Animal Reproduction Science 87 (2005) 73–92

Infertility in the dromedary bull: a review

of causes, relations and implications
A.A. Al-Qarawi
Department of Veterinary Medicine, College of Agriculture and Veterinary Medicine,
University at Al-Qassim, P.O. Box 1482, Buraydah, Saudi Arabia

Received 2 June 2004; received in revised form 21 October 2004; accepted 9 November 2004

Abstract

Research into infertility in the dromedary bull, as reported during the last two decades, is reviewed
with emphasis on causes and effects. Reproductive activity of such animals is naturally limited by
a breeding season, though with enough encouragement some may mate with oestrous females out
of season but a full fertilization potential can in no way be expected. It is essential that any female
presented to a bull is capable of reproducing. The presentation of a subfertile or infertile female due
to infection or physiological abnormality will adversely affect the female’s ability to conceive and,
therefore, the apparent fertility rates of the bull she was put to. The average number of successful
services a bull could be expected to perform is two per day. Dromedary bulls with large testes have
larger sperm outputs and can cope with more than two females per day providing that they are given
adequate periods of rest, 1–2 days every 10 days or so, in conjunction with appropriate nutrition
throughout the season. Anabolic steroids or testosterone therapies, which are sometimes used in an
attempt to improve male characteristics and bull libido, are not recommended for dromedary bulls
in breeding work. Such steroids result in a decrease in testicular size and weight with fewer sperm
per gram of testicular tissue being found and the sperm produced also have lower motility rates. Pain
associated with the act of mating a she-camel, due to injuries or inflammation in the scrotum, testes,
prepuce and sheath, can cause a permanent reduction in bull libido. Camel bulls achieving pregnancy
rates more than 60% have had consistently higher spermatozoal concentrations and kinematic variables
derived by the computerized cell motion analyzer (CMA) system. As far as physical capabilities are
concerned, 3-year-old dromedary bulls, which have reached puberty, have been shown to be perfectly
capable of fertilizing a female, but they have a limited sperm production to perform consistently

E-mail address: alialqaraqi@hotmail.com.

0378-4320/$ – see front matter © 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.anireprosci.2004.11.003
74 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92

throughout the season in a large herd. By 4.5–5.0 years of age, they are capable of producing adequate
numbers of sperm to mate as many as females as an adult bull but fertilizing capacity is not attained
until 6 years of age on average. Hyperoestrogenaemia, associated with autoimmune thyroiditis and
trypanosomiasis, suppresses the secretion of testicular testosterone and augments the release of testic-
ular histamine, which appears mandatory for quantitative reduction/loss of advanced spermatogenic
cells in infertile dromedary bulls.
© 2004 Elsevier B.V. All rights reserved.

Keywords: Infertility; Causes; Implications; Dromedary bulls

1. Introduction

The reproductive efficiency of camels under natural conditions is generally considered

to be low. For example, in Tunisia, Djellouli and Saint-Martin (1992) reported an annual
calving rate of approximately 40% for 679 exposed females in 30 herds and a mortality rate
between birth and 1 year of age of 17%. Saley (1990) reported calving rates as low as 45%
in East Central Niger and mortality rates of 10–25% at 0–6 months of age. The reasons
for these low reproductive rates could be blamed on the female dromedary due to delay in
the onset of puberty, a gestation period of 13 months, a prolonged period (8–10 months)
of lactation-related anoestrus leading to a long interval between births, high incidence of
genital tract abnormalities resulting in failure of fertilization and/or implantation in the order
of 15–20% and finally increased rates of early embryonic deaths especially during the hotter
months of the year (El-Wishy, 1987; Arthur, 1992; Musa et al., 1993; Tibary and Anouassi,
1997; Sghiri and Driacourt, 1999; Al-Eknah, 2000; Skidmore, 2003). However, half of the
responsibility for the success or failure of a covering lies with the bull and half with the
female. If a she-camel, therefore, is covered by an infertile or subfertile bull due to infection
or physiological abnormality her chances of producing a calf are markedly reduced though
no fault of her own. Research into causes of infertility in dromedary bulls is yet rather
limited and is largely limited to the reproductive physiology and artificial insemination in
such animals (El-Wishy, 1988; Ismail, 1988; Musa et al., 1993; Tibary and Anouassi, 1997;
Salhab, 2001; Skidmore, 2003). In the present paper, research with respect to all possible
extrinsic (external) and intrinsic (internal) factors affecting reproductive performance in
dromedary bulls during the last two decades will be reviewed to present a survey of the
actual state of knowledge and to indicate possible voids, specially with reference to mating
efficiency and development of reliable semen evaluation method, as a point of departure for
additional research in this area.

2. Extrinsic causes

Extrinsic causes affecting the reproductive efficiency of a dromedary bull include breed-
ing season, lack of use, the presentation of subfertile or infertile she-camels, poor bull
management and methods of semen collection and evaluation.
 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92 75

