BULL INFERTILITY 0749-0720/97 $0.00 + .

20

BULL LIBIDO jSERVING

CAPACITY
Peter J. Chenoweth, BVSc, PhD, MACVS

Cattle, in common with other food animal species, show polygynous

mating behavior, a system that allows individual males to impregnate a
number of females. 75 Modern production systems, such as those gener-
ally employed in the beef cattle industry, rely primarily on natural
breeding39 despite higher male-to-female breeding ratios and shorter
breeding periods than those typical for wild cattle. This places greater
emphasis on factors important for male reproductive success: sperm
output, semen quality, and male sex drive (or libido). Those factors
associated with semen quality and sperm production may be assessed
with the breeding soundness examination (BSE), a procedure that does
not, however, routinely assess sex drive. In many cases, beef bulls do
not receive any form of assessment prior to sale or use. 39

DEFINITIONS

Sexual behavior in the bull may be defined as that behavior associ-

ated with the detection and service of estrous females. Libido, or sex
drive, is defined as the "willingness and eagerness" of a bull to attempt
to mount and service; mating ability refers to the ability of the bull to
fulfill this aspiration. 19 Serving capacity is a measure of the number of
services achieved by a bull under stipulated conditions5 and thus in-
cludes aspects of both libido and mating ability. Reaction time is the
time that elapses between male cognition of an appropriate stimulus
and the completion of copulation. 26

From the College of Veterinary Medicine, University of Florida, Gainesville, Florida

VETERINARY CLINICS OF NORTH AMERICA: FOOD ANIMAL PRACTICE

VOLUME 13· NUMBER 2· JULY 1997 331

332 CHENOWETH

Cattle Reproductive Behavior

The major sexual attractant for bulls is female mounting activity.

Bulls are primarily attracted to the inverted "U" structure that resembles
the rear end of a stationary female. When this exists, whether or not the
female is in estrus is largely irrelevant26; in fact, steers are often used
successfully as mount animals in semen collection centers. Pheromones
also playa role in allowing bulls to detect receptive females, although
this mechanism evidently requires close physical contact for activation
in cattle. 48 The major special sense used by bulls to detect estrous females
is vision. 22, 38, 41
The task of identifying estrous females is facilitated by the tendency
of females in both late proestrus and estrus to form a mobile sexually
active group (SAG), which usually stays within visual contact of the bull
or bull group.19, 88 Females in heat generally become more active and
vocal than at other phases of the estrous cycle. Bulls tend to be most
attracted to females who are newly in estrus; provision of a fresh
stimulus can restore libido in satiated males. 42, 74 Bulls test the receptivity
of females by making real or sham mounting attempts, by chin resting,
and by licking and sniffing around the female's perineal region. The last
actions are often followed by a characteristic curling of the upper lip,
termed the flehmen response, an action that is most probably associated
with the transfer of fluids to the vomeronasal organ and their assessment
for pheromonal activity.48
The female exerts considerable control over mating by determining
the timing of sexual access and even the identity of the successful
male(s). Bulls are polygynous and form hierarchical groups, with social
status influencing reproductive success. 19, 74 Pre stimulation of males in-
creases their sexual response. 57 For young bulls, a learning process is
often necessary before competent and confident mating ability is
achieved. 12, 23 Bulls have a fibroelastic penis and copulation generally
occurs rapidly (1 to 2 seconds) once intromission is achieved. 81 Bulls are
capable of short bursts of great reproductive activity, dependent on their
inherent sex drive and stimulus pressure. Breeding trials with both
natural and induced-estrus females indicate that it is not uncommon for
bulls to serve more than 30 times within a 24-hour period.

TESTS FOR BULL SEX DRIVE

Libido, or sex drive, is a behavioral trait that is measurable. Testing

procedures for bulls generally rely on the exploitation of several or more
of the following findings, some of which have already been discussed.
1. Libido in bulls has a significant genetic component. 4, 6, 12, 25, 47, 66
2. Bulls are polygynous and tend to distribute their services among
receptive females. 5
3. The greatest single stimulus for a bull to attempt to mount and
 BULL LIBIDO/SERVING CAPACITY 333

service is the immobile rear end of a female, or something similar

in appearance. 9, 26, 86
4. Prestimulation of bulls increases their sexual response. 9,57
5. Competition among bulls can increase their sexual response. 39,59,61
Many procedures have been used to assess sex drive in bulls and
other male livestock. 19, 20 Although the simplest method to assess this
trait would appear to be observation of male sexual activity during
pasture breeding, this approach has generally yielded disappointing
quantitative results. Within the constraints of current knowledge, the
use of a formalized, pen-type testing regimen appears necessary for the
generation of comparative, quantitative results. Initial work on bull
sexual behavior arose from difficulties in maintaining sex drive in dairy
bulls in artificial insemination (AI) centers.ll Later work emphasized the
role of sexual behavior in optimizing sperm harvests. 1, 42 In Sweden, a
testing procedure (the "libido index") was devised to assess both libido
and mating ability in AI sires,47 and this procedure was modified to
become the "libido score" test for assessing range-type beef bulls. 67
These latter authors placed individual, prestimulated bulls each with an
unrestrained, estrogenized female in two adjacent pens for a short dura-
tion (5 minutes). This was modified into an expanded 0 to 10 scoring
system,17 which described degrees of sexual interest (including service)
within a 10-minute test.
A "serving capacity" test was developed in which bulls were group-
tested with restrained, nonestrous females at bull-to-female ratios (BFRs)
of either 5:2 or 5:3 for 40 to 60 minutes. 5 A "serving capacity score" was
given based on services performed only. A comparison of reaction time,
serving capacity score and libido scores has been conducted in young Bas
taurus bulls. 26 In this study, restrained females were equally attractive to
bulls whether they were in induced estrus or not, and a 10-minute test
provided as much comparative information on bull sex drive as did a
30-minute test. The use of estrous females may also be unnecessary for
the assessment of sex drive of Bas indicus bulls, providing females are
adequately restrained. 65 A composite procedure employing elements of
both the libido and serving capacity score systems has been described,ls
and procedures exploiting various features of these tests have been
used.13, 37, 52, 69, 76

PREDICTABILITY AND REPEATABILITY OF TESTS

The ideal test for the assessment of sex drive in bulls would be
simple, quick, highly repeatable, predictive of reproductive performance,
and esthetically acceptable. At present, no single current procedure
meets all of these criteria. However, relative differences in sex drive
between bulls can be reliably estimated?4 For example, moderate pheno-
typic correlations (r = .67 and .60, respectively) were obtained between
libido and serving capacity scores in yearling bulls tested on different
334 CHENOWETH

days,26 although reaction times to service in the different tests were not
significantly correlated. With Bos indicus bulls, the repeatability of libido
scores was relatively low (r = 0.44),72 although this procedure was
regarded as being superior in repeatability and logistics to the serving
capacity method for assessing sex drive in mature Bos indicus bulls. 65
A disadvantage of the serving capacity method was that it yielded
no comparative information on the degree of sexual interest shown by
bulls who did not complete service; a particular concern with young,
inexperienced bulls. When 26 yearling Bos taurus bulls were assessed a
total of eight times (two tests per day on four occasions within 2 months)
for libido and serving capacity scores, four tests were required before
test results did not differ (W. E. Berndtson, personal communication,
1986). A learning process can occur with young, inexperienced bulls,
and this can affect test results. This phenomenon has been observed in
several trials 13, 26, 39 in which young bulls obtained low serving capacity
scores that improved with subsequent mating experience. One study13
suggested that young virgin bulls who show poor serving capacity
should be offered sexual experience and then be retested to determine
their true worth. Another study 15 suggested that virgin Santa Gertrudis
bulls younger than 2 years of age might not display their inherent
serving capacity in such testing environments.
In general, best success has been obtained when assessments of
libido or serving capacity are used to place bulls into categories or
groups. Eight Hereford bulls maintained their relative ranking for both
libido scores and fertility when assessed at both 16 and 40 months of
age. 53 In another study,9 high correlations (r = .82 to .91) were obtained
for rankings in mating activity between simulated pasture tests and
subsequent pen tests of 12 Bos taurus bulls.

