Vol. 70 December.2000 No.

12

Veterinury Bulletin

Research on dromed ary reproducti

The past two decades and future prospective

G.N.Purohit and P. K. Pareek

Department of Veterinary Obstetrics and Gynaecology, College of Veterinary & Animal Science,
Raj. Agric. University Bikaner, Rajasthan,INDIA -334001

Abstract
A bibliometric analysis of documents published on the subject of dromedary reproduction (n:489) in
the past two decades and three years (1977-1999) revealed that highest emphasis had been affached to
female reproductive physiology (26.17%), followed by female examination (13.49%), female
reproductive pathology (13.08Yo) andmale reproductive physiology (12.06%). Other research remained
focussed on male reproductive anatomy (8.79o/o), artificial breeding including embryo transfer (8.79%),
female surgery and obstetrics (5.52%), male breeding soundness examination (4.49%), male
reproductive pathology @.29%) and female reproductive anatomy (3.27%).

The main thrust for research on reproductive physiology was mostly between 1988 to 1996. Embryo
transfer research was published more between 1990-1999. Male reproductive physiology was studied
more between 1977-1986 whereas female reproductive physiology was reported most between 1989-
1999. The male reproductive anatomy received greater attention than female reproductive anatomy, but
conffarily female reproductive physiology got more attention than male reproductive physiology. Also,
female reproductive pathology was reported more than male reproductive pathology.

66.46% of the total literature on dromedary reproduction appeared in 59 different journals whereas
33.53% of the research data appeared in publications other than journals. Only 15.54% of the research
data on dromedary reproduction appeared in journals of reproduction whereas 50.92 Yo appeared in other
journals. The currently available bibliographies are mentioned.
,,

The top 6 counffies contributing research on this topic are India, Egypt, Sudan, UAE, Saudi Arahia and
Israel. The important highlights of research in each sector of dromedary reproduction and the important
milestones are discussed. The direction of reproductive research in future is considered.

Introduction
The Arabian camel (Camelus dromedarius) is used for sport besides
its use for transport, m,ilk, meat, hair and hides. Kesseba (1991) esti-
mated the world population of camels as 18.5 million, with
dromedaries accounting for 95Yo (Bactrian camels the other 5olo),
whereas Tibary & Anouassi (1997) estimate the population of drom-
edaries to be 20 million. Low reproductive performance has
remained a major obstacle to the growth of populations of drome-
daries. The history of camel research and literature has been
discussed by Wilson (1991), who drew affention to the acceleration
in the number of publications and changes in emphasis of research in
the Arabian camel. Reproduction in the dromedary has been poorly
studied, as are other aspects of camel. There have been a very few
reviews (Higgins, 1983; El-Wishy, 1987; 1988; Khanna et al., 1987;
Wilson and Bourzat, 1988; Wilson, l99l; Arthur, 19921' Wernery and
Wernery, 1992; Arthur and Tigani,1993; Saint Martin, 1993; Chaud-
hary, 1995; Faye et al.,1995; Agag,1996; Purohit,1999a) on camel
reproduction. During the past two decades plenty of work on the
understanding of reproduction in the dromedary has been done.
There have been 6 camel conferences or seminars (in Egypt, Libya,
Dubai, Israel, India and Morocco) and more than 4 workshops (in
Ethiopia Somalia, Sudan and Paris). The International Camel con-
ference 2-6 February,1992 at Dubai, UAE and the workshop " Is it
possible to improve the reproductive performance of camel? -Paris
1990, have been the important research exchange gatherings in cov-
ering reproduction in this species. The present review analyses the
research on dromedary reproduction during the past twenty-three
years (1977-1999).
1266 Veterinary Bulletin 2000 Vol. 70 No. I2

Materials and methods female than on the male (except during 1995). This also appears to
have emanated from the increased interest on the rearing, breeding
The basic data utilized in the present study was generated by scan- and application of biotechniques for reproductive efficiency
International) and, Animal improvement in females.
ing of the important research
ies. The recent publication of
The greatest number of research publications appeared in 1990 and
by Tibary & Anouassi (1997) in 1992.
thereafter
was an important support for the bibliometic and research scanning.
The data has been scanned into 4 groups according to: l) Area of
reproduction studied which includes male reproductive anatomy,
Journals of Publication
male reproductive physiology, male breeding soundness examina-
The research data for the period was scattered, and, on identification
tion, male reproductive pathology, female reproductive anatomy,
ion, it was found that it appeared in 59 different journals, both national
and and international. Only a small portion (15.54yo) ofthe data on repro-
for duction appeared in national and intemational journals of
reproduction whereas 50. er joumals. Of these
other journals Indian an contributed equally
18.47o/o (46 of 249) and respectively. Nearly
20.24%(99 arY reProduction
appeared in Conferences' Of
these, symp ed a major share

