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ABSTRACT
Artificial insemination (Al) and embryo transfer are the two biotechnologies that have been applied to reproduction
in camelidae. Although during recent years Al has been developed in dromedary and bactrian camels, as well as in South
American camelids, methods of semen collection and evaluation are yet to be standardised. Moreover, poor sperm motility,
inefficient techniques of semen cryopreservation, inadequate procedure for timing insemination and ovulation are a few
of the turning points which when resolved in future will make Al a common place in camelidae. Global semen transport
and priorities for trait propagation will then come up. Development of MOET is the outcome of research in new and old
world camelids during the last decade. Although embryo recovery rates have been improved from 1.55/donor to 6.5/
donor, superovulatory treatments are yet far from being perfect. High variability in embryo recovery rates are thought to
be due to failure of ovulation, or abnormal leutinisation, superovulatory treatment or poor fertility and breeding management.
Pregnancy rates with fresh embryo transfer are currently too low (8-50%) and cryopreservation of embryos need to be
developed fully. Induction of ovulation is the common reproduction management technique used in camelids. Other
reproductive biotechnologies are in the primary stage of development. ln vitro fertilisation of follicular oocytes has been
sparsely reported with development of embryos upto B-16 cell stage using epididymal sperm. Limited availability of slaughter
house ovaries is one constraint which can be resolved by collection of COC's by in vivo methods. Work on such a
technique is in progress and standard protocols are in the offing. New biotechniques that are likely to be developed in
camelid reproduction in the nearfuture are gender preselection, nucleus transfer and embryo cloning, genome analysis
and gene mapping, in vitro embryo production, gamete cryopreservation and gene transfer.
Reproductive biotechnologies that have extent due to various limiting factors. Some two
been applied for camelids include artificial decades ago when the camel racing turned into
insemination (Al), multiple ovulation and embryo a serious sport (Haydn Evans and Wernery,
transfer, in vitro fertilisation and controlled 1995) concentrated efforts to improve the speed
breeding. In light of the various developments and characteristics of the dromedary began. These
constraints in the above techniques it appears and other factors led to the specif ic use of males
mandatory to analyse the current status of these and the need for artificial insemination. Gordon
biotechnologies in camelidae. ln this review an (1997) expressed that research to develop an
attempt has been made to analyse the current Al system for use in the Arabian camels is
artificial insemination, multiple ovulation and necessary to improve the productivity in terms
embryo transfer, in vitro fertilisation and controlled of meat quality, milk, riding and other purposes.
breeding and biotechniques likely to be developed Serious efforts on improvement th,rough Al in
in f uture like gender preselection, nucleus transfer dromedary camels of many of the developing
and embryo cloning, genome analysis and gene countries had been lacking on account of their
mapping and gene transfer in this species. sole use for draught purpose, however, interest
on breed conservation is now emanating. In the
Artificial Insemination bactrian camel the technique of breeding with
Although the f irst calf f rom Al was born to Al is a development of the past two decades
a bactrian camel in 1960's (Elliot, 1961) and (Chen and Yuen 1984, Chen et al, 1985; Xu ef
reports date back to as early as 1966 a/, 1985, 1987,1990 and Zhao et a|,1992, 1994,
(Fernandez-Bae,a 1 966; Fernandez-Baca and 1996). Al in camelids has been reviewed by
Calderon, 1968) in new world camelids but Al in Chaudhary (1995) and Tibary and Anouassi
camelidae has not been developed to a greater (1ee7).
SEND REPRINT REQUEST TO G.N.PUROHIT i-l
a
Methods of semen coilection and evaruation Anouassi (1992) summarised the work and
Both electroejaculation (Musa et al, lggz expressed that for short term preservation of
and 1993) and artif icial vagina (AV) (Khan and dromedary and bactrian semen ail extenders
Kohli, 1973; Singh, l g8g) have been used for containing lactose or egg yolk are suitable
collecting semen in the camelids. collection of although laiciphos appeutlo be better (sieme ef
semen in camelidae presents various problems a/, 1990). The dilution recommended is 1:1 and
due to dribbling nature of the ejaculation and the 1:3 and semen should be diluted onlV after
animal's be n of semen using the complete liquefaction and cooled slowly to 4-5"c.