FERTILITY RELATIONSHIPS

Cattle fertility is multifactorial, with involvement of both male and

female factors. From the male aspect, good evidence exists that bull
libido is one of the more important contributing factors. For example, in
one study better first-cycle pregnancy rates were obtained in heifers
mated with higher serving-capacity bulls than in those mated with bulls
of low serving capacity? More recently, differences in pregnancy rates
were demonstrated between high, medium, and low serving-capacity
Hereford bulls.lO Other studies have shown advantages in herd fertility
by using higher sex-drive bulls. 54, 58, 62 Close relationships were reported
between bull rankings for fertility, libido score, and testosterone re-
sponse to gonadotropin-releasing hormone (GnRH)?3 In a Florida study,
bull libido and semen quality both significantly influenced pregnancy
rates achieved by naturally mated Brangus bulls, with libido having the
most effect (A. C. Warnick, personal communication, 1990).
Other studies have indicated either that bull libido assessment pro-
vided greater prediction of bull fertility than did semen assessment
 BULL LIBIDO /SERVING CAPACITY 335

alone,B3 or that the BSE assessment alone was lacking in its ability to
predict bull fertility.64 Using multisire mating and progeny identification
by blood typing, the number of services performed in libido/serving
capacity tests was positively correlated with fertility up to a certain level
(approximately four services), above which fertility actually declined
with subsequent services. 31
However, other studies have revealed poor or inconclusive relation-
ships between bull libido / serving capacity assessment and herd fertility
or pasture performance. 13, 30, 33, 36,59,72 In some studies, although higher-
libido bulls serviced more often and serviced more females than did
lower-libido bulls, significantly more pregnancies did not result.13, 36, 39
The reasons for these apparently contradictory findings may be
sought in differing approaches and methods. In some trials, bulls were
not placed under sufficient breeding stress to demonstrate real differ-
ences. These differences may have been more apparent with use of
higher BFRs or shorter breeding periods. In addition, bulls of potentially
low fertility were excluded from cooperative breeding trials. Investiga-
tors have sought the "holy grail" of demonstrating that a single trait
(e.g., bull sex drive) would have a consistent, decisive influence on
herd fertility-an approach that is invariably doomed to failure. Factors
associated with BSE (scrotal circumference, sperm motility and morphol-
ogy) can separately influence fertility63 and are apparently not linked
with sex drive. 13, 25, 27, 28, 62 Bulls may be superior in one trait or several,
but their fertility can be compromised by deficiencies in others.
This was demonstrated in one stud y 36 in which 92 beef bulls were
classified as satisfactory or questionable for BSE and as high (score 9 to
10) or medium (score 7 to 8) in libido prior to single-sire mating with
groups of estrus-synchronized heifers. Here, pregnancy rate was 9.10/0
higher for satisfactory BSE bulls than for questionable BSE bulls, but did
not differ between bulls of high and medium libido score, even though
high-libido bulls serviced more females and served more times than did
medium-libido bulls. This paradox apparently occurred because a lower
percentage of serviced females in the high-libido group became preg-
nant. Differences between bulls in sex drive were clearly masked by
differences in their BSE components.
The ability of bulls to service females is related not only to their
inherent sex drive, but also to their mating ability. Problems in mating
ability may be due to a number of physical and pathologic causes
including skeletal and penile abnormalities,2° and these may contribute
significantly to bull infertility. The libido/serving capacity test procedure
can play an important role in the detection of such problems.

FACTORS INFLUENCING BULL SEX DRIVE

Age and Rearing Effects
Age and/or experience of bulls can influence their mating ability
and thus their perceived level of sex drive. Mating ability does appear
336 CHENOWETH