Result and discussion

Area of ReProduction

nant langUage, used in 91.5o/o of the research publications, was

English.
cial breeding including and embryo
transfer(43\4S9). Among reproductive
physiology received high ed by female The top ten publishing joumals included Journal of Camel Practice
Lxamination(13.49%) an gy (13'087o)' and Research (29) Journal of Reproduction & Fertility (24), Indian
Joumal of Animal Science (22), Animal Reproduction Science (21),
Other research remained focussed on male reproductive anatomy
(8.79o/o), female surgery and obstetrics (5.52%o), male breeding Assuit Journal of Medical Science (21), Theriogenology (16), Acta
soundness examination (4.49oA, male reproductive pathology Anatomica (14), British Veterinary Journal (13), Veterinary Record
(4.29o/o) and female reproductive anatomy (3.27yo)' (11) and Veterinary Medical Review (9) in their descending order in
terms of number of publications (mentioned in parentheses). These
thrust in research journals excep! Acta Anatomica and Assuit Journal of Medical sci-
1996. Malerepro- ence (which published more on male anatomy), published papers
77-1986, whereas mostly on female reproductive physiology and patholgy followed by
more frequently male reproductive physiology and collectively contributed
36.80o/o(180\439) of the published literature during the period'
between 1989-1999. Male and female reproductive anatomy was
little snrdied except during 1985, possibly because only the basic
studies were required, whereas female reproductive pathology was It is clear that data on reproductive research in the dromedary is being
continuously studied, the impetus being received since 1989. It is
known that only a small number of males are required to mate with
large number of females, and when infertile, males can be replaced,
whereas with females correction of the pathological condition is
essential, and has led to more studies on the female. The male repro-
ductive anatomy received greater attention than female reproductive
anatomy, but contrarily reproductive physiology and reproductive competition between on) forces many authors
pathology was studied Reproductive from developing cou onal publication of their
physiology of female ecialities like results. Further, these m looking at publications
induced ovulation, ill luteal phase, on cattle that much of the information is published in the form of pro-
seasonality of breeding, longer gestation period and poor reproduc- ceedings, thesis, annual reports which are difficult to locate and
tive efficiency. These and other factors like long term preservation of acquire. The same appears to be true for research on dromedary'
semen (this makes the breeding available for use Moreover, due consideration should be given to the research which
at any part ofthe year I nature ofbreeding) have is canied out but never published, such practices appear to be com-
led to more interest in female reproduction' The

become available on Intemet if not in most libraries. A list of cur-

this trend is likely to continue in future. given in Table l. Attempts should be
iographies on compact discs, and they
The number of publications on male reproduction was slightly more le. An electronic camelid bibliography
or equal to those on female reproduction until 1988. However, since is reported to be developed (Agag 1996). A list of currently available
1989 there has been three to four times more publications on the bibliographies is given (Table 2).
Research on dromedary reproduction - The past two decades andfuture prospective r267

Table 1. Web sites available on camel literature

S. No Name and address of the web site

I htp ://www. came lnet.e thz.ch/
2 http ://www. africaonline.co.ke/AfrricaOnline/derby/about.html
a
J http ://www.nwf.org/ni ck/camelco. htm I

4 http ://www.agri.uaeu.a c.ae I agril apineVcamels.html

5 http ://www.nuffi c.nl/ciran/ikdm/ I -3larticles/Kohler.html

6 htg ://www. llamaweb. com/cameUwelcome.html

7 http ://www. arab-heritage.com/camelrace.htm

8 htp :/iwww. austcamel.com

9 htp ://www. arab.net/camels/welcome.html
l0 htp ://sites.netscape.neVsomalivet/camel
ll http ://www. ansi. okstate.edu/breeds/otheriCAMEL/drom/

t2 http :i/www.netiran.com/Htdocs/Clippings/Sociali9803 0I XXSO03.html

l3 http ://www. allcamels. com

Table 2. Bibliographies cunently available on camel literature

Bibliography Name Year of publication Publishing Agency Citations/pages

Camelids l98l ACSAD, Arab 2539 cit.
bibliography Centre for the
studies of Arid
Zones & Drylands,
Damascus, Syrian
Arab Republic
Bibliographie Sur I 990 Cirad EMVT BP 4812 cit.
Le dromedaire et le 50355, 34032
chameau Montpellier, Cedex,
France.

Research Scenario 1995 National Research 676 cit.

and Bibliography of Cente on camel,
Research Jorbeer, Bikaner,
Publications in Rajasthan India.
Indian camels
Future Outlook r996 Cairo University, 1095 cit.
through Cairo, Egypt.
documentation of
camel research in
Egvpt
Electronic camel 1996 Department of 32768 querries
bibliography on Theriogenology,
camel reproduction Behna University,
Egvpt.
Camel 1997 Jordan University 450 pages
Encyclopaedia ofScience and
Technology, Jordan

Contributing Countries
India and Egypt appear onlhe top of the list of countries contributing
towards research with nearly equal number of publications (89 and
79 respectively). This appears to be related to the fact that both the
countries have equal number of their national journals (7 and 5) in
which they have contributed nearly equal number of publications (49
and 42) forming about 50% of their total research publications.
Together these countries have contributed 34.35 % of the total
research carried out and published. After India and Egypg Sudan
have contributed 12.47% of the total research publications followed
by United Arab Emirates (11.04%) which is now emerging as a
center of excellence in this field. Saudi Arabia (5.72Vo) and Israel
(4.29%) are the other important countries contributing research.
Overall the above six countries have contributed a major share
(66.05%) of the total research canied out in the past twenty-three
years.
Continent wise analysis reveals that the African continent has con-
tributed the maximum 40.28%o (197\489), followed by Middle East
29.03% (142\489) South Asia 20.85% (102\489) and others 9.81%
(48\489) of the total research carried out. Countries like France, Ger-
mdfl), Australia and U.K. where dromedaries were previously
unknown have contributed with significant findings during the recent
past.

1268
Research Highlights and Areas of Future Interest
On scanning the research findings the key research outcome was
identified and areas of future interest recognized, which are discussed
by topic.
Male reproductive anat omY
The fact that spermatogenesis is present all year round in the drome-
dary (Singh and Bhardwaj, 1978) was an important research finding.
Besides this other studies included those on testicles (Singh and
Bhaedwaj, 1978; El-Jack, 1980) and work during 1988 and 1989
confirmed an important previous finding that seminal vesicles are
absent in the dromedary and that the prostate is the largest and the
only palpable gland (Ali et al.,l978ub:' Djang et al.,1988). Based on
histo-chemical and ultrastructural studies the steroidogenic activity
of the poll glands was suggested, and, it was also suggested that the
gland accumulates testosterone (Tingari and George,1984;' Tingari,
et a|.,1984).
Male Reproductive P lrys io I ogt
Embryonic development of male genitalia and the mechanism and
chronology of testicular descent remains unstudied. Some authors
refer to the dromedary as a non-typical seasonal breeder (Ali et al.,
1978; Osman et al.,1979) because there is no part of the year when
the testes are incapable of spermatogenesis. The anatomical and
endocrinological evidence of seasonality was demonstrated in the
male and the androgens, thyroid (Yagil et al.,1978), pituitary activity
(which shows refractoriness to GnRH during summer), and cortisol
(which appears to increase significantly after rutting Agarwal et a[.,
l99l) studied. Studies on prolactin levels showed that they were high
during non-breeding (Osman and Ploen, 1986) season. Other studies
included semeniferous epithelial cycle, androgen production by poll
glands, mating and copulatory behaviour, including soft palate ejec-
tion, agressiveness and marking.
The mechanisms of onset of seasonality, role of epididymis in sperm
maturation and the role of poll glands still remain poorly explained
and need attention of researchers in future.
Male Examination
Milestones in male examination including special examination of
genitals using techniques like ultrasonography were recently
described in detail (Tibary and Anouassi 1997), whereas others
investigated methods of semen collection, like those by artificial
vagina (AV) (Merkt, et a1.,1990; Willmen, et a1.,1993) and electro-
ejaculation (Tingari et al., 1986). As known, the semen contains a
gelatinous fraction (Tingari et al.,1986) highly viscous in nature and
the ejaculation is dribbling in nature. Biochemical fractions of the
semen have been identified (El-Manna et al., 1986; Tingari et al.,
1986). Other research included objective (Tibary et al., 1992) and
subjective (Musa et al., 1993) evaluation of sperrn motility and
normal sperm morphology (Tingari, l99l; Tingari et a|.,1986).
The major problem with breeding soundness examination is the lack
of standard methods, not only for semen collection, but also for
examination. This should form the core of future research if camels
are to be artificially bred and propagated globally by means of artifi-
cial insemination.
Male Reproductive P athologt
There were very few reports.. It was emphasized that the affections of
the penis and prepuce are seldom found due to the well-protected
anatomic position, however prepeutial affections were described
(Gahlot, 1992, Nigam 1992). Orchitis, cryptorchidism, testicular
hypoplasia (Homeida et al., l985ab; Homeida et al., 1985) epidi-
dymal affections were other described conditions. Surgical
procedures on the male were described in detail (Tibary and
Anouassi, 1997) including vasectomy, epididymectomy and others.
Looking to the extensive use of males with excellent racing perform-
ance for breeding, it becomes more important to better understand
v
Veterinary Bulletin 2000 Vol.70 No. l2
male reproductive disorders and to treat such valuable animals.
Studies on male pathology should be encouraged for future studies.
F emale Repr odttc tive Anat omy
Important studies include scanning electron microscopy of the ovi-
duct (Nayak, 1977) and endometrium (Fetaih et al., 1992) md
detailed mammary gland morphology (Smuts and Bezuidenhou!
1987). Other studies included those demonstrating that the corpora
lutea are present in mated females or pregnant animal's only (El-
Wishy, 1992; Tibary and Anouassi, 1996) and that they regress
within 10-12 days. The anatomy of the four types of follicular struc-
tures on the ovary was described by Tibary and Anouassi (1996).
Female Reproductive P hys i ol og,t
Female reproductive physiology has received most attention during
the past two decades in an attempt to increase understanding of the
female reproductive processes and to enhance reproductive perform-
ance. It was fairly concluded that phases of the oestrous cycle
described for species with spontaneous ovulation (oestrus and dioe-
strus) do not exist in Camelidae, unless the female is bred and has
ovulated. In the absence of mating there is only a succession of fol-
licular waves with highly variable rhlthm (Tibary and Anouassi,
1996). The signs of oestrus were described (Elias et al., l984ub;
Elias et al.,1985; Homeida et a1.,1988), however only 55o/o of the
dromedaries were found to exhibit clear signs (Joshi, et al., 1978) and
it was concluded (Anouassi et al.,1994; Tibary and Anouassi, 1996)
that the signs cannot be used to predict the time for breeding. Sea-
sonal effects on fertility and ovarian follicular activity has been
recently described to be due to low aromatase activity in the granu-
losa cells of early breeding season follicles (Shigri and Driancourt,
1999).
The milestones of research understanding were the induced nature of
ovulation, finding of an epidermal membrane amongst fetal enve-
lopes, the exclusive left hom pregnancies and the difference in the
luteolytic properties of prostaglandin's released from the left and the
right uterine horns.
The hypothesis of induced ovulation was confirmed by endocrino-
logical and clinical studies (Anouassi, 1984) and it is now established
that copulation causes the hormonal changes (LH surge) necessary
for ovulation. Although not precisely known in the dromedary,
studies on other species including the Bactrian camel suggest that the
factor that induces ovulation is present in the seminal plasma (Pan er
al., 7986; Chen er al., 1985; Anouassi et al., 1992) and is a protein
(Pan er al., 1995). Others have suggested a neurohormonal pathway
(Anouassi, 1984). The incidence of ovulation following mating
varies from 60-100% (Anouassi et al., 1992;Tibary and Anouassi,
1996 and Skidmore et al.,1996b). Most authors suggest the interval
from breeding to ovulation as between 32-40 ohrs (Marie and
Anouassi, 1986; Marie and Anouassi, 1987; Tibary..and Anouassi,
1996). Spontaneous ovulation (Marie, 1987) has been speculated to
be leutinization of follicles in non-mated females (Tibary and
Anouassi, 1996). The optimum time to mate or to attempt ovulation
in the female dromedary was described in a recent studv (Skidmore
et al.,1997c).
Some other interesting research findings were, that the active corpora
luteum does not inhibit the gromh of new follicles (Tibary and
Anouassi, 1996), difference in the luteolytic properties of the pros-
taglandin from the left and the right uterine horn (Fernandezet al.,
l9T9,demonstrated in the alpaca). The matter of debate on exclusive
left horn pregnancies rvas concluded on the basis of ultrasonographic
studies that showed that the pregnancy rate is similar whether ovula-
tion occurs in the left or the right ovary (Tibary and Anouassi, 1996;
Anouassi et al.,1994\.
An extra fetal membrane termed the epidermal membrane was
reported to be present in camels. It is not known when this membrane
first appears, but in dromedaries it can be identified at 3 months of
gestation (Hussein et al., l99l). Due to this membrane the fetus is

Research on dromedary reproduction - The past two decades andfuture prospective
never directly in contact with the amniotic fluid, and the membrane
is thought to lubricate the birth canal and prevent dehydration of the
fetus. A recent study described the placentation during the first 60
days of gestation (Skidmore et al.,l997b,a; Skidmore et al.,1997a)
and the maternal recognition of pregnancy (Skidmore et al.,1997b).
Other research studies and findings included those on the endo-
crinology of the oestrous cycle and pregnancy including LH (Bravo
et al., 1990; Agarwal and Khann4 1998a), FSH (Anouassi et al.,
1987) oestrogens (Elias et al.,1984a; Cristofori et a1.,1986; Cristo-
fori and Quarant4 1990; Alfurazi, 1998), progesterone (Elias et al.,
1984a; Tibary and Anouassi, 1996; Alfuraiji, 1998), thyroid hor-
mones (Agarwal et al., 1986; Agarwal et al.,
1989) and
corticosteroids. It has been shown recently that release of PGF2-
alpha controls luteolysis in camels, however in contrast to other
ruminants, release of endometrial PGF2-alpha in non-pregnant
camels does not appear to be under the control of oxytocin (Skidmore
et al., 1998). Studies on embryo development showed that the
embryo descends between day 6 and 7, then following elongation,
the embryo implants at around 50 days (Skidmore et al.,1992b; Skid-
more et al., 1996a). Trans-uterine migration of embryos takes place
and fetal growth occurs mostly between the last 3 months of gestation
(El-Wishy et al.,l98l). Studies on parturition and the post-partum
period (Mus4 1983; Musa and Makawi, 1985; Elias and Cohen,
1986; Chriqui, 1988) were important and necessitate their mention.
Pregnancy diagnosis using hormone assay has been described with
limited success (Agarwal et al., 1998a; Mohmat et al., 1997). The
nycto-hemeral rhythm of melatonin secretion in the camel has been
described recently (Vyas et al., 1997) as has been the patterns of
maternalmicro-vasculature, blood flow system, size and form of cap-
illaries of the placenta (AbdelNaeim et a|.,1998).
Female Examination
By far the most important milestone in female dromedary examina-
tion was the development of ultrasonography. After it became
available for mares in the late 1970s, Schels and Mostafawi (1978)
were probably the first to describe the use of the technique for preg-
nancy diagnosis in the camel and since then it has been used for
physiological studies on ovarian and uterine activity (Chriqui, 1988;
Tibary and Anouassi, 1996), diagnostics and pathological studies.
This technique and other means of female examination including
drugs for chemical restraint, and other practical procedures were
described (Tibary and Anouassi, 1997). Other studies include vaginal
and uterine cytology (Musa, 1983; Aaden et al.,1990) but concluded
to be of little diagnostic significance, uterine culture and biopsies
(Hegary and Salim, 1979, Ali et a1.,1987; Wernery and Ali, 1989;
Enany et al.,1990) which depicted various infectious agents and the
changes in the endometrium, techniques of hormone assay using
radio-immunoassay and enzyme immunoassay (Anouassi, 1984;
Anouassi et al., l99l) and cytogentic studies (Vijh er al., 1990;
Tibary and Bakkoury 1994). The use of fibroscopy has been
described for the study of embryonic development, and gross evalu-
ation of the uterine cavity (Tibary and Anouassi, 1997) whereas
laproscopy has been described in new world camelids (Bravo and
Sumar, 1989; Anderson et a1.,1996) but not in the dromedary.
Female Reproductive P athologt
There a serious lack of studies on this aspect. The causes of infer-
"vas
tility were poorly described. Conditions like repeat breeding in
dromedaries are difficult to describe, but females that fail to conceive
after 2 breeding may be considered in this category (Tibary and
Anouassi, 1997). Reasons for fertilization failure have been sparsely
described (Tibary and Anouassi, 1996; Egbe-Nwiyi zurd Chaudhary,
1994). Conditions like cystic ovaries, par-ovarian cysts were
described from abattoir studies (El-Wishy and Homeida 1984; El-
Wishy, 1987; El-Khouly et al.,1990; El-Wishy, 1990; Ribadu et al.,
l99l; Al-Ani et a1.,1992; Ahmed and Noda, 1993), conditions like
ovarian hypoplasia (El-Wishy. 1990), hydro-bursitis (Tibary and
Anouassi, 1997), ovaro-bursal adhesions (Musa, 1984) ovarian
tumors (El-Wishv and Homeida- 1984: El-Khoulv et a1..1990). El-
\
1269
Wishy (1990) described to be mostly teratomas), and pyometra
(Ribadu et al., l99l; Chauhan and Kaushik, 1992) were described.
Recent studies however cover some causes of abortion including bru-
cellosis (Wilson et al.,1980; Yagoub et al.,1990), uterine infections
(Wemery and Ali, 1989; Wemery, l99l; Wernery and Wemery,
1992), toxoplasmosis, trypanosomiasis (Egbe-Nwiyi and Chaud-
hury, 1994), twinning, stress and abdominal trauma. Vaginal
prolapse (Arthur and Rahim, 1982; Gitao and Akabwai, 1989; Arthur
and El-Tigani, 1990), uterine torsion Oligam et al.,1977; Arthur and
El-Tigani, 1990; Elias, l99l), fetal maceration (Gahlot et al., 1983;
Ribadu et al.,l99l) and uterine prolapse were a few other conditions
reported.
Female Reproductive Surgery and Obstetrics
l99l; Hopkins et al.,
Caesarian section (Purohit et a1.,1985; Elias,
l99l; Purohit et al.,1999b), ovariectomy and hysterectomy have
been described in the dromedary but the procedures were poorly
explained. Female obstetrics and surgery have been described (Mus4
1979a; Gera and Datt, l98l; Tibary and Anouassi, 1997) but this area
lacks support of studies, except for those on normal parturition
(Mus4 1979b;Musa, 1983). Fraternal twins have been recently put
on record (Purohit, 1999) so has been hydrocephalus (Abubakar, er
al.,1998) and hypocalcaemia (Straten, 1999).
Aspects of female reproduction appear to be complex in camels and
there is a need for further research.
Artificial Breeding including Embryo Transfer
Reproductive biotechnologies have been recently reviewed by
Purohit (1999a) Advances in artificial breeding in dromedaries arose
from the desire to breed from animals performing excellently on the
racetrack. Artificial Insemination using liquid or frozen semen is still
in the preliminary stage owing to various constraints. Semen has
been collected using an artificial vagina (Singh, 1989; Hassan et al.,
1995), the dilutions recommended are 1:3 (Anouassi, e/ a1.,7992:
Musa et al., 1992) and there is difficulty in long-term preservation
due to the gelatinous nature of the semen. Commercial bull extender
has been recommended for dilution (Sieme et a|.,1990). Recto-vag-
inal deposition of semen is recommended (Anouassi er al., 1992).
The problems of semen collection, evaluation, dilution, freezing,
optimum dose for insemination, site of insemination and others are
yet to be resolved properly before dromedary insemination becomes
commercially viable.
Collection of embryos for multiple ovulation and embryo transfer
(MOET) have been attempted since 1970's, the first live llama from
non-surgicaltransfer was bom in 1985 (Weipz and Chapman, 1985),
it is claimed that the first live dromedary calf was bom from embryo
transfer in 1990 at UAE although not repofted. Reports on embryo
transfer depict that probably the first were carried out'in Israel (Yagil
and Etzion, 1984) in 1984. Plenty of reports have gathered thereafter
(Anouassi and Ali, 1990; Cooper et al., 1990; Cooper et al.,1992:'
McKinnon and Tinson, 1992 Skidmore et al., 1992a; Anouassi and
Haaz4l993; Ismail et a1.,1993; McKinnon et a1.,1994; Vyas et al.,
1998). Ovine FSH was used for super-ovulation in doses of 20-30
Units for six divided doses, the protocols being similar to those used
for the bovine species (Cooper et al.,1990; Cooper et al., 1992; Skid-
more e/ al.,1992a). The results with these treatments and those using
porcine FSH (McKinnon et al., 1994) have been poor in terms of
embryo recovery. Ilorse chorionic gonadotropin (eCG) has also been
used in doses of 1500-6000 I.U as a single dose after a progestagen
treatment (Anouassi and Ali, 1990; Cooper et a\.,1990; Yagil and
Van- Creveld, 1990; McKinnon and Tinson, 1992; McKinnon e/ a/.,
1994) however the responses have shown individual variations.
Breeding of superovulated donors is difficult and should be started
when follicles are above l0 mm in diameter. Interesting research
findings include the introduction of the embryo into the uterus
between day 6-7 (Tibary and Anouassi, 1997), and folding and elon-
gation of embryos by day 9 after ovulation (Anouassi and Ali, 1990;
McKinnon et al., 1994). Embryo recovery has previously been
Research on dromedary reproduction - The past two decades andfuture prospective
never directly in contact with the amnioJic fluid, and the membrane
is thought to lubricate the birth canal and prevent dehydration of the
fetus. A recent study described the placentation during the first 60
days of gestation (Skidmore et al.,l997b,a; Skidmore et al.,1997a\
and the maternal recognition of pregnancy (Skidmore et al.,1997b).
Other research studies and findings included those on the endo-
crinology of the oestrous cycle and pregnancy including LH (Bravo
et al., 1990; Agarwal and Khann4 1998a), FSH (Anouassi et al.,
1987) oestrogens (Elias et al.,1984a; Cristofori et a1.,1986; Cristo-
fori and Quarant4 1990; Alfurazi, 1998), progesterone (Eliu et al.,
1984a; Tibary and Anouassi, 1996; Alfuraiji, 1998), thyroid hor-
mones (Agalwal et al., 1986; Agarwal et dl., 1989) and
corticosteroids. It has been shown recently that release of PGF2-
alpha controls luteolysis in camels, however in contrast to other
ruminants, release of endometrial PGF2-alpha in non-pregnant
camels does not appear to be under the control of oxytocin (Skidmore
et al., 1998). Studies on embryo development showed that the
embryo descends between day 6 and7, then following elongation,
the embryo implants ataround 50 days (Skidmore et al.,l992b; Skid-
more e, al.,1996a). Trans-uterine migration of embryos takes place
and fetal growth occurs mostly between the last 3 months ofgestation
(El-Wishy et al.,l98l). Studies on parturition and the post-partum
period (Mus4 1983; Musa and Makawi, 1985; Elias and Cohen,
1986; Chriqui, 1988) were important and necessitate their mention.
Pregnancy diagnosis using hormone assay has been described with
limited success (Agarwal et a1.,1998a; Mohmat et a1.,1997). The
nycto-hemeral rhythm of melatonin secretion in the camel has been
described recently (Vyas et al., 1997) as has been the patterns of
maternal micro-vasculature, blood flow system, size and form of cap-
illaries of the placenta (AbdelNaeim et a1.,1998).
Female Examination
By far the most important milestone in female dromedary examina-
tion was the development of ultrasonography. After it became
available for mares in the late 1970s, Schels and Mostafawi (1978)
were probably the first to describe the use of the technique for preg-
nancy diagnosis in the camel and since then it has been used for
physiological studies on ovarian and uterine activity (Chriqui, 1988;
Tibary and Anouassi, 1996), diagnostics and pathological studies.
This technique and other means of female examination including
drugs for chemical restraint, and other practical procedures were
described (Tibary and Anouassi, 1997). Other studies include vaginal
and uterine cytology (Musa, 1983; Aadenet a1.,1990) but concluded
to be of little diagnostic significance, uterine culture and biopsies
(Heguy and Salim, 1979, Ali et a1.,1987; Wernery and Ali, 1989;
Enany et a1.,1990) which depicted various infectious agents and the
changes in the endometrium, techniques of hormone assay using
radio-immunoassay and enzyme immunoassay (Anouassi, 1984;
Anouassi et al., l99l) and cytogentic studies (Vijh er al., 1990:
Tibary and Bakkoury 1994). The use of fibroscopy has been
described for the study of embryonic development, and gross evalu-
ation of the uterine cavity (Tibary and Anouassi, 1997) whereas
laproscopy has been described in new world camelids (Bravo and
Sumat, 1989; Anderson e/ al.,1996) but not in the dromedary.
Female Repr oductive P at hologt
There was a serious lack of studies on this aspect. The causes of infer-
tility were poorly described. Conditions like repeat breeding in
dromedaries are difficult todescribe, but females that fail to conceive
after 2 breeding may be considered in this category fiibary and
Anouassi, 1997). Reasons for fertilization failure have been sparsely
described (Tibary and Anouassi, 1996; Egbe-Nwiyi and Chaudhary,
1994). Conditions like cystic ovaries, par-ovarian cysts were
described from abattoir studies (El-Wishy and Homeida, 1984; El-
Wishy, 1987; El-Khouly et al.,1990; El-Wishy, 1990; Ribadu et al.,
l99l; Al-Ani et a1.,1992; Ahmed and Noda, 1993), conditions like
ovarian hypoplasia (El-Wishy, 1990), hydro-bursitis (Tibary and
Anouassi, 1997), ovaro-bursal adhesions (Musa, 1984) ovarian
tumors (El-Wishy and Homeida 1984; El-Khouly et a1.,1990). El-
1269
Wishy (1990) described to be mostly teratomas), and pyometra
al.,l99l; Chauhan and Kaushik, 1992) were described.
(Ribadu et
Recent studies however cover some causes of abortion including bru-
cellosis (Wilson et al.,1980: Yagoub et al.,1990), uterine infections
(Wemery and Ali, 1989; Wemery, l99l; Wemery and Wernery,
1992), toxoplasmosis, trypanosomiasis (Egbe-Nwiyi and Chaud-
hury, 1994), twinning, stress and abdominal trauma. Vaginal
prolapse (Arthur and Rahim, 1982; Gitao and Akabwai, 1989; Arthur
and El-Tigani, 1990), uterine torsion (Nigam et al.,1977; Arthur and
El-Tigani, 1990; Elias, 1991), fetal maceration (Gahl ot et al., 1983;
Ribadu et al.,l99l) and uterine prolapse were a few other conditions
reported.
Female Reproductive Surgery and Obstetrics
Caesarian section (Purohit etal., 1985; Elias, l99l; Hopkins et al.,
l99l; Purohit et al., 1999b), ovariectomy and hysterectomy have
been described in the dromedary but the procedures were poorly
explained. Female obstetrics and surgery have been described (Mus4
1979a; Gera and Datt, I 98 I ; Tibary and Anouassi, 1997) but this area
lacks support of studies, except for those on normal parturition
(Mus4 1979b;Musa 1983). Fraternal twins have been recently put
on record (Purohit, 1999) so has been hydrocephalus (Abubakar, er
al.,1998) and hypocalcaemia (Snaten, 1999).
Aspects of female reproduction appear to be complex in camels and
there is a need for further research.
Artificial Breeding including Embryo Transfer
Reproductive biotechnologies have been recently reviewed by
Purohit (1999a) Advances in artificial breeding in dromedaries arose
from the desire to breed from animals performing excellently on the
racetrack. Artificial lnsemination using liquid or frozen semen is still
in the preliminary stage owing to various constraints. Semen has
been collected using an artificial vagina (Singh, 1989; Hassan et al.,
1995), the dilutions recommended are l:3 (Anouassi, er al., 1992;
Musa et al., 1992) and there is difficulty in long-term preservation
due to the gelatinous nature of the semen. Commercial bull extender
has been recommended for dilution (Sieme et a\.,1990). Recto-vag-
inal deposition of semen is recommended (Anouassi e/ al., 1992).
The problems of semen collection, evaluation, dilution, freezing,
optimum dose for insemination, site of insemination and others are
yet to be resolved properly before dromedary insemination becomes
commercially viable.
Collection of embryos for multiple ovulation and embryo transfer
(MOET) have been attempted since 1970's, the first live llama from
non-surgical transfer was born in 1985 (Weipz and Chapman, 1985),
it is claimed that the first live dromedary calf was $om from embryo
transfer in 1990 at UAE although not reported. Reports on embryo
transfer depict that probably the first were carried out')n Israel (Yagil
and Etzion, 1984) in 1984. Plenty of reports have gathered thereafter
(Anouassi and Ali, 1990; Cooper et a1.,1990; Cooper et al., 1992;
McKinnon and Tinson,1992; Skidmore et al.,1992a; Anouassi and
Haaza, 1993; Ismail et al., 1993; McKinnon et al., 1994;Yyas et al.,
1998). Ovine FSH was used for super-ovulation in doses of 20-30
Units for six divided doses, the protocols being similar to those used
for the bovine species (Cooper et al.,1990;Cooper et al., 1992; Skid-
more et al.,1992a). The results with these ffeatments and those using
porcine FSH (McKinnon et al., 1994) have been poor in terms of
embryo recovery. Horse chorionic gonadotropin (eCG) has also been
used in doses of 1500-6000 I.U as a single dose after a progestagen
treatment (Anouassi and Ali, 1990; Cooper et a1.,1990; Yagil and
Van- Creveld, 1990; McKinnon and Tinson, 1992; McKinnon et al.,
1994) however the responses have shown individual variations.
Breeding of superovulated donors is diffrcult and should be started
when follicles are above l0 mm in diameter. Interesting research
findings include the introduction of the embryo into the uterus
between day 6-7 (Tibary and Anouassi, 1997), and folding and elon-
gation of embryos by day 9 after ovulation (Anouassi and Ali, 1990;
McKinnon et al., 1994). Embryo recovery has previously been
1270
reported to be as low as 1.55 per donor (Cooper et a1.,1990; Skid-
morc et al., 1992) are now known to bp improved to 6.5 per donor
(Tinson et a1.,1998) but the protocols are still far from perfect.
Freezing of the dromedary embryo has been attempted (McKinnon
and Tinson,1992; McKinnon et a1.,1994) but yet not standardized,
although some reports have started to appear (Skidmore and
Loskutoff, 1999). Invttro fertilization [VF) in camelid species as a
whole is in the preliminary stage with only a few reports (Bou et al.,
1993; Del Campo et al., 1995; Brogliatti et al., 1996; Purohit et al.,
1999a). Preliminary studies on oocyte maturation (Bou e/ al.,1993)
showed that a longer time (36 hrs) is required for maturation of
oocytes. Reports on development of blastocysts invitro are cunently
not available in the dromedary although work is in progress.
Besides these biotechniques, photoperiodic confiol of reproduction
has been recently reported with limited success (Agarwal & Khann4
1998b). Use of advanced molecular techniques in camels include
DNA analysis techniques (Sher'eif and Alhadrami, 1996), parentage
and identity testing (Scott et al., 1992), analysis of archaeological
remains (Stephans, 1995), chromosomal homology between drome-
dary and humans (Abdo et al.,1997), blood protein polymorphism
(Ouragh & Bengoumi,1996) and sequencing analysis of prion genes
(Kaluz et al.,1997).
Alex Tinson of UAE and Bruce Harrison of Australia have recently
claimed to produce a live dromedary calf from a pre-sexed embryo.
The scientists also claim to have obtained twin pregnancies in two
animals from a single bisected embryo (Journal of Camel Practice
and Research, June 1999).
Despite all ofthe work mentioned above, there is work yet to be done
artificial insemination, embryo transfer, and in vitro fertilization in
the dromedary before it becomes an important commercial reality
from a research tool. Research on various aspects related to these
techniques although in progress at various laboratories, but need to
be geared up.
Conclusions
The review clearly reflects that the data on dromedary research
remains scattered and hence efforts should be made to collate the
dat4 which would include creation of global and regionaljournals on
camels, as well as those specifically on camel reproduction. There
should be wider circulation of conference and seminar proceedings,
as well as encouragement of scientists to publish their unpublished
data. Also, camel research data should be made available on the
Internet and on CD-ROMs.
Male reproduction has been less studied hence work on this area
should be encouraged. The areas of future interest should include
development of standard methods for semen collection, dilution,
preservation and insemination, studies on the role of the epididymis
in sperm maturation, role of poll glands and mechanisms of onset of
seasonality. Female reproductive research should bedirected
towards detailed studies on reproductive pathology and reproductive
physiology.
Research on reproductive bio-techniques should include standardiza-
tion of embryo transfer techniques including standard protocols for
superovulation, embryo collection and embryo cryo-preservation.
IVF and subsequent embryo transfer offers a potential tool for
improving genetic stock and should be developed for dromedaries, as
it has for other species. A global gene pool for dromedary semen and
embryos should be set-up.
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@ CAB Intemational2000