AV used t=,t-t'r?""J;""?,
d ro m e d a ry 5 'irLX:
Dilution in order to obtain a known concentration
such as 50x1 06 per cent per ml has been
Anouassi e et al, l ggb; Willmen advocated by some authors (Anouass i et al,
1992). skimmed mirk extenders used for equine
semen were also found to be satisfactory. lf
semen is to be used within a few minutes of
collection, storage at 37"c or room temperature
is recommended (Anouassi et al, f SbZy. For
longer preservation in a liquid form (upto 4g
hours) semen shourd be coored srowry to 4 io soc,
which can be accomplished by placing the tube
containing extended semen In a water bath which
et al, 1996a; Sumar, 19gg) the inner liner of the is then kept in a refrigerator (Tibary and
AV having annular constrictions. A few Anouassi, 1 gg7).
modifications to this technique has been recenily
The dilution of semen for cryopreservation
reported (Lichtenwarner ef ar, 1gg6a,b). coilection
varies from 1:1 to 1:g for the llama (Graham et
of semen using electro-ejaculation have been al, 1978) and from 1:2 to 1:3 in the dromedary
described in the dromedary (Er-Manna ef ar,1gg6;
and bactrian camel (Chen et al, lggO; Graham
Graham et al, 1g7B; Merkt et al,1gg0; Merilan ef et al, 1978; Zhao et al, 1994,1 gg6) the extenders
used for deep freezing of semen have been
adapted from other spectes containing S_7%
glycerol as cryoprotectant (Che n et al, f ggO; Sun
et al, 1990; Musa et al, 1992, 1gg3; Sie me et al,
1990; Zhao et al, 1991) and the semen is
packaged in 0.25, 0.5 and 4 ml plastic srraws
(Musa et al, 1992; Sieme et al, 19gO; Willmen
and calderon, 196g) but not recommended (Tibary ef
and Anouassi, 1gg7), because of the poor quality
al, 1993) pellets (Graham et al, 197B) or
ampoules (Chen et al, 1990; Zhao et al, 1gg4,
of the ejaculate obtained and the distress caused
1996) or 1 .5 ml straws (Chen et al, 1990; Zhao
to the animar. other ress important methods of et al, 1994, 1996).Semen has been cryo_
semen collection in camelids include rectal preserved using instant freezilg emproying
dry
ice and liquid nitrogen. gesidls many otf,"r,
techniques a simple technique ol treezing semen
packaged in 0.25 or 0.5 ml straws nas been
recently described (Tibary and Anouassi, 19g7)
wherein dilution is compreted as soon as the
semen is riquefied, the diruted semen is coored
semen coilection have been described in detair by, down to 5"c over one hour and maintained at this
Tibary and Anouassi (1997).
temperature lor Z hours before packaging in
straws. The straws are placed on a ,".[ 4 cm
Semen Preservation above the liquid nitrogen surface for 1o minutes
Both short and long term preservation of and then transferred direcily into riquid nitrogen.
camelid semen have been described. Tibary and
A standard protocol for cryopreservation of
2lJunetggg
Journalof Camel practice and Research
camelid semen is yet to be developed
although f urther attention as
rack of adequate methods for
semen has been reported to be stored
(Tibary and Anouassi, 1997) to B years for 6 semen collection, poor post ejaculation
(Graham sperm
et al, 1978) without loss of motilify. No motirity, rack of standard techniques
ieport roi rreezing
long distance transport of f rozen semen of semen and difficulty in transcervical
resurtant birth of carves are avairabre.
and passage in south American camef pipette
The thawing stressed the need for further research ids. They
temperature for small straws is 37"C to:
seconds (Tibary and Arouassi 1gg7)
for 30 l.Standardise semen cof lection techniques;
,
ampoules and large straws are thawed
whereas 2.Define procedures for semen handling
at 40_55"C and
rn a water bath, for 30 seconds evaluation; 3.Standardise semen cryopreser_
to 1 minute. vation techniques; 4.Standardise insemination
Insemination procedures for optimal pregnancy
rates.
ts-
Oestrus females are difficult to detect the above, work on matJ ano ?emale Besides gamete
behavioural signs of oestrus should and physiology is required. Whife
be combined considering Al in
with assessment of ovarran activity camels for impro.ving their genetic qualit-y,
by rectal worth noting that serection lor mirr 'proi"ction it is
palpation and/or ultrasonography (Tibary
Anouassi, 1gg7). The semen deposition and could pose probtems due to the fact
site tnat camefs
recommended is the uterus, just are very late maturing and the males
cranial to the used would
internal cervical os using an insemination become very old before their darght"r,.
gun or performance are available and
pipette by re_ctal or vaginal technique
(Tibary and as such l;;; i"r,
Anouassi, 1gg7). studies on minimar storage of semen by cryopreservation
number of a viabre sorution and therefore
should be
spermato zoa per insemination are stress be attached
.required
lacking arthough for the bactrian to it (Gordon, 1gg7).
camer 400
spermatozoa are recommended (Chen mirion
1 9BS, 1 990; Zhao et
et al, Multiple Ovulation and Embryo Transfer
al, 1 991 , 1 gg4, 1 996) while
for dromedary a dose of SO million Use of multiple ovulatibn and
is suggested (Anouass t et ar, 1g92).