2.1. Breeding season

Reproductive activity in the camel bull is naturally limited by a rather short breeding sea-
son (Table 1). The variation in both the onset and duration of the breeding season amongst
different countries demonstrates that local environmental factors must initiate sexual activ-
ity. One view is that changes in day length stimulate seasonality (Williamson and Payne,
1978; Chen and Yuen, 1979) but, obviously, in dromedaries situated near the equator, fac-
tors such as rainfall (Bono et al., 1985), pasture condition and nutrition (Merkt et al., 1990;
Skidmore, 2003) are more important. Observations in Emirates (Tibary and Anouassi, 1997)
and Tunisia (Kamoun and Wilson, 1994) indicated that well-fed and well-watered female
camels show ovarian activity throughout the year and the determinant factors of the ob-
served seasonality in conception rates are mainly due to a considerable decrease in libido of
the male and increase in early embryonic death during the summer months. The dramatic
increase of testosterone in the blood (Gombe and Oduor-Okele, 1977; Yagil and Etzion,
1980; Agarwal et al., 1991; Azouz et al., 1992) and the testis (Bedrak et al., 1983; Frielander
et al., 1984; Al-Qarawi et al., 2001a) during the rutting season above the basal levels, which
prevail during the rest of the year, could be responsible for the failure of the male to mate,
decreased number of sperm/ejaculate, decreased total sperm number, increased number of
mounts per successful ejaculation and finally increased reaction times in bulls out of season
(Abdel-Raouf et al., 1975; Hemeida et al., 1985a; Tingari et al., 1984; Abou-Ahmed et
al., 1988; Muslah et al., 1992; Abdel-Rahim and El-Nazier, 1996). As a result, fertilization
potential out of season is significantly reduced.
A stud male is, however, said to be capable of mating with and fertilizing an oes-
terous female at any time of the year (Arthur et al., 1985; Arthur, 1992). A female
oestrogen-dependent pheromone was implicated in the rapid and simultaneous response
of the dromedary male with increased libido and testosterone levels in the blood (Elias
et al., 1985). Moreover, breeding was possible when males were kept in a cooler con-
trolled environment during the extreme heat and humidity of August in Emirates (Tibary
and Anouassi, 1997; Skidmore et al., 2002). Indeed, if the dromedary bulls are overused
during the non-breeding season, the effect is even more detrimental than overuse during

Table 1
Rutting season in the dromedary bulls
Country Period Authors
Egypt December–April Shalash (1965); Abdel-Raouf et al. (1975)
Emirates November–February Tibary and Anouassi (1997); Skidmore (2003)
India November–March Singh and Prakash (1964); Matharu (1966); Sharma and Vyas (1981);
Rai et al. (1988)
Israel January–April Yagil and Etzion (1980)
Morocco November–April Charnot (1965)
Pakistan December–March Yasin and Wahid (1957)
Saudi Arabia November–March Tingari et al. (1984); Arthur et al. (1985); Abdel-Rahim and El-Nazier
(1990)
Somalia April–May Mares (1954)
Tunisia November–April Burgermeister, 1974; Djellouli and Saint-Martin (1992)
76 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92

the breeding season and a full work load can in no way be expected. The ability to mate
and fertilize a female depends on the individual animal but in majority of the cases females
that conceived during the hotter months of the non-breeding season had experienced early
embryonic death and resorption within the first 45 days of gestation (Skidmore, 2003). The
effect of heat and humidity on embryo survival is also well illustrated by embryo transfer
results. In a study by McKinnon and Tinson (1992), no pregnancies were obtained after
transfer of embryos during July and August in Emirates. The natural breeding season, with
its maximum libido as well as optimum fertilization and embryo survival rates, is a natural
mechanism, which ensures that calves are born during the spring and early summer so as
to maximize their chances of survival.

2.2. Lack of use

Reproductive efficiency in any animal is reflected by its performance in the breeding

program. A dromedary bull may not be used in a certain year by design due to financial
or management considerations by the owner. Disease may also preclude a bull from use
due to the risk of a direct transfer to she-camels in the case of venereal (e.g. brucellosis) or
inherited (e.g. cryptorchidism) diseases, or it may limit his ability to perform in the case of
non-contagious diseases (e.g. orchitis). Alternatively, the dromedary bull may have suffered
from disease or infection (e.g. trypanosomiasis) during the previous year and is not to be
used in the following season in order to allow full recovery, or because the long-term effects
of disease on his reproductive performance deem it inappropriate to use him until he is fully
recovered. Diseases of the dromedary bull’s reproductive tract will be discussed later under
intrinsic factors. Finally, semen evaluation, which is part of good practice and should be
carried out regularly at the beginning of each season (Al-Qarawi et al., 2002), may have
indicated very poor quality and the bull may be taken out of use until the cause has been
isolated and the problem solved.

2.3. Subfertile or infertile she-camel

Both male and female camels are equally responsible for the production of an offspring.
A bull is only as good as the female he is expected to cover and vice versa. It is essential
that any female mated by a bull is capable for reproducing and does not suffer from any
reproductive problem. If the female camel herself is subfertile or infertile due to intrinsic
abnormalities, lack of success cannot be blamed on the bull. Based upon the extensive field
studies (Musa, 1984; Shawki et al., 1985; Ali et al., 1987; Chauhan et al., 1987; Wernery and
Ali, 1989; Enany et al., 1990; Wernery, 1991; Chauhan and Kaushik, 1992; Wernery and
Renate Wernery, 1992; Abdel-Rahim, 1996) and abattoir surveys (El-Wishy, 1989, 1990;
El-Khouly et al., 1990, 1991; Kibadu et al., 1991; Ridan et al., 1991; Al-Ani et al., 1992;
Fetaih et al., 1992), the most common genital diseases causing subfertility or infertility in
female camels include infectious (ovaro-bursal adhesions, acute or chronic endometritis
and pyometra), congenital (ovarian agenesis, dermoid cysts, double cervices and persistent
hymen) and finally physiological (ovarian cysts and abnormal vaginal projections unique
to camels) abnormalities.
 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92 77

2.4. Poor bull management

All aspects of bull’s management will affect his ability to cover female camels suc-
cessfully. According to our clinical experience and the data published in this area, bull
management as far as its effects on reproductive efficiency are concerned can be subdivided
into the following sections.

2.4.1. Nutrition
Appropriate nutrition throughout the year is essential in order to ensure that the camel
bull is in optimal physical condition in readiness for the season. Obese or excessively
thin bulls suffer from low libido and nutrition is a major determinant of body condition
(Arthur, 1992). A body condition score of 3 (the hump is prominent on the back of the bull
with the skin well extended over it showing a clear slope to the head of the tail and the
transverse processes cannot be seen or felt with folds of fatty tissue present at the tail head
area) reported by Abdel-Rahim and El-Nazier (1990) is to be aimed for. A standard set of
practices involving free-range grazing for 7–8 h followed by confinement to the fenced farm
areas established near local towns have been adopted in different areas of the Middle East
(Tibary and Anouassi, 1997). The proximate composition of major fed plants grazed by
camels in Saudi Arabia was determined by Abdel-Rahim et al. (1994), who concluded that
their nutritional value is low and they need supplementing with better quality feed especially
with protein. It has been noted in Saudi Arabia (Bakkar and Basmail, 1988; Bhattacharrya
et al., 1988) and Jordan (Al-Ain, 2002) that dromedary camels supplemented with barley
and lucerne have had better condition scores and fertility levels compared to those grazed
only. As far as specific deficiencies are concerned, only limited research has been carried
out in camelids. It is known that general severe nutritional deficiencies are associated with a
delay in puberty, testicular atrophy and a reduction in sperm production (Arthur et al., 1985;
Tibary and Anouassi, 1997). Severe deficiencies in Vitamin A are specifically associated
with a reduction in the testis weight and spermatogenesis (Ismail et al., 1988). Deficiencies
in iodine and selenium have also been associated with marked reduction in the ejaculated
volume and sperm numbers (Ahmed and Nada, 1993; Barsham et al., 2002).
Obesity, associated with excess fat deposition within the scrotum and perineal region,
will result in increase in testes temperature, loss of libido and marked reduction in sper-
matogenesis (Wilson, 1992).

2.4.2. Excess breeding labor

Considering the relatively short breeding season together with the excessive numbers of
females to be mated by one male (1 male per 100–120 females), excess work load could be
a cause of herd infertility under range conditions (Arthur, 1992; Djellouli and Saint-Martin,
1992). Nowadays, most camels in several parts of the Middle East have been transferred to
more intensive management systems where the male is isolated and mating occurs under the
owner’s supervision (Abdel-Rahim and El-Nazier, 1992). However, the amount of work or
number of females a bull may be expected to cover successfully during a season, whatever
the management system, is highly variable depending upon individuals. It is one of the
responsibilities of the bull manager to know the capabilities of his bulls and to work within
these limits. The work load of a bull depends largely on the ability of his testes to produce
78 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92

sperm. This is a function of testes size that could be assessed by calipers or untrasonically
(Al-Qarawi and El-Belely, 2004). Dromedary bulls with large testes have larger daily sperm
outputs (Osman and El-Azab, 1974; El-Wishy and Omar, 1975) and can cope with a heavier
work load than those with small testes. Testes size is also a function of age (Ismail, 1979,
1982), which has an important bearing on fertility, especially at the extremes of youth and
old age.
Sperm concentrations of a camel bull are usually in the range of
(200–600) × 106 sperm/ml and for successful fertilization, assessed by the use of ar-
tificial insemination, (400–600) × 106 sperm/ml are required (Tibary and Anouassi, 1997).
On an average, 50–60% of the sperm collected can be classified as normal progressively
motile sperm capable of fertilizing an ovum (Sieme et al., 1990; William et al., 1993; Hassan
et al., 1995). The average daily sperm production for a dromedary bull is (800–2000) × 106
depending upon season, environment, age, etc. (Ismail, 1982). From these figures it can
be calculated that in theory the average number of successful services a dromedary bull
can be expected to perform per day is between one and three. There are, however, other
considerations to take into account when looking at work loads.
It is interesting to note that total sperm production per week is the same regardless
of whether a dromedary bull is used daily or on alternate days. However, use on a daily
basis results in a lower concentration of sperm/ml (Osman and El-Azab, 1974). This may
be of no consequences in bulls with high daily sperm production ((1400–2000) × 106 ),
as concentrations will still be acceptable, but daily use of bulls with lower daily sperm
production figures ((800–1000) × 106 ) may have a deterimental effect on fertility rates
(Chen et al., 1985).
Overuse will not only deplete the bull’s sperm reserves but will also result in the ejac-
ulation of immature sperm. Dromedary sperm have to spend a period of 48–72 h within
the epididymis of the testis in order to mature in readiness for fertilization (Tingari and
Moniem, 1979; Osman, 1986; Osman and Ploen, 1986a). If this period of time within the
epididymis is reduced because of overuse, then the sperm ejaculated will have a limited
fertilizing potential.
Excess work loads in a dromedary bull may also result in a lack of libido (Arthur, 1992).
As a result of this, the bull will be slow to breed or may even fail to ejaculate. In such cases,
it is best to take the bull out of work for a short period of time, and reintroduce him a week
or so later. If libido is still low it may well be indicative of further problems.

2.4.3. Chemicals and drugs

The male camel loses up to 16–25% of its body weight during the rutting season be-
cause its sexual activity prevents normal feeding together with the imposed work load
(Khanna, 1990; Abdel-Rahim and El-Nazier, 1992). Anabolic steroids, such as nandrolone
and trenbolone, are sometimes used by the camel pastoralists in an attempt to improve
weight gain, muscle growth and performance in stud camels during the breeding season.
Camel breeders also have used these anabolic steroids to improve deficient sex derive in
bulls out of season. In man (Roche, 1980; Brander et al., 1982) and the stallion (Morel,
1999) such use of anabolic hormones is associated with infertility and a similar associa-
tion has been indicated in dromedary bulls. The results of El-Beley and Al-Qarawi (2002)
revealed that the intramuscular injection of nandrolone phenylpropionate (600 ␮g/kg body
 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92 79

weight) daily for 7 days in three stud bulls had resulted in a decrease in testicular size in the
order of 25–30%. Spermatogenesis may also be reduced, with fewer sperm concentration
((150–170) × 106 sperm/ml) being found, and the sperm produced also had lower motility
rate (30–40%). Anabolic steroids, thus, are not recommended for camel bulls in breeding
work. These drugs not only have an immediate effect, but may also have a long-term effect,
at least until they are cleared from the bull’s system (Ali and El-Sheikh, 1992).
Testosterone propionate is widely used in nomadic camel herds to enhance libido at the
beginning of the breeding season especially in stud males aged over 12 years and as such is
relatively effective. However, the owners of these herds advocated that testosterone therapy
does have serious potential side effects as far as fertility is concerned. Testicular testosterone
is largely responsible for sperm production and also feeds back negatively on pituitary
function to reduce the release of LH and FSH, and hence controlling its own production.
This negative feedback ensures that the hypothalamus–pituitary–testicular system does not
overrun itself as reported in dromedary bulls (Bedrak et al., 1983; Al-Qarawi et al., 2001a),
bovine bulls (Sharp, 1984), stallions (Bedrak and Samuel, 1969) and rams (Walton et al.,
1980). If, however, testosterone is added exogenously it affects the delicate balance of
the whole axis. Four dromedary bulls suffering from deficient sex drive were selected
during the rutting season by El-Belely and Al-Qarawi (2003) and injected intramuscularly
with testosterone propionate (50 mg twice weekly for 4 weeks). Testosterone therapy had
provoked the behavioural signs of libido in such animals but a reduction in sperm counts in
the order of 20–25% was evident. Testosterone treatment could reduce the release of FSH,
which is known to start the process of spermatogenesis in dromedary bulls (Al-Qarawi et al.,
2001a), developing spermatogonia to secondary spermatocytes, and hence reduced sperm
concentrations. Testosterone is, therefore, associated with low fertility due to low sperm
counts and hence is not advised for use in dromedary camels in breeding work.
In addition, the use of any other drug or treatment that causes a drop in appetite, diarrhoea
or lack of condition is ill-advised during the breeding season and should only be used under
veterinary supervision. Some anthelmentics (Kayum et al., 1992; Raisinghani, 1992) and
acaricides (Kumar et al., 1992) have also been associated with a temporary decline in
fertility, and thus, many camel breeders arrange their parasite control regimes to ensure that
dromedary bulls are not treated during the breeding season.

2.4.4. Breeding discomfort

Full physical examination of the camel bull is essential before purchase to ensure that
no abnormalities are present. It is also advisable that bulls undergo regular clinical and
anatomical examinations at the beginning of each season to ensure that no problems have
arisen that may cause pain during mating (Tibary and Anouassi, 1997). Pain associated with
the act of covering a she-camel can cause a permanent reduction in bull libido. The male
camel shows particular reproduction behaviour during rutting season and becomes dominant
and aggressive. It frequently balloons the soft palate and tends to bite other animals, nearby
standing humans and fixed objects (Abdel-Rahim and El-Nazier, 1992). This predisposes
him to fractures of the mandible and injuries of the soft palate (Purohit et al., 1984; Ramadan,
1992) reducing his willingness to mate. When it bites other males, it may lead to injuries
in the scrotum, testes, prepuce and sheath (Gahlot, 2001) causing pain to these males at
mating. Finally, when using an artificial vagina (AV) care should be taken that the internal
80 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92

temperature is not too hot. Temperatures above 41–43 ◦ C (Musa et al., 1992; El-Belely and
Al-Qarawi, 2004) will cause the bull pain and will reduce his future willingness, not only
to use an AV, but also in natural service.

2.5. Methods of semen collection and evaluation

Semen acquisition can be considered an important step towards assessing breeding

soundness and fertility examinations in camel bulls under field conditions. From a practical
point of view and based on consistency of response, collection of semen from such animals
is undertaken by the use of AV during mating or via electro-ejaculation (EE). Several studies
in dromedary camels (Tingari et al., 1986; Anouassi et al., 1992; Musa et al., 1993, Hassan
et al., 1995; Abdel-Rahim, 1997; Aminudeen et al., 2001; Skidmore et al., 2001) have used
the light microscope and did not find any significant effect of the collection method (AV or
EE) on the major macro- and micro-characteristics of semen (Table 2). Recently, Al-Qarawi
et al. (2002) have considered spermatozoal concentrations and kinematics as the major cri-
teria of computerized semen analysis for predicting fertility of the camel bull (Table 3). The
elements of subjectivity, and its attendant drawbacks of human error and bias, are intro-
duced when using the traditional microscope by the previously mentioned authors, thereby
limiting its reliability as a fertility probe especially when comparing the fertilizing ca-
pacity of semen collected by both techniques. More recently, El-Belely and Al-Qarawi
(2004) provided new evidence that spermatozoal concentrations and velocity variables
(straight line velocity, VSL; curve line velocity, VCL; and average line velocity, VAP)
as well as other kinematics (amplitude of the lateral head displacement, ALH; linearity,
LTN; and straightness, STR) derived by the computerized cell motion analyzer (CMA) sys-
tem from semen collected by AV are superior to that collected by EE (Table 4). Camel bulls
achieving pregnancy rates more than 60% have had consistently higher spermatozoal counts
and kinematic attributes (Al-Qarawi et al., 2002; Table 3) indicating that fertility rates of
semen collected by AV are likely to be higher than those of semen collected by EE methods.

3. Intrinsic causes

Intrinsic causes affecting reproductive performance in the dromedary bull include age
along with hormonal, testis, epididymis, penis and prepuce abnormalities.

Table 2
Range of average characteristics in ejaculates of the dromedary bullsa
Attribute Artificial vagina Electro-ejaculation
Semen volume (ml) 3.0–9.5 3.8–4.2
Sperm concentration (×106 (ml)) 350–450 320–400
Sperm motility (%) 40–60 40–50
Stained (dead) spermatozoa (%) 16–18 18–20
Total sperm showing morphological abnormalities (%) 18–25 26–30
Sperm showing acrosomal abnormalities (%) 8–10 8–12
a Tingari et al. (1986); Anouassi et al. (1992); Musa et al. (1993); Hassan et al. (1995); Abdel-Rahim (1997);

Skidmore et al. (2001).

 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92 81

Table 3
Correlation between fertility rates and spermatozoal concentrations as well as kinematic variables measured by
the cell motion analyzer (CMA) system in dromedary camels (Al-Qarawi et al., 2002)
Parameter Sperm no. Motility Progressive VSL VCL VAP ALH LIN STR
(106 ml−1 ) (%) motility (%) (␮m/s) (␮m/s) (␮m/s) (%) (%)
Motility 0.581a 1.000
Progressive motility 0.396a 0.735a 1.000
VSL 0.322b 0.291b 0.385a 1.000
VCL 0.164 0.313b 0.391a 0.086 1.000
VAP 0.217 0.308b 0.317a 0.219 0.314a 1.000
ALH 0.153 0.469a 0.724a 0.286b 0.308b 0.319b 1.000
LIN 0.385a 0.415a 0.655a 0.394a −0.421a 0.154 0.425a 1.000
STR 0.282b 0.311b 0.158a 0.401a 0.115 −0.561a 0.214 0.109 1.000
Fertility rate 0.509a 0.381a 0.859a 0.416a 0.253b 0.241b 0.824a 0.910a 0.320b

No. of analyses = 42 (3 ejaculates × 14 animals), VSL, straight line velocity; VCL, curve line velocity; VAP,
average line velocity; ALH, amplitude of the lateral head displacement; LIN, linearity; and STR, straightness.
a Significant at 1% level.
b Significant at 5% level.

3.1. Age

Age is an important aspect in considering the potential fertility of a camel. Young and
old dromedary bulls may have problems with taking on a full breeding labor with consistent
success rates. Based upon assessing testicular morphometry and rutting behaviour, a young
bull may be sexually active and used for service as early as 3 years of age as reported in
Saudi Arabia (Arthur et al., 1985; Abdel-Rahim, 1997; El-Agawany et al., 1998), Egypt
(Abdel-Raouf and Owaida, 1974; El-Wishy and Omar, 1975; Ismail, 1982), India (Matharu,

Table 4
Mean ± S.E. values of spermatozoa concentration and kinematics measured by objective (CMA) and subjective
(visual) methods in camel semen collected by AV or EE (El-Belely and Al-Qarawi, 2004)
Semen characteristics AV method (n = 24 ejaculates) EE method (n = 24 ejaculates)

CMA Visual CMA Visual

Volume of ejaculate (ml) 7.3 ± 1.1 3.8 ± 0.6
Sperm concentration (×106 ) 525 ± 45 376 ± 39 384 ± 33 349 ± 41
Motility (%) 72.3 ± 5.7 50.8 ± 7.4 58.3 ± 4.6 45.2 ± 5.8
Progressive motility (%) 68.4 ± 5.2 43.1 ± 6.3 52.7 ± 4.2 40.4 ± 5.6
VSL (␮m/s) 125.6 ± 11.9 96.8 ± 9.6
VSL (␮m/s) 132.5 ± 12.6 107 ± 10.0
VSL (␮m/s) 129.4 ± 12.2 100.9 ± 9.5
ALH 7.7 ± 1.1 5.4 ± 0.8
LIN (%) 94.0 ± 12.0 86.0 ± 9.0
STR (%) 97.0 ± 14.0 94.0 ± 11.0

CMA, cell motion analyser; AV, artificial vagina; EE, electroejaculation; VSL, straight line velocity; VCL, curve
line velocity; VAP, average line velocity; ALH, amplitude of lateral head displacement; LIN, linearity; STR,
straightness.
82 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92

1966; Nada, 1966; Singh and Bharadwaj, 1974; Djang et al., 1988) and Kenya (Gombe
and Odour-Okele, 1977). However, Skidmore (2003) found that young bulls aged 3–4
years old, which is long before physical maturity is attained, would have little chance of
mating with females in a large mixed herd. Moreover, Arthur (1992) and Al-Qarawi et al.
(2002) reported that pregnancy rates in clinically healthy females, mated by dromedary
bulls aging between 3 and 5 years old were significantly lower than in those mated by bulls
exceeding these ages. Al-Qarawi et al. (2001a) have developed a new scheme for evaluating
spermatogenesis in conjunction with measuring testicular hormones during the pubertal
period in dromedary bulls (Table 5) and concluded that as far as physical capabilities are
concerned, 3-year-old dromedary bulls, which have reached puberty, have been shown to be
perfectly capable of fertilizing a female, but they have a limited sperm production to perform
consistently throughout the season in a large herd. By 4.5–5 years of age, they are capable
of producing adequate numbers of sperm to mate as many females as an adult bull but full
fertilizing capacity is not attained until 6 years of age on average. These conclusions were
also emphasized by Al-Qarawi et al. (2000), who reported for the first time in dromedary
bulls that the major elements of testes (Na, Ca and Cu) epididymides (P and Fe), prostate
(Zn) and bulbourethral glands (K and Mg), which affected sperm motility and metabolic
activity, are secreted and peaked at around 6 years of age, when sexual maturity is reached.
At the other end of the spectrum old age may be a problem. Age related decline in fertility
is associated normally with general age related problems rather than reproductive capacity
per se. Arthritis causing pain on mounting is a major cause of lack of libido and, there-
fore, low fertilization rates in older dromedary bulls (Yagil, 1985). Al-Qarawi et al. (2000)
found that testicular degeneration associated with a marked decline in plasma testosterone
concentrations, concomitant with a clear disturbance in the mechanisms controlling normal
cation distribution in the testes at over 18 years of age could provide an explanation for
the reduced gonadal sperm reserve, the increased percentage of spermatozoal abnormal-
ities and the poor axonemal motility characteristic of semen collected from aged camels
(Singh and Bharadwaj, 1974; Osman and Ploen, 1986b; Tingari et al., 1986; Willmen et
al., 1993; Skidmore, 2003). However, the effect of age is very different in individual camel
bulls (Tibary and Anouassi, 1997) and older bulls should not be precluded from use, as such
animals have had many years in which to prove their worth as far as their own performance
and that of their progeny are concerned. If using an older camel bull his semen should be
evaluated regularly and monitored closely to allow reduction in his work load if a drop in
any of the semen evaluation parameters is detected.

3.2. Hormonal abnormalities

The influences and interrelationships of hormones secreted by the hypothalamus–

pituitary–testicular axis on the process of spermatogenesis, and hence potential fertility,
were clarified in dromedary bulls only by Al-Qarawi and El-Belely (2004). Sertoli cells
are the main source of seminal and plasma oestradiol 17␤ in dromedary bulls (Al-Qarawi
et al., 2001a). Autoimmune thyroiditis with increased thyroid autoantibodies (Al-Qarawi
et al., 2001b) and blood born parasites activating the immune system to produce cytokines
from macrophages (Al-Qarawi et al., 2004) constitute the two main conditions causing
hyperactivation of Sertoli cells and consequently hyperoestrogenaemia. Such defective ste-
Table 5
Morphometric and cellular changes (mean ± S.E.) in the testes of the Camelus dromedarius around the time of puberty (Al-Qarawi et al., 2001a)
Parameter Age group (years)

<1 (n = 5) 1.5 (n = 6) 2.5 (n = 10) 3 (n = 8) 4.5 (n = 8)

Testicular weight (g) 4.8 ± 0.5 6.9 ± 0.8 8.3 ± 1.0 11.9 ± 1.2 53.8 ± 5.6**
Diameter of the seminiferous tubules (␮) 98.5 ± 8.4 108.8 ± 10.1 119.6 ± 12.0 136.2 ± 13.3 183.4 ± 18.4**
Area occupied by the interstitial tissue in relation to the semi- 17.5 ± 1.6 12.4 ± 2.0 23.5 ± 2.2 31.8 ± 3.1 32.1 ± 3.3

A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92

niferous tubule (%)
Size of leydig cells (length ␮m × width ␮m) 12.3 ± 1.2 × 6.0 ± 0.5 13.5 ± 1.4 × 7.2 ± 0.7 14,6 ± 1.5 × 7.5 ± o.8 22.4 ± 2.2* × 9.8 ± 0.9* 23.5 ± 2.3 × 10.2 ± 1.1
Numbers of Sertoli cells in 100 seminiferous tubules Nil Nil 38.5 ± 1.8 58.9 ± 3.2 84.8 ± 4.9
Size of Sertoli cells (length ␮m × width ␮m) Nil Nil 25.6 ± 2.4 × 6.6 ± 0.6 26.8 ± 2.4 × 6.8 ± 0.7 36.5 ± 3.8* × 10.2 ± 1.1*
Percentage of the seminiferous tubules containing
Spermatogonia 84.5 ± 8.6 82.4 ± 8.1 73.4 ± 7.5 60.4 ± 6.6 34.2 ± 4.2
Primary spermatocytes 11.3 ± 1.2 12.9 ± 1.3 15.5 ± 1.3 21.2 ± 2.3 28.1 ± 2.9
Secondary spermatocytes 4.2 ± 0.3 34.7 ± 0.5 7.9 ± 0.7 10.5 ± 1.3 21.9 ± 2.2
Spermatids Nil Nil 3.2 ± 0.2 4.8 ± 0.5 8.6 ± 0.6
Spermatozoa Nil Nil Nil 3.1 ± 0.3 7.2 ± 0-.5

Parameter Age group (years)

5 (n = 7) 5.5 (n = 5) 6 (n = 6) 6.5 (n = 6) ≥7 (n = 5)
Testicular weight (g) 61.5 ± 6.3 70.8 ± 7.2 81.2 ± 8.3 92.3 ± 9.5 93.4 ± 9.3
Diameter of the seminiferous tubules (␮) 245.8 ± 25.0 272.3 ± 28.1 296.5 ± 30.1 295.1 ± 29.2 301.3 ± 30.2
Area occupied by the interstitial tissue in relation to the semi- 33.9 ± 3.2 23..4 ± 3.3 31.8 ± 3.2 31.9 ± 3.2 32.5 ± 3.4
niferous tubule (%)
Size of leydig cells (length ␮m × width ␮m) 22.9 ± 2.9 × 10.1 ± 1.0 24.7 ± 2.8 × 10.3 ± 1.3 25.1 ± 2.5 × 10.2 ± 1.1 24.8 ± 2.4 × 10.3 ± 1.2 23.6 ± 2.2 × 10.4 ± 1.3
Numbers of Sertoli cells in 100 seminiferous tubules 185.2 ± 8.9** 359.5 ± 26.3 598.3 ± 58.2 595.4 ± 58.5 596.6 ± 60.3
Size of Sertoli cells (length ␮m × width ␮m) 38.7 ± 3.9 × 10.6 ± 1.7 38.6 ± 3.8 × 10.5 ± 1.6 37.9 ± 3.6 × 10.3 ± 1.2 38.2 ± 3.7 × 10.0 ± 0-.9 38.5 ± 3.9 × 10.2 ± 1.3

Percentage of the seminiferous tubules containing

Spermatogonia 25.1 ± 3.0 14.7 ± 1.3** 6.7 ± 0.9 6.3 ± 0.6 6.1 ± 0.6
Primary spermatocytes 31.4 ± 3.1 10.2 ± 1.1** 10.8 ± 1.3 10.2 ± 1.1 10.4 ± 1.3
Secondary spermatocytes 21.0 ± 1.8 7.1 ± 0.6** 5.2 ± 0.8 7.0 ± 0.6 6.8 ± 0.7
Spermatids 11.2 ± 1.2 39.6 ± 4.1 8.5 ± 1.3** 8.6 ± 0.9 8.3 ± 0.8
Spermatozoa 11.3 ± 1.3 28.4 ± 2.9 68.8 ± 5.9** 67.9 ± 6.8 68.4 ± 6.9
Differences amongst various age groups were significant at 5% (*) and 1% (**) levels.

83
84 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92

riodogenesis suppresses secretion of GnRH and exerts a negative feedback mechanism on

LH (Schanbacher, 1979; Amann, 1983; Payne, 1990) leading to depression in Leydig cell
function and maintaining low levels of testicular and plasma testosterone in males of several
species including dromedary bulls (Van der Mollen et al., 1981; West and Brenner, 1990).
Also, hyperoestrogenaemia could augment the proliferation of testicular mast cells and
increased histamine secretion, which appears mandatory for quantitative reduction/loss of
advanced spermatogenic cells in infertile dromedary bulls (Al-Qarawi and El-Belely, 2004).
Treatment of a dromedary bull (Muslah et al., 1992) or a stallion (Blue et al., 1991; Boyle
et al., 1991), experiencing low circulating testosterone concentrations, with GnRH has been
tried successfully in attempts to increase testosterone concentrations and sperm production.
Administration of mast cell blockers, tranilast or ebastin, in azoospermic or oligozoosper-
mic human patients could significantly increase sperm density and sperm motility (Matsuki
et al., 2000; Hibi et al., 2001) via inhibition of testosterone metabolism. Such antihistaminic
therapy is essential for restoring complete spermatogenesis in the azoospermic or oligo-
zoospermic dromedary bulls having low peripheral testosterone concentrations (Al-Qarawi
and El-Belely, 2004).

3.3. Testis abnormality

Testis abnormalities, as with most anatomical abnormalities, are either inherited or caused
by a disease, the significance of which varies from life-threatening to a minor flaw that may
be of little consequence as far as reproductive performance is concerned but may still reduce
the bull market value.

3.3.1. Cryptorchidism
Descent of the testes from the body cavity into the scrotum occurs by mid-gestation in the
bovine bull and ram (Senger, 1999) and before or after birth in the stallion (Morel, 1999). In
the camel, the testes enter the scrotum when the young dromedary is in the second or third
year of age (Smuts et al., 1987). Cryptorchidism is the failure of the testes to enter the scrotum
and this may be unilateral or bilateral. Both types were detected in 1.8% of 155 slaughtered
camels (Hemeida et al., 1985b). In farm animals, except the camel, the cryptorchid testis
remains in the body cavity or situated just above the inguinal ring (Vlissingen et al., 1988),
but in camels it is descended through the inguinal ring and situated subcutaneously in the
region between the sigmoid flexure of the penis and the external inguinal ring (Ismail,
1982). Bilateral cryptorchidism results in sterility of the bull due to complete degeneration
of the seminipherous tubules. Although the use of a unilateral cryptorchid as a stud animal
is possible, it is not advisable, as the condition is heritable. It is normally advised that
dromedary bulls without full descent of the testes by 3 years of age should be castrated in
an attempt to eliminate the trait from the population.

3.3.2. Hypoplasia
Testicular hypoplasia is characterized by inadequate development of primitive sperm
cells or spermatogonia within the testes of the fetus. The extent of hypoplasia, partial
or complete, does vary between individual sufferers and may be due to spermatogonia
degeneration in later life (Ladds, 1993). In the dromedary bull, hypoplasia is evident as
 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92 85

seemingly shrunk testes within the scrotal sack and this is due, in part, to fewer sperm/gm
of testicular tissue and to small epididymides as there are few if any sperm maturing within
them (Abdel-Raouf, 1964). Unilateral and bilateral hypoplasia were decribed in 1.5% of
155 specimens in an abattoir survey (Hemeida et al., 1985c).

3.3.3. Testicular disease

The most common systemic diseases that may cause orchitis (inflammation of the testis)
and degeneration in most camel raising areas include trypanosomiasis, caused by Try-
panosoma evansi (Omer et al., 1998; Al-Qarawi et al., 2004), brucellosis, caused by Bru-
cella abortus or B. melitensis (Ahmed and Nada, 1993; Agab, 1977) and filariasis, caused by
Diptalonema evansi (Youssef, 1976; Shawki et al., 1983; El-Wishy, 1988). The mechanisms
involved in disruption of fertility in the dromedary bull are clarified only in trypanosoma in-
fection via monitoring plasma hormonal concentrations and semen characteristics (Table 6)
by Al-Qarawi et al. (2003). The direct antitesticular effect may be associated with bouts of
hyperthermia for considerable period of time causing degeneration of seminiferous tubules
and associated Sertoli cells (Group 2C). The indirect effect may be mediated through altered
Sertoli cell function due to formation of immune complexes (Group 2A) or could be a result
of pituitary damage along with marked decline in testicular concentrations of both estradiol
17␤ and testosterone (Group 2B). Clinical signs of orchitis due to acute trypanosomiasis
and brucellosis are swollen and firm testes that feel warm to the touch and are extremely
sensitive as shown by the stiff and wide-based gait of the affected bull (Tibary and Anouassi,
1997). Degeneration as a result of orchitis is characterized by soft and flabby testes with
50–80% reduction in sperm counts and motility percentages together with 20–30% increases
in sperm abnormalities (Table 6). If inflammation, and hence degeneration, persist, then the
testes may be felt as small hard structures within the scrotum. Al-Qarawi et al. (2004)
have recommended that dromedary bulls should not be used immediately after recovery
from trypanosomiasis as 75–90 days should be allowed after recovery for the resumption
of spermatogenesis and maturation of adequate sperm numbers within the epididymides.

Table 6
Different hormonal and seminal patterns (mean ± S.E.) in dromedary bulls infected with Trypanosoma evansi
(Al-Qarawi et al., 2004)
Parameter Chronic infectiona Acute infection
Group 2A (n = 4) Group 2B (n = 6) Group 2C (n = 3) Group 2D (n = 5)
Oestradiol-17␤ (pg/ml) 300.6 ± 38.6a 25.3 ± 3.2c 20.5 ± 2.4b 95.3 ± 10.2c
Testosterone (ng/ml) 0.8 ± 0.2c 0.6 ± 0.1c 3.9 ± 0.4a 4.2 ± 0.5a
Sperm count (×106 ml−1 ) 58.1 ± 11.0b 36.0 ± 8.0b 4.0 ± 1.0a 115.0 ± 16.0c
Motility (%) 65.3 ± 7.2a 30.8 ± 4.3b 10.8 ± 2.5a 71.3 ± 8.2c
Live sperm (%) 68.5 ± 9.5a 44.1 ± 11.8b 16.2 ± 5.8a 76.5 ± 9.7c
Abnormalities (%) 7.8 ± 1.4b 8.9 ± 1.8b 35.8 ± 4.6c 2.9 ± 0.6a

(a) vs. (b) and (b) vs. (c), significant at 5% level; (a) vs. (c) significant at 1% level.
a Bulls were grouped according to the different hormonal patterns into: Group 2A, animals displaying about

4-fold increases in oestradiol-17␤ concentrations and basal levels of testosterone; Group 2B, animals maintain-
ing basal levels of both hormones; Group 2C, animals showing basal levels of oestradiol-17␤ and testosterone
concentration close to those of the controls.
86 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92

Traumatic orchitis is frequently encountered in large mixed herds during the rutting
season due to bites suffered while fighting (Nigam, 1992; Gahlot, 2001). In such cases,
the scrotum is punctured, the deep wound usually involves the tuncia vaganalis and, if left
untreated, abscess, scirrhous cord, gangrene and adhesions supervene (Chouhan et al., 1981;
Tibary and Anouassi, 1997).

3.3.4. Testicular tumours

Though rare, tumours of the testes and associated structures are not known in camel
bulls. Those reported by Hemeida et al. (1985b) included sporadic cases of Sertoli cell or
interstitial cell tumours and seminomas (malignant tumours of spermatogonia). However,
dermoid cysts and teratomas were specially evident in cryptorchids (Tibary and Anouassi,
1997).

3.4. Epididymis abnormalities

Blockage of both epididymides due to spermatic granulomas or spermatocoele in the

caput epididymides has been reported in three dromedary bulls (Al-Qarawi and El-Belely,
2004) characterized by a normal libido but azoospermic semen. Swelling and inflammation
of the cauda epididymides are specially evident in dromedary bulls infected with filariasis
(Shawki et al., 1983). The complete absence of one or both epididymides is rare in the
dromedary bulls (Tibary and Anouassi, 1997).

3.5. Penis and prepuce abnormalities

Abnormalities of the penis or prepuce are frequently faced by veterinarians and are
normally associated with trauma or injury to this area. Phimosis occurs due to ischaemic
necrosis of the penis anterior to the segmoid flexure following application of a tight strap
taken around the posteroventral body and sheath when putting camels into carts (Gahlot,
1992; Nigam, 1992).
Paraphimosis may be associated with infection after castration, which stimulate ex-
cessive swelling. In rare cases, after copulation or sand masturbation, the dangling penis
is injured leading to paraphimosis (Choudhary et al., 1981; Purohit et al., 1984; Gahlot,
1996). However, sometimes injury to the penis occurs when it protrudes through a ventral
incision made to drain sheath abscesses. A protruded penis may become infected, which
may lead to necrosis and gangrene if untreated (Nigam, 1992).
Sand masturbation in camels is a vice which causes ascending urinary infection, leading
ultimately to formation of a mucous plug into the urethra, causing urinary obstruction and
subsequently urethral rupture (Gahlot, 1992). Urinary calculi may be formed due to other
reasons not established so far (Kock, 1985) but causing localized to extensive necrosis
of urethral mucosa and urethral rupture (Purohit et al., 1984). Rupture of the urethra and
subcutaneous infiltration of urine may be also caused by stricture of the girth straps (Nigam,
1992).
Not only may the penis itself be infected, but also it is the major means by which
venereal infections can be passed from bulls to she-camels and vice versa. The bull often
does not show any clinical signs of contamination, but infection is traced back through
 A.A. Al-Qarawi / Animal Reproduction Science 87 (2005) 73–92 87

symptoms shown by she-camels he has mated. A number of venereal diseases are transferred
via the bull’s semen in particular brucellosis and trypanosomiasis (Bauman and Zessin,
1992; Ahmed and Nada, 1993; Agab et al., 1996). They are now notifiable diseases in the
Kingdom of Saudi Arabia and other Gulf countries, and valuable bulls require negative
brucellosis and trypanosomiasis certificates before being imported into the country. Truly
sexually transmissible bacteria, that is, those transferred into the uterus of the she-camel at
covering, include Streptococcus pyogenes,E. coli,Pseudomonas aeroginosa and Klebsiella
aeruginosa (Refai, 1990). The presence of these infections in the female camels causes
endometritis (Youssef, 1992).

4. Conclusions

The causes of infertility, or the failure to produce an offspring, whether on temporary or

permanent basis, are numerous. Some are treatable but many may cause the camel bull to
be useless for breeding. It is essential, therefore, that all potential breeding bulls are given
a thorough examination prior to purchase in order to ensure that it is capable of fulfilling
their reproductive potential.
The results show that with good management and use of reliable semen collection and
evaluation techniques it is possible to increase reproductive efficiency of camel bulls. How-
ever, there is a large gap in the area of nutrition and disease control and their relationship
to production that needs to be filled to maximize the reproductive performance of these
animals.

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