basis of DNA from 15 calf crops and 275 bull natural breeding opportunities (Table 1). Reproductive failure, meaning
that no calves were produced, occurred in 4.4% of the bull
seasons (12 out of 275 bull breeding season opportunities).
In 40% of the calf crops at least 1 bull sired only 1 calf and
at least 1 bull sired more than 50 calves. DNA information
was unable to uniquely assign paternity to an average of
3.8% of progeny across all ranches, with 2.6% on Ranch A,
3.0% on Ranch B, and 5.7% on Ranch C.

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Van Eenennaam et al.

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

A, B, C
A, B, C
A, B, C
A, B, C
A, B, C
A, B, C


Calf crop

No. of

Minimum bull Maximum bull Mean bull age

age, yr
age, yr
SEM, yr
4.3 0.3
3.5 1.4
4.3 0.9
3.9 1.2
4.7 1.2
4.2 1.8
4.3 0.3
4.3 0.3
3.0 1.1
4.3 0.3
4.3 1.4
4.6 1.6
4.3 1.2
4.6 1.7
5.5 0.9
4.1 0.7
4.8 0.5
3.9 1.6
4.3 0.3
4.3 1.0
4.2 0.6
4.0 0.7
4.5 0.2
4.4 1.7

no. calves

Per bull
Minimum no. Maximum no. Mean no. calves
19.9 3.8
19.6 18.2
19.8 3.8
18.4 22.1
24.2 4.7
16.8 14.5
16.7 10.0
21.8 9.3
16.5 7.4
20.2 12.9
26.5 14.4
22.8 6.2
20.3 3.0
19.9 3.8
14.4 5.7
18.6 6.0
19.9 3.8
21.1 13.2
19.9 3.8
20.1 1.9
19.7 7.7
20.5 12.2
19.2 5.0
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.

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-

d weight of progeny per bull was similar across ranch

(4,687 1,890, 5,713 1,077, and 3,425 1,404kg;
P = 0.35), year (4,762 388, 4,410 368, and 4,654
386 kg; P = 0.66), and season (spring, 4,882 414 kg;
fall, 4,336 304 kg; P = 0.20) but varied widely between bulls (P < 0.01), ranging from mean totals of 676
to 8,838 kg. The mean number of calves per bull was
highly related (P < 0.01) to total production, explaining 96.9% of the variation (R2= 0.969), with each calf
contributing 220 2.6 kg. Mean I205 was also related to
total production (P < 0.05) but by itself explained only
2% of the variation (R2 = 0.02). Similarly, Ncalf was
highly correlated (r = 0.98) to T205 per bull compared
to the lower correlation (r = 0.15) to I205.
Prolificacy and Bull Age. There was no significant relationship between Ncalf and bull age. As shown in Fig. 2,
bulls of increasing age tended to have a higher maximum
Ncalf, resulting in a higher variance in Ncalf. Young bulls
in their first breeding season (N = 24) ranged in age from
1.4 to 2.9 yr (mean of 2.4 0.3 yr). The mean number of
calves per young bull ranged from 1 to 40 (mean of 14.9
9.8). Prolificacy in subsequent breeding seasons was positively linearly related (P < 0.05) to first breeding season
prolificacy, explaining about 20% of the subsequent varia-

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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).


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).

Carcass weight and the $Beef (BE) index value were

also both negatively related (P < 0.05) to prolificacy,
likely because CW is a component in BE and is therefore highly correlated (r = 0.81) to it. Additionally, when
CW and BE were regressed together, only CW was significant. When both SC and CW were regressed together
against prolificacy, only SC (P < 0.05) remained significant compared to CW (P = 0.28). Scrotal circumference
and CW were not correlated (r = 0.01).
Regression responses similar to those seen for Ncalf
were seen for SC and CW when regressed on T205, where
T205 = 7,723 + 4,189 (1919) SC (P = 0.04,
R2 = 0.13, SE = 4,345),
T205 = 14,097 179.7 (77.7) CW (P = 0.03,
R2 = 0.10, SE = 4,987).
The similar regression responses for T205 and Ncalf
were not surprising because of the high correlation (r=
0.98) between these 2 variables. No other EPD examined were related to prolificacy.
Individual 205-d weight was positively correlated with weaning weight (P < 0.01), yearling weight
(P < 0.01), CW (P < 0.01), $Feedlot index value (P
< 0.01), and BE (P< 0.01) EPD but not to any other
EPD. Individual 205-d weight was not highly correlated with either Ncalf (r = 0.22) or T205 (r = 0.26).
The importance of Ncalf compared to that of I205 on
T205 is demonstrated by comparing the R2 (0.9793)
for the regression of Ncalf (P < 0.01) on T205 to the
R2 (0.9812) for the combined regression of Ncalf (P <
0.01) and I205 (P= 0.09) on T205, showing the very
small improvement resulting from the inclusion of I205
in the regression.

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Van Eenennaam et al.

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

single week or the sum of several weeks were closely

related (P < 0.01) to prolificacy assessment based on
the total calf crop. No HP bulls were reassessed to LP
using the subset of calves born in wk 2, 3, and 4, and
only 1.4% were reassessed using only those born in wk
3 (Table 2). Between 17% and 19% of the HP bulls were
reassessed to MP using the subsets. Nonetheless, either
of these reduced assessments (wk 2, 3, and 4 or only wk
3) offers a reduced sampling (and thus cost) method of
determining prolificacy. Reassessment of LP bulls to HP
also occurred at a low rate (3% for both subsets).
DNA testing can be used to accurately assign paternity and has been documented here and by others
(Holroyd et al., 2002; Van Eenennaam et al., 2007;
Gomez-Raya et al., 2008), providing an opportunity to
investigate bull performance in commercial multisire
breeding pastures. We have shown that prolificacy is
the main driver of bull productivity as measured by total
weight of calves weaned per bull.
The results of bull prolificacy studies conducted in
varying environments and management systems consistently reveal that prolificacy varies considerably among
individual bulls. Holroyd et al. (2002) also used DNA
paternity in multisire commercial ranches but with Bos
indicus or Bos indicus cross bulls in extensive conditions in northern Australia and found very similar results
to our temperate intensive production systems, including
a 6% frequency of reproductive failure, i.e., bulls that
sired no calves in a given breeding season, compared to
the 4.4% observed in the current study. They found several phenotypic traits related to bull performance with
little practical application because of the limited amount

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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
prolificacy calves born in wk 2, 3, and 4, %
assessment1 High

Figure 5. Bulls categorized into 3 equal-sized groups (high, medium,

and low prolificacy) based on their total number of progeny in each calf crop
had different calving distributions with implied differences in breeding and
conception distribution. Error bars represent SEM.

of variation explained by these traits. Bamualim et al.

(1984) found bull age (between 2 and 5 yr of age) was
not significant on pregnancy rate of Bos indicus cross
bulls; however, they may not have had sufficient cow
numbers to adequately challenge bull fertility capacity.
A significant relationship was found between SC
EPD and bull prolificacy in the current study, but no other
breeding group management activities explained a significant amount of variation in calf output. Scrotal circumference measurements have been previously associated with
Ncalf (Coulter and Kozub, 1989), although Holroyd et al.
(2002) generally found no relationship between actual SC
measurements and prolificacy with the exception of 5/8
Brahman bulls. This relationship between SC EPD and
prolificacy during a natural service breeding season has not
been previously reported.
Scrotal circumference EPD have been positively associated with sperm motility and total BSE score (Moser
et al., 1996). Favorable influence of scrotal circumference and SC EPD on heifer maturity has been reported
(Brinks et al., 1978; Toelle and Robison 1985; Smith et
al., 1989; Moser et al., 1996; Martnez-Velzquez et al.,
2003). Scrotal circumference is also a component of the
breeding soundness examination that has been related
to bull fertility (Kealey et al., 2006), and SC estimates
testicular tissue volume, which impacts semen quantity.
The most valuable SC measurements are those taken at
approximately 1 yr of age.
Difficulties in assessing fertility in both male and female cattle are well documented, and genetic improvement is further hindered by the low heritability of fertility traits (<0.15; Mackinnon et al., 1990). Bull and cow
fertility are low but positively correlated (r = 0.16), suggesting indirect trait selection in males for fertility traits
could be an approach to improve female fertility. Scrotal

Reassessed using only

calves born in wk 3, %
High Medium

on sampling entire calf crop.

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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Van Eenennaam et al.

that required to sample the entire calf crop in this study.

Because of age and source verification marketing, birthdates are often known, and samples could be collected
at marking time from a designated group of calves. Even
without whole-herd birthdate records, strategic planning
during the third week of calving could include recording or marking calves born during that time period for
later sampling. Given the moderate repeatability of prolificacy, this type of reduced sampling could provide an
approach to identify bulls with the greatest likelihood of
being either highly or lowly prolific. The costs involved
in DNA collection include not only the costs of the tests
(currently ~$15/head for parentage testing;, accessed January 14, 2014), which are likely
to continue to decrease in the future, but also the costs
associated with unique animal identification, labor to
process each animal and collect the DNA sample, and
costs required to manage the records and integrate the
DNA information back into herd management decisions.
The value of parentage information needs to outweigh the costs of genotyping. One study examined the
value of DNA paternity identification on commercial
beef cattle operations. The assumption of their model,
based on 15 microsatellite loci, was that the information would be used to cull bulls that were producing
low weaning weight calves (Gomez-Raya et al., 2008).
Although this might be important if all bulls are producing an equal number of progeny, as can be seen from
the data in this study, some of the bulls that produced
bulls with the lowest weaning weight were siring a large
number of calves and hence were not the least profitable
bulls. The benefit that could be derived from these parentage data would conceptually be the removal of low
prolificacy bulls and increasing the cow:bull ratio of the
more prolific bulls, thereby saving on the costs associated with maintaining an inactive or low prolificacy bull.
Using the data obtained in this study, it can be estimated that if the entire calf crop were sampled to obtain prolificacy estimates, then the cost per bull to obtain
prolificacy data would be approximately 20 times the
cost of the test (1 bull + 19 offspring), or $300/bull in
the case of a $15 test. Less expensive alternative sampling strategies could be envisioned, including sampling
all bulls and only those calves born in wk 3 (~20% of the
calf crop) or, alternatively, sampling only those offspring
produced by young sires in their first breeding season
based on the observation that young bulls categorized as
either HP or LP tended to remain in those categories in
subsequent breeding seasons. However, given the small
number of young bulls involved in this study (8 each initially categorized as HP and LP), care should be taken in
over interpreting these results. In addition to prolificacy
data, the DNA information could also be used to calculate genetic merit estimates of these commercial bulls

and identify those producing superior or problematic

classes of calf (e.g., high birth weight calves).
These results reveal that highly prolific bulls sire a
disproportionately large number of the preferred more
valuable, early born calves (Funston, 2012) in well-managed herds that have a large numbers of females cycling
early in the breeding season. In self-replacing beef systems, replacement heifers are often selected on the basis
of age to enhance the potential for conception early in
their first breeding season. This study showed that only
a small percentage of such replacement heifers would
likely be sired from LP bulls, providing indirect selection on male fertility. Using DNA paternity assignment
to evaluate the relationship between heifer fertility and
sire prolificacy would provide information of economic
interest given the high costs of raising replacement heifers and the overriding importance of fertility to the beef
enterprise (Melton et al., 1979; Melton 1995).
The DNA paternity testing of the calf crop can be
used to determine sire prolificacy, and testing of fewer
calves offers opportunities for reduced costs. The number of calves sired by natural service bulls in multisire
breeding pastures is highly variable between bulls but
similar across ranches and was positively associated with
SC EPD. Varying prolificacy of herd bulls has implications for the genetic composition of replacement heifers,
with the genetics of those highly prolific bulls siring a
lot of calves likely to be disproportionately represented
in the replacement heifer pool. To be cost-effective, the
costs of parentage testing need to be recouped by the
value derived from this resulting information. One such
use might be the cost savings associated with the removal of low prolificacy bulls, although the feasibility of this
approach would depend on the continued ability of the
more prolific bulls in one year to be able to successfully
service an increased cow:bull ratio in the following year.
Prolificacy was found to be moderately repeatable (0.33)
in this field study of commercial herd sires.
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