,p"rr"t ozoa transfer has been attempted since embryo
various (Novoa and the 197O
- problems are i
ll"
a
(Super-Ov) in a
result of using porcine FSH others on the
Superovulation the camelid decreasing dole schedule over The super-
Attempts'of superovulation in poor and embryo recovery in dromedaries' o{ the.,hCG
species have generally yielded B-11
of an ill-defined ovulation *u, inOuced on day
inconclusive retults because mating was dole thrice
Both FSH and eCG treatment at oestiur "no
oestrus cycle lGorOon, 1997)' following 5000
Due to a short ar l|hourly intlivaf s on day 1g-14 A total
have been ur"Jtotsuper rvulation'
21
cycle and induced lU of hCG at the first mating' the 7
,of
donors
luteal phase, ill-defined oestrus
phase been embryos were recovered out of
nature of ovulation, the lr teal l9u"
(Co1r91 et flushed non-surgically' The numbet ?l ,:1t"lt
induced using progestagen rmplants embryos was maximum In
and 1995) ' yielding betwe
al, 994; BourkJ et "l' 1992a
1
without
"iZ-12
this group compared equal divided
to doses of
arOerovulation qan be obtained
but the best 400 mg of Fortoiropin or 5!1nO
ot FS[-P-' Of the
progesterone o' oih"r treatment
treated
Follotropin
fs eacn in FSH-P and yielded 2
females have
results are obtained when treated "nitui.
group) only r iemafe of FSH-P groupgroups did
no follicular (Tibary and Anouassi'
embryos, rest att temates in both
the \
"ttu"t""t
1997). Other methods of inducing a luteal phase
Using.220 in
by non- not yield ry^l:]-P
include mating with a t ale followed "nv "tnorvo'
decreasing 12 hourly intervais
over a period of 5
daV 7 and initiation ovulation
surgical flushin! ot the uteri 1n days Correa et at(f
ggZ) found a mean
on day 9 followed by usual laproscopy) of f 'J+3'1 in
of superovutatlon non-surgical rate (observed by
mating, tr"utt"nt with f'CC and(lsmail et al' Itamas and the response was
higher compared
embryo r,".ou" ry 6-7 day?,later or 500 lU of
with this method were to 500 lU of eCG daily for 3 days (rvulation rate of
1993). Embryo
'Ltou"ty corpora luteal were eCG + 156 *;;isi tor 4days
of 21
1-.6i,' although a mean 5'33 A total
1.5+0.5 and i.oto'7, respeciively)' g to 47"/"
reco rded. embryos were collected correspondin by 750 lU
CL observed. Ovulation was induced
Superovulation with FSH of hCG on day 5'
^^..-:+^
ing 20 to 30 ^{ nn
units of Porctne
or over 6 daYs (two.injections
tocols similar to those used
SuPerovulation with eCG
-,a eCGhasbeenSuccessfuIlyusedfor
in' g 2 daYs befo.re and ] duv f rom 1500-
course of Progestagen induction of ,up"tovulation in dosage
1990, 1992; Skidmore 6000lUinthedromedary(Anouassia,ndAli,
et al' 1994;
1990; Cooper Lt McKinnon
in good superovulation "t,1990;
ggZ;Yagil and Creveld' 1990) 500-
Musa et al, f
et al' 1997) 750
;,',i: li:3'i:?";,Ili'{ 2000 lU in tne tlama ind (Cor rea
ef a/' 1995; Del Campo
due to poor lU in the alpaca (Bourke
embryos, respectively) probably decreasing et ai,1995). eCG is generally given-as a single
in
breeding *"n-Jg"ment' bvine
FSH the day of completion
dose one day before or on
or f ixed doses- of 1-3 mg b'i'd' for treatment
3-5 days The
progesterong. of a 5 to r S J"yt progesterone treatment' different
following a 10-15 day yielded equally follicular response was variable withltJ of ecG
have also been reporfed to haue et al' doses ot eCc ('4 + 3'2 withi15o0
(McKinnon and Ali'
good Po.nse
1997)' Small doses and 5 .7 + S.i with 2000 lU' Apouassiovulation
1994; ssi'
a single dose) followed 1990), which may be due to vaiiablepopulation
of ovi and individual u"iiution. The follicular
by an U of eCG have resulted in camelids and
'Yos recovered Per treated has been poorty studied so far population )
in an however' the hence ,"utont due to the follicular with limited
female (Skidmore et al' 1992) cannot oe ascribed to. Vyas 998)
(1
due to the effect of eCG of eCG in lndian
effects ur" iit"fy to be data reported poor response
FSH (1-3 mg bid) in
than that of FSH' Porcine dromedari"s. fwo dromedaries treated
with 3000
days after a 10-
decreasing doses over 3' 5 or 7 lU of eCG on O"y 7 of 100 mg lM
daily injections
have also shown
15 day proggsterone treatment et al' of progesteron" resulted in a mean ovulatory
(McKinnon
to sup"rouriJte a dromedary response of 5'5, however' no embryo could be
1994). Vyas (1 998) have reported
a beneficial
,r*.
a2
1992a', 1994 and 1995a,b)'
Superovulation bY immunisation
i1-+
supernatant.