to have a learned component in bulls,12, 13, 29, 49, 71, 82 even though exposing
young Polled Hereford bulls to heifers postweaning did not influence
their subsequent libido or mating ability.51 In this trial, individually
penned bulls initially showed greater serving capacity than did group-
penned bulls, but this difference did not persist. Male-male mounting in
group penned bulls was not indicative of libido, serving capacity, or
mating behavior with heifers. Social restriction of young bulls was
therefore not detrimental to their mating ability.
In young tropical beef bulls, libido score increased with bull age
between 16 and 31 months of age. 69 Bull age affected sexual behavior
traits in crossbred bulls, with yearling bulls showing lower libido and a
higher proportion of mounts than did older bulls. 31 In Florida, sexual
performance assessments generally increased with age in young (12 to
24 month) Bos taurus bulls, although not in Bos indicus bulls. 29 The latter
group, however, generally displayed a lower level of sexual activity
than did the former. More information is needed to differentiate the
effects of age and experience on bull sexual behavior from those effects
caused by environmental and managerial influences.
In this respect, prolonged nursing was considered to retard, or
compromise, the expression of normal sexual behavior in Angus bulls32
as did the feeding of high concentrate levels in crossbred bulls. 46 How-
ever, postweaning dietary energy levels were not found to affect sex
drive in young bulls of synthetic breeds. 63 Negative relationships were
obtained between sex drive and production traits (average daily gain
and final test weight) in yearling beef bulls in one study,66 whereas
underfeeding had no adverse effects on bull sexual behavior in another.87
Zebu bulls raised on open range exhibited slower sexual responses
compared with those reared more intensively,56 although no permanent
sexual inhibitions were attributable to rearing methods in bulls. 54 How-
ever, temporary sexual inhibitions in bulls may compromise pregnancy
rates in herds that have restricted breeding seasons. 19

Bull-to-Female Ratio

One study reported that most bulls achieved good reproductive

efficiency when used at BFRs of both 1:44 and 1:60, with no consistent
effect being attributable to either BFR or bull grouping. 80 Individual bull
reproductive capability had more effect on reproductive success than
did either BFR or single versus multisire breeding combinations. Com-
parison of the reproductive performance of young bulls mated with
estrus-synchronized females at BFRs of either 1:20 or 2:40 indicated that
single-sire mating was most efficient,35 whereas no difference occurred
in calving rate when Angus and Polled Hereford and Santa Gertrudis
bulls were mated either singly or in pairs at a BFR of 1:40. 35 In studies
employing bulls with estrus-synchronized females, one suggested low-
ered fertility at a 1:50 BFR,44 whereas in another pregnancy rates were
more influenced by individual bull fertility than by BFR.70 Several stud-
 BULL LIBIDO/SERVING CAPACITY 337

ies have indicated that BFR is not an important consideration when

conducting tests for sex drive in young, experienced bulls/62,76 provided
group sizes are relatively small. One study obtained better results when
bulls were tested individually/51 although intermale aggression and inter-
ference in the group tests may have influenced these results. Such
interference occurs more often when the spacing between stimulus fe-
males is too small, or when bulls compete for female access, and it may
be detrimental when bulls are tested in groups consisting of more than
two bulls. 26

Social Effects

Social ranking of bulls within groups can influence their sexual

activity.8, 19,34,80 Dominance is expressed more strongly and linearly in
older bulls (those 3.5 to 4 years of age or older) and appears to be more
related to seniority than to age or body weight. 8 The effect of social
interactions among bulls on herd fertility may be greater at lower BFRs
than with higher levels of breeding stress. 8
Dominance rank was negatively correlated with sex drive in one
study of yearling bulls. 66 If dominance and sex drive are different traits,
then the dominant bull (or bulls) could impair herd fertility both through
failure to service females and by preventing less dominant bulls from
serving. Evidence exists for such effects in extensive beef operations/55
in which the social dominance ratio of bulls had some relationship with
herd fertility. Such effects are probably most evident when older and
younger males are combined in the breeding pasture/8 although mixing
different bull genotypes can apparently cause similar effects. 21
Social effects may, however, also benefit bull sexual behavior. Such
an effect was observed in one study in which greater sexual activity
occurred when young, inexperienced bulls were tested in groups (3 to
5) than when they were tested individually. 54 Similarly, yearling bulls of
higher serving capacity achieved more services in double-sire tests than
when tested alone. 39 Beneficial social effects on bull sex-drive expression
include the observations that both prestimulation of bulls and competi-
tion among them can positively influence test results. 23

Genetic Effects

Reports indicate that dairy breeds may be more sexually active than
beef breeds1, 40 and that Bas indicus bulls generally show lower, and more
variable, levels of libido than do Bas taurus bulls. 2, 24, 29, 43, 50, 79, 85 A number
of such breed differences are probably attributable either to genotype-
environment influences or to breed idiosyncracies in response to testing
procedures. 43, 69,72 In several studies in tropical Australia, Brahman and
Brahman-crossbred bulls obtained the lowest libido scores and Afri-
cander bulls and their crosses achieved the highest, whereas European-
338 CHENOWETH

bred bulls were intermediate. 24, 69 In US studies, higher sex-drive scores

were also obtained for Bos taurus bulls than for Bos indicus bulls. 29, 43, 77
Despite these results, a comparison of trials in which bulls were
placed with estrus-synchronized females indicated that Bos indicus-
derived bulls were as efficient as European-breed bulls in detecting,
serving, and impregnating estrous females, despite a lower service rate. 23
This discrepancy may be partly explained by observations that Bos
indicus bulls tend to be selective and shy breeders, and that they gener-
ally do not perform well in pen tests to assess sex drive/9 even though
they can be very active and efficient detectors of estrus in the pasture.
In Florida, Bos taurus (Angus, Hereford) bulls obtained superior results
in sex-drive tests compared to tropicalized Bos taurus (Senepol, Romosi-
nuano) bulls, with Bos indicus (Brahman, Nellore X Brahman) bulls
generally obtaining the worst results. 29 Commonly used testing proce-
dures for sex drive may disadvantage Bos indicus bulls. Modifications
that have been suggested to improve the performance of such bulls
include the use of unrestrained estrous females and the avoidance of
extraneous distractions during the test. 21
A number of studies indicate that strong microgenetic influences
occur with bull sex drive. In Scandinavia, monozygous twin bulls raised
on differing nutritional regimens displayed greater similarity in mating
behavior and temperament within pairs than between pairs,4 suggesting
strong genetic influences on these traits. Paternal half-sibs of Swedish
bulls differed significantly in libido with greater variation between sire-
son groups than within them. 47 Studies showing that crossbred bulls
generally exhibited higher sex drive in pen tests than did their parental
purebreds indicate that genetic effects, in this case heterosis, influence
bull sex drive. 24,69 In Colorado, line of breeding (inbred lines or crosses
among inbred lines) was an important source of variation in bull sex
drive, indicating that lines had previously been selected or differentiated
on the basis of sexual behavior.25 Similarly, differences in libido scores
were observed between breeding lines and sires-within-lines in young
bulls of British breeds. 66 Here, high sex drive was not synonymous with
either superior production traits (average daily gain or final test weight)
or high social ranking. Sire strongly influenced serving capacity in young
Angus bulls. 12
A number of studies have indicated that traditional measures of
breeding soundness such as scrotal circumference and semen traits are
not significantly correlated with sex-drive estimates in bulls,6, 13,25, 62
indicating that these are separate traits. A heritability estimate of 0.59 ±
0.16 was obtained for serving capacity in a study of 157 paternal half-
sib bull groups in Australia. 6 In this study, the inclusion of body weight
as a covariate did not alter the result, and serving capacity was not
associated with temperament rating. This differed from a previous study
in which the "sexual appetite" of Fleckvieh bulls was related to their
temperament. 84
 BULL LIBIDO/SERVING CAPACITY 339

PITFALLS IN TESTING BULLS FOR SEX DRIVE

Successful testing of bulls for libido and mating ability requires

careful planning. The following represent some of the pitfalls that should
be avoided in testing procedures that employ restrained, nonestrous
females.
1. Testing of bulls that are excessively apprehensive or agitated.
Precautions should be taken to handle cattle quietly and to avoid
unnecessary distractions during the test.
2. Testing of bulls immediately after their subjection to other proce-
dures such as electroejaculation, vaccination, or parasite control
measures.
3. Testing under adverse environmental conditions such as in ex-
treme heat, cold, or inclement weather.
4. Testing of bulls in groups in which one or more bulls are mark-
edly dominant, such as with mixed-age groups of bulls. The
exposure of only two bulls to the test at a time, and subsequent
retesting with a different bull, helps to minimize this problem.
However, a dominant bull can exert an inhibitory effect from a
distance (e.g., from an adjacent pen). Stimulus females should be
located at an appropriate distance from each other (5 m or more).
5. Use of inadequate stimuli. Restrained females should be incapa-
ble of excessive movement, or some bulls may be deterred.
Overly restless or agitated females should be replaced. The ser-
vice crates used should not impede mounting and service.
6. Spreading of venereal diseases. Every precaution should be taken
to ensure that diseases such as vibriosis and trichomoniasis are
not transmitted by such procedures.

ANIMAL WELFARE CONSIDERATIONS

The use of restrained nonestrous females as stimuli for bull sex-

drive testing, a feature of a number of published procedures, has led to
understandable concerns for the welfare of the females so employed.
These concerns have led to the development of guidelines for the proper
conduct of such tests, such as those adopted by the Australian Veterinary
Association,3 which are stated below:
1. The service crates used should be properly designed for adequate
restraint of previously unhandled females.
2. Bull groups should be managed so that a number of bulls do not
simultaneously compete for a single female.
3. Significant size disparities between males and females should be
avoided, as should be the conduct of tests on uneven terrain or
unsound footing.
4. Females of suitable temperament should be selected when possi-
ble. Mild sedation should be a routine option.
340 CHENOWETH

5. Females showing signs of agitation or distress should be replaced

expeditiously.
6. Stimulus females should undergo regular application of lubricant
to their vulvas and posterior vaginas.
7. Females that have been serviced a number of times should be
monitored subsequently for untoward sequelae and treated ac-
cordingly.
In addition, consideration should also be shown for the welfare of
the bulls being so tested. Some appropriate precautions are as follows:
1. Do not congregate bulls in yards or pens any longer than is
necessary because such congregation increases the chances of
bull injury. This is particularly true for older bulls but also
applies to mixed-age bull groups.
2. Separate any bull that is showing aggressive behavior that threat-
ens other animals.
3. Avoid using service crates that have projecting edges or other
design features that may cause injury.

ALTERNATIVE ASSESSMENT PROCEDURES

The indirect determination of bull libido (e.g., via blood hormone

levels) is an attractive proposition because it could reduce or eliminate
the time, labor, and esthetic or welfare concerns inherent with current
methods of assessing sex drive in bulls. It would also allow assessment
of bulls that, for reasons not readily apparent, do not give a good
response in such tests. However, attempts to link either sporadic or
sequential luteinizing hormone (LH) or testosterone (T) levels with bull
sex drive have generally been disappointing/6, 37, 45, 78 probably because
of the episodic nature of hormone release and the inhibiting effects of
handling or restraint of the anima1. 78 By inducing LH or T release with
parenteral administration of gonadotropin-releasing hormone (GnRH),
some of these difficulties have been avoided. In one study,73 a significant
relationship occurred between induced T levels and bull fertility;
whereas in another,68 positive relationships were obtained with induced
LH levels. However, other studies have obtained disappointing results
when attempting to relate GnRH-induced levels of testosterone or LH
with bull sex drive. 14,16

SUMMARY

Bull libido, or sex drive, is a measurable trait with a large genetic

component. It also represents an important aspect of bull reproductive
performance, with positive effects on herd pregnancy rates and their
patterns. Comparative, quantitative assessment of this trait requires a
formalized testing procedure such as a test for libido or serving capacity
 BULL LIBIDO/SERVING CAPACITY 341

score. Such tests are useful not only for obtaining quantitative informa-
tion, but also for the detection of physical and pathologic problems that
may interfere with normal bull mating ability. These tests should be
conducted in such a manner that animal welfare is not unnecessarily
compromised. The quantitative results should be interpreted with cau-
tion, especially when young, inexperienced bulls or those of Bos indicus
breeds are being assessed. Libido does not necessarily work in concert
with other traits known to separately influence bull fertility (e.g., BSE
traits and social dominance). Until a single procedure is found that can
adequately assess all of these factors, optimal bull appraisal requires
separate evaluation of each of these factors.

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