Theriogenology 81 (2014) 1823

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Theriogenology
journal homepage: www.theriojournal.com

40th Anniversary Special Issue

Understanding and evaluating bovine testes

John P. Kastelic*
Department of Production Animal Health, University of Calgary, Calgary, Alberta, Canada

a r t i c l e i n f o

a b s t r a c t

Article history:
Received 8 July 2013
Received in revised form 19 August 2013
Accepted 2 September 2013

The objective is to briey review bovine testes and how they are assessed, with an
emphasis on articles from Theriogenology. Scrotal circumference (SC) is the most common
method to assess testicular size; it varies among individual bulls and breeds and is highly
heritable. In general, a large SC is associated with early puberty, more sperm, a higher
percentage of morphologically normal sperm, and better reproductive performance in
closely related females. Consequently, there are minimum requirements for SC for
breeding soundness. In prepubertal bull calves, there is an early rise (1020 weeks of age)
in LH, which is critically related to onset of puberty and testicular development. Feeding
bulls approximately 130% of maintenance requirements of energy and protein from
approximately 8 to 30 weeks of age increased LH release during the early rise, hastened
puberty (approximately 1 month), and increased mature testis size and sperm production
(approximately 20%30%). However, high-energy diets after weaning (>200 days) often
reduced sperm production and semen quality. A bulls testes and scrotum have opposing
(complementary) temperature gradients, which keep the testicular temperature 2
C to
6
C cooler than core body temperature for production of fertile sperm (increased testicular temperature reduces semen quality). Infrared thermography, a quick and noninvasive
method of assessing scrotal surface temperature, may be benecial for evaluations of
breeding soundness. The primary clinical use of ultrasonography in assessment of reproductive function in the bull is characterization of grossly detectable lesions in the testes
and scrotum. In conclusion, testis size and function are critical for bull fertility, affected by
nutrition, and readily assessed clinically.
2014 Elsevier Inc. All rights reserved.

Keywords:
Bull
Testicle
Sperm
Breeding soundness evaluation
Puberty

1. Introduction
To optimize reproductive function for either natural
service or articial insemination, bulls should produce
large numbers of morphologically normal fertile sperm.
Therefore, knowledge regarding bovine testes and how
they are evaluated, is of great importance. The purpose of
this article is to briey review factors controlling gonadal
development and puberty, the importance of testis size in
breeding soundness, and mechanisms regarding testicular
thermoregulation. In addition, methods for evaluating
testicular size, temperature, and integrity are also

* Corresponding author. Tel.: 1 403 210 8660; fax: 1 403 210 9466.
E-mail address: jpkastel@ucalgary.ca
0093-691X/$ see front matter 2014 Elsevier Inc. All rights reserved.
http://dx.doi.org/10.1016/j.theriogenology.2013.09.001

discussed. Because this article is being prepared as part of

an anniversary issue of Theriogenology, emphasis is given to
publications from this journal.

2. Measurement of testicular size

A comprehensive review of bovine testicular measurements published in Theriogenology in 1979 [1], still has
considerable relevance. In that review, it was noted that
testicular weight provides an accurate estimate of sperm
producing potential [1]. Furthermore, because testicular
weight cannot be directly measured in a noninvasive way
in breeding bulls, indirect measurements of the testes (and
scrotum) have been developed, including testis length and
diameter, paired testis width, and scrotal circumference

J.P. Kastelic / Theriogenology 81 (2014) 1823

(SC) [1]. Linear measurements are done with a caliper,

whereas SC is done with a tape (various commercial tapes
are available). Although it has been suggested that using a
caliper to measure testis size is better than measurement of
SC [2], in eld conditions, measurement of SC remains the
most common approach. In that regard, SC is regarded as an
accurate predictor of paired testis weight (correlations
ranged from 0.890.95 in dairy bulls and were 0.95 in beef
bulls [1]). Furthermore, the correlation between SC and
sperm output in young dairy bulls was 0.81 [3]. However,
there are indications that shape of the testes could affect SC
and sperm production. In one study [4], 27 Holstein bulls
were categorized according to testicular shape. In that
study, bulls with long and slender testes had the lowest SC
but produced the greatest number of sperm, whereas bulls
with short, spheroid testes had the largest SC, but produced
the fewest sperm. This is an interesting observation that is
certainly worthy of follow-up study.
There were profound changes in SC with age (distinct
sigmoid curve): slow growth before 25 weeks of age, very
rapid growth during the peripubertal phase, slow growth in
mature bulls, and if bulls were retained for many years, a
subsequent decline (because of senescence) after approximately 144 months of age [1]. In addition to the effects of
age, there was considerable variation among bulls in SC;
because the average (of several studies) heritability for SC
was approximately 0.67 [1], it was concluded that rapid
progress could be made by selection of bulls with large SC.
In a subsequent study involving more than 3000 yearling
beef bulls, paternal half-sib estimates of heritability were
0.41  0.06 for age-constant scrotal circumference, 0.34 
0.06 for testicular length, and 0.37  0.06 for calculated
paired testicular volume [5]. Furthermore, it was noted that
SC might be a reliable measure of puberty and bulls with a
smaller SC were much more likely to have unsatisfactory
semen quality [1]. Finally, SC was positively associated with
growth characteristics and with conception rates in closely
related female animals [1], although in a subsequent study
[6], there was very little association between growth traits
and SC in young beef bulls.
There are substantial differences among breeds in SC. In
the 1979 review paper on assessing testicular size, it was
noted that there was a need to collect additional breedspecic SC data [1]. In a subsequent study [7], SC was
measured on 7918 beef bulls (2-year-old) of several breeds.
The authors recommendations of minimum acceptable SC
for 2-year-old bulls were: Simmental, 36.0 cm; Angus and
Charolais, 35.0 cm; horned and polled Herefords and
Shorthorn, 34.0 cm; and Limousin, 33.0 cm.
3. Puberty
Puberty in bulls is usually dened as collection of an
ejaculate that contains at least 50  106 sperm, of which at
least 10% are progressively motile [8]. Alternatively, an SC of
28 cm has been used as a measure of puberty (reviewed in
Rawlings et al. [9]).
In prepubertal bull calves, there is an early transient rise
(between 10 and 20 weeks of age) in gonadotropin secretion (particularly LH); there is ample evidence that this is of
critical importance with regard to puberty and testicular

19

development [9]. In a comparison of beef bulls reaching

puberty early versus late (41.9  0.3 and 48.3  0.7 weeks
of age, respectively), serum LH concentrations were greater
(P < 0.05) in the early maturing bulls at 12, 13, 15, 17, and
48 weeks of age [10]. In a subsequent similar study, beef
bull calves that reached puberty either early (36.6
44.2 weeks, n 12) or late (46.448.9 weeks, n 8), serum
LH concentrations in the former group of bulls were
signicantly greater at 12, 14, and 16 weeks of age [11].
Furthermore, in that study, mean serum LH concentrations
at 4, 10, and 40 weeks of age and LH pulse frequency at 10
and 20 weeks of age were negatively correlated with age at
puberty.
There is considerable interest in early-life prediction of
future reproductive events. Sexual development and
reproductive function were studied in 22 Angus  Charolais and 17 Angus bulls from 6 to 16 months of age [12]. Age,
weight, SC, and paired-testes volume were all good predictors of pubertal and mature status, with moderate to
high sensitivity and specicity (71.6%92.4%). In another
study using response to exogenous GnRH as a means of
predicting onset of puberty in bulls [13], peak serum LH
concentrations and area under the LH response curve were
lower (P < 0.05) in early- versus late-maturing bulls at 4
and 20 weeks of age. Therefore, the authors concluded that
the response of LH to GnRH in calves could be useful to
select bulls with early puberty. In another study involving
264 young beef bulls, 200-day-old calves with SC 23.0 cm
had a 95% probability of achieving SC >34.0 cm by 1 year,
whereas those with an SC <23 cm at 200 days had only a
54% probability of having an SC >34.0 cm by 1 year [14].
In a study characterizing sexual development in earlyand late-maturing Nelore (Bos indicus) and Canchim (3/8
Bos indicus  5/8 Bos taurus crossbred) bulls, earlymaturing bulls were lighter and had a smaller SC at puberty than late-maturing bulls [15]. Therefore, it was
concluded that sexual precocity was not related to lower
thresholds for body weight and SC. Furthermore, yearling
SC could be used to predict onset of puberty, whereas age,
weight, and SC were equally good predictors of sexual
maturity in B. indicus bulls.
In an attempt to hasten the onset of puberty, bull calves
were given 3 mg of bovine LH (n 6) or 4 mg of bovine FSH
(bFSH; n 6) once every 2 days, from 4 to 8 weeks of age
[16]. In this study, the age at which the bulls rst had an SC
28 cm (as a measure of puberty), occurred earlier (P <
0.05) in bFSH-treated than control calves (39.3  1.3 vs.
44.8  1.3 weeks, respectively). Furthermore, at 56 weeks,
bulls given bFSH had signicantly more Sertoli cells and
elongated spermatids and spermatocytes. Therefore,
exogenous bFSH from 4 to 8 weeks of age enhanced
testicular growth (SC), hastened puberty (based on SC
28 cm) and promoted spermatogenesis. In contrast,
although exogenous bovine LH increased serum LH concentrations and appeared to enhance spermatogenesis to
some extent, the improvements were not signicant.
4. Effects of nutrition on testicular development
In a series of experiments to determine the effects of
postweaning nutrition on reproductive development and

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J.P. Kastelic / Theriogenology 81 (2014) 1823

performance, beef bulls were fed high nutrition (80% grain

and 20% forage) or medium nutrition (primarily forage)
from weaning (67 months) to 12 to 24 months of age
(reviewed in Coulter and Kastelic [17]). Bulls fed the highenergy diet had a faster growth rate and thicker backfat,
decreases in both daily sperm production (10%30%) and
epididymal sperm reserves (30%90%), and a higher percentage of morphologically abnormal sperm (up to 50%
with a crater defect), compared with bulls fed the mediumnutrition diet. However, the two groups had similar paired
testes weight. Perhaps the increased dietary energy resulted in more scrotal fat, which reduced heat dissipation from
the testes, thereby resulted in increased testicular temperature with adverse effects on spermatogenesis and
sperm quality [17]. In another study, high-nutrition diets
(containing substantial concentrate) resulted in bulls with
approximately 10% larger SC than those fed mediumnutrition diets (forage-based), but paired testes weight
was not affected [17]; perhaps the increased SC was
because of fat.
Until recently, there was a paucity of data regarding the
effects of nutrition from birth to maturity on sexual
development, sperm production, and semen quality. In an
older study, Holstein bulls were fed various diets from 1 to
80 weeks of age [18]. In that study, bulls that received low
nutrition (w60% of total digestible nutrients [TDN] requirements) had smaller testes and delayed puberty,
whereas those that received high nutrition (approximately
160% of TDN requirements) had larger testes and earlier
puberty compared with bulls that received control nutrition (100% of TDN requirements). More recent indirect
evidence regarding the effects of early-life nutrition was
that testicular size of bulls from 2-year-old dams was
smaller than that of bulls from older dams [5], presumably
due to maternal milk production. Age adjustment factors
for yearling SC did not differ (P > 0.20) among breed groups
and averaged 0.032 cm/day. Adjustment factors for age of
dam were 1.3, 0.8, 0.4, and 0.0 cm for sons of 2-, 3-,
4-, and 5-year-old dams, respectively [5].
In a series of four experiments, beef bulls were fed
varying diets (approximately 70%, 100%, and 130% of recommended amounts of energy and protein) to determine
the effects of nutrition during calfhood (w1030 weeks)
and during the peripubertal period (w3070 weeks) on
sexual development and reproductive function (reviewed
in Barth et al. [19]). Overall, bulls fed 130% energy and
protein during calfhood had greater LH pulse frequency
early in life (<25 weeks), and at approximately 70 weeks,
larger testes and greater epididymal sperm reserves. In
contrast, bulls fed 70% of requirements had decreased LH
secretion before 25 weeks, delayed puberty, and smaller
testes with fewer epididymal sperm at 70 weeks. For
example, for bulls fed low, medium, or high nutrition from
10 to 70 weeks of age, age at puberty was (mean  SEM)
326.9  5.5, 304.7  7.4, and 292.3  4.6 days, respectively
(differences not signicant) and paired testis weights were
523.9  25.8, 552.4  21.1, and 655.2  21.2 g, respectively
(signicantly greater for high vs. low nutrition). Furthermore, it was noteworthy that in bulls fed 70% of requirements from 8 to 26 weeks of age and subsequently fed
130% of requirements, testicular development was not

improved and puberty was still delayed; therefore, supplemental feeding did not overcome earlier decits [19].
Although growth rate between 6 and 16 months did not
affect sexual development and reproductive function in
beef bulls, greater body weight was associated with earlier
puberty and maturity and with larger testes at 16 months
(correlation coeffecients approximately 0.40.6; P < 0.05)
[12]. Therefore, high-nutrition diets enhanced reproductive
function only when they were fed at less than 6 months of
age [12]. Average daily gains of approximately 1.0 to 1.6 kg/
day did not cause excessive fat accumulation in the
scrotum, increased scrotal temperature, or reductions in
sperm production or semen quality [12].
5. Testicular size and breeding soundness evaluation
SC is an integral component of a breeding soundness
evaluation [20,21]. Because SC increases with increasing
age, the minimum SC varies according to the age of the bull
[21]. Furthermore, some organizations use breed-specic
minimum standards for SC [22]. In one study involving
3648 bulls evaluated for breeding soundness [23], 76.2%
were classied as satisfactory potential breeders, with 7.1%
of all bulls deemed unsatisfactory based on an inadequate
SC. In another breeding soundness evaluation study [22]
that involved 2898 yearling beef bulls, 10% to 13% of all
bulls evaluated were deemed unsatisfactory (depending on
the minimum standard used for SC). In a retrospective
study of 205 beef herds, based on 2990 breeding soundness
evaluations, the percentage of morphologically normal
sperm was lower (P < 0.006) in bulls with an SC 34 versus
>34 cm [24]. Furthermore, cows exposed to bulls with a
smaller SC were less likely to be diagnosed pregnant (P <
0.047) and had a longer median interval from rst bull
exposure to calving (P < 0.016) than bulls with a larger SC
(overall, means of 92.2% and 304 days for pregnancy rate
and interval from bull exposure to calving, respectively).
6. Testicular temperature
It is well known that a bulls testes must be 2
C to 6
C
cooler than core body temperature for production of fertile
sperm and increased testicular temperature reduces semen
quality [25]. There are several features that regulate
testicular temperature. Scrotal skin is usually thin, relatively hairless with numerous subcutaneous blood vessels,
all of which promote heat loss [25]. Furthermore, the
scrotal neck allows testes to move away from the body,
thereby keeping them cooler [25]. The so-called testicular
vascular cone is located above each testis (Fig. 1); it is
comprised of the highly coiled testicular artery surrounded
by the pampiniform plexus, a complex venous network
[26]. The testicular vascular cone acts as a counter-current
heat exchanger, transferring heat from the artery to the
vein [26]. Scrotal sweating and panting also help to cool the
testes [26].
Several studies have been conducted on scrotal and
testicular temperatures and the role of blood vessels in
achieving these temperatures have been described. Top to
bottom differences (gradients) in temperature were 1.6, 0.4,
and 0.2
C for scrotal surface, scrotal subcutaneous, and

J.P. Kastelic / Theriogenology 81 (2014) 1823

21

temperature gradients were consistent with their vasculature. The scrotum is vascularized from the top to bottom,
whereas the testis is essentially vascularized from the
bottom to the top [29]. Regarding the latter, the testicular
artery exits the bottom of the testicular vascular cone, goes
down the testis, and at the bottom of the testis, breaks into
several branches that spread dorsally and laterally along
the posterior surface of the testis, and then enter the
testicular parenchyma [29]. Blood within the testicular artery is approximately the same temperature at the top and
bottom of the testis, but cools between the bottom of the
testis and where it enters the testicular parenchyma [29].
Therefore, the scrotum and the testis are warmest at the
origin of their blood supply (top of scrotum and bottom of
testis), but the temperature decreases further from that
point (i.e., bottom of scrotum, top of testis) [28,29].
Remarkably, these opposing (indeed complementary)
temperature gradients result in a testicular temperature
that is approximately uniform in vivo [28,29].
7. Evaluation of scrotal surface temperature with
infrared thermography

Fig. 1. Vascular cast of the spermatic cord, testis, and epididymis in a 2-yearold beef bull. The artery and vein were ushedand injected with red and
blue latex, respectively, and the tissues digested with sodium hydroxide.
Note the testicular vascular cone, consisting of the coiled testicular artery
surrounded by the venous pampiniform plexus, is above the testis. The
testicular artery exits the bottom of the testicular vascular cone, passes to
the bottom of the testis (underneath the corpus epididymis), and then is
readily visible on the posterior surface of the testis, as it branches and
spreads upwards, before entering the testicular parenchyma.

intratesticular temperatures, respectively [27]. The top of

the scrotum is the warmest and the bottom the coolest,
whereas the testes (when they are removed from the
scrotum) are cooler at the top than the bottom [28]. These

Using infrared thermography to evaluate scrotal surface

temperature in bulls with apparently normal scrotal thermoregulation, scrotal thermograms had left to right symmetry, with a 4
C to 6
C decrease in temperature from the
top to the bottom of the scrotum [30], whereas more
random temperature patterns were interpreted as
abnormal thermoregulation (testes or epididymides;
Fig. 2). Nearly every bull with an abnormal thermogram has
reduced semen quality [30]; however, not every bull with
poor quality semen has an abnormal thermogram. Therefore, although infrared thermography is useful as a
screening tool during breeding soundness evaluation of
bulls, it cannot be used in lieu of semen collection and
evaluation [30]. In one study, 30 yearling beef bulls, all
designated as acceptable based on a standard breeding
soundness examination, were individually exposed to
approximately 18 heifers for 45 days [31]. When the heifers
were examined 80 days after the end of the breeding season, pregnancy rates were similar (83% vs. 85%) for bulls

Fig. 2. Infrared thermograms of the posterior aspects of the scrotum in two beef bulls. Left: This bull has a normal temperature pattern; decreasing temperatures
from top to bottom and left to right symmetry. Right: This bull has a grossly abnormal temperature pattern, characterized by large areas of uniform temperature.

22

J.P. Kastelic / Theriogenology 81 (2014) 1823

with a normal or questionable scrotal surface temperature

pattern, respectively, but were higher (P < 0.01) than those
for bulls with an abnormal scrotal surface temperature
pattern (68%) [31].
8. Ultrasonographic imaging of bull testes
Although diagnostic ultrasonography is frequently used
(in research and clinical applications) for imaging the
reproductive tract of female cattle, it has not been used as
widely for imaging the reproductive tract of bulls [30]. In
one study, a single 3-min examination of bull testes using a
standard ultrasound scanner had no signicant effect on
testis size, sperm production, or sperm morphology [32].
Ultrasonography has been used for evaluating testes [30].
In an early study to assess the value of testicular ultrasonography, it was concluded that routine testicular ultrasound examination did not provide additional information
regarding breeding soundness compared with standard
evaluation techniques (namely, physical examination of the
reproductive tract and collection and evaluation of sperm
[33]). Although ultrasonography was a useful noninvasive
means of measuring testis size in bulls, it was no better
than SC [33].
In a study to characterize ultrasonographic echotexture
of the testis [34], 10 beef bulls were examined using ultrasonography every 2 weeks from 2 to 46 weeks of age, at
which time, all calves had an SC of 28 cm (consistent with
puberty). Image intensity of the testes increased from 6 to
8 weeks of age and approximately 20 to 46 weeks of age,
with an inection at 30 weeks (P < 0.05). Similarly, in
another study [35], testicular echogenicity increased between 20 and 40 weeks of age. In subsequent studies, beef
bulls (n 152) were examined from approximately 20 to
70 weeks of age to evaluate testicular ultrasonogram pixel
intensity during sexual development [36]. In that study,
testicular echogenicity started to increase (i.e., testis
appeared more white) 16 to 12 weeks before puberty and
peaked from 4 to 0 weeks before puberty. Although testicular echogenicity increased (i.e., the parenchyma appeared
more white) as a bull approached puberty, echogenicity was
not better than SC as a predictor of puberty [36].
Ultrasonography can be used to detect and characterize
testicular pathology. It is noteworthy that areas of increased
echogenicity (testicular brosis) are common, especially in
young bulls, but are not associated with decreased semen
quality (e.g., percentage of morphologically abnormal
sperm [30]). Unfortunately, neither visual evaluation nor
computerized pixel analysis of testicular ultrasonic echotexture were consistently predictive of semen quality in
bulls [30]. Therefore, it was concluded that the primary
clinical use of ultrasonography in assessment of reproductive function in the bull is characterization of grossly
detectable lesions in the testes and scrotum [30].
9. Discussion and conclusions
Testicular size, which is closely related to spermproducing potential, is most commonly assessed using
measurement of SC. Because SC varies substantially among
bulls and is relatively highly heritable, it is possible to

rapidly increase SC using genetic selection. In young bulls,

SC can be used to predict the onset of puberty. Feeding bulls
younger than 6 months of age, energy, and protein in excess
of minimum requirements hastened puberty and increased
mature SC and sperm production. There are minimum
standards for SC for a bull to be declared breeding-sound. In
general, bulls with a larger SC usually have a higher percentage of morphologically normal sperm. Testes must be
cooler than body temperature for the production of normal,
fertile sperm. Scrotal and/or testicular thermoregulation
can be assessed using infrared thermography, whereas lesions in the testes and scrotum can be characterized using
ultrasonography.
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