June lggg /5
Journalof Camel Practice and Research
lii;J:"",Xiffi';'i"x;,;X': ,xltii:t
tH;'H "l l'";^""'iff:U'H'[: iH:T'!:':""1:ii:"1i::;:
rile,.::TT,"J3:il:3iiJ:r*:lHli![i[iil*:.fiiltt"t*'***l';li#:T[:!:J,
j*:?"3HlFi,tl*;rrn:tTJ"tlililt',;"Ff?p[;!::flilry6ii:ri,';.ti:r"J;]
:lri of such trearments lM-:l'lj:sesterone
and Anouassi, 1ee7) rnsprle ovurate and turn :::i';;:l;"'"i"jbo-'s
:;;;;r;; roricres,mav not
ifil'
"{::,li*li''l**,m;:*,W*',H,tiiii
"+?"Jlii""'u"Jtl!'"1""""'""'u trearment used i:t^l"Y'r;r, rhe reported.pr"sn,llSv rates lrom
."t"
u.o "l!ill-i"il;"lJ-'1!":_"!!'! *llU,:*m*:t[iH *U:tt#:i: ":1,?
"un "it!"t '1"
L:::T;lT";i,';Jii"lnr",ifjmtn:nl; l",8iff u;;;;;;;,"1"" "," kn:y1to ' be arrected
i:r{*r{Tl1dilillilrdfi :in"ii'r:L'l",.#n::],ii,",'=m*:"f,],li:*f i
lf ilYfl ilil ?3,J',11?l*"",", ,n" , recipilnt, however. rh-e,sde
or transfer,
';:l#"ll::
);;;;;ti;.
with
?;ffi
increased natural
,l;'J":";:l:i rlltit*:rfTi,'J:i'":?l';l:
malrnl
rransrer or em
surs :rgfi:iigi'1'l;"""1,"iffi'J&il3ry;i'ill
,^"li*Uru';h;y],",1;::lfiHfjJ'.%:i
Bp,h
Y"l,'-:'?i:l
Tinson, 1 992;
T;,
M( itJ;:,:"$ :iiln+Hi:[g11"jj"+1{* j["J,'"':.'tfilj
oi f or
e
",nOryo
but the n;-;;;i;al described in freezins camerro
;:*::r"Ut;en reported
and Researcr
Journal of Camel Practice
6/June tggg
probe (5 or
survive the low epidural anaesthesia, a vaginal
relatively large size which does not 1997)'
and
7.5 mhz) with a aspiration needle aspirate the
pump is
Ireezing process (Tibary and Anouassi' introduced through vaginal wall to
The major problems encountered with follicular contents' A recovery rate of
40-50"/o was
efficient development of embryo transfer reporteO by these workers whereas
a recovery
technology in camelidae are superovulation' rate of 64% *"' in the llama (Brogliatti
ovuIationandbreedingmanagementand '"ported from abattoir
research et al,1996). Retrieval of oocytes
cryopreservation of embryos' More bvarie u et al (1993) in
dynamics
should be directed towards follicular drome et a/ (1992 and
of follicular
nypothalamopituitary f"g'lation 1994) kers using slicing
ixed time Al'
wave cycles, ovuiation induction'
"nO f
metho SA fraction V' 25 PM
in vivo' effect of
*orpnophysiology of embryos and sodium pyruvate and 50 pg/ml gentamicin
cold temperatures on:amelid embryos 6'4
sulphate) reported an average recovery^ofwith
studies on follicular population to simplify
and
et a/ (1999)
tackle the existing oo.Vt". per Ilama' Purohit (fortif ied with
improve the proceduies and limited data using TCM 199
problems. antibiotics and BSA) described an0'7. average
(1017)
recovery ot +.0 (g214), 2'33(.1416) and
ln vitro fertilisation with oocytes per ovary by follicular aspiration'
tn vitrofertilisation has been attempted dissection and slicing oi dromedary
rom
ovaries.f
Embryos have been
limited success in camelids' dead animals. Slicing was considered unsuitable
oocytes
proOuced in vitro f rom llama follicular due to extravasation of blood into the medium of
using ePididYmal
t al' 1994)
of the oocytes were
collection. Sixty four per cent
but no live birt d' In the
attached to it'
ing to the denuded anO frLO no cumulus mass
dromedary IVF a
ed (Bou et
8-16 cell stage in OocYte Maturation
very low and
a/, 1993). Current success rates are Not much work has been done on the
no embryo has been obtained from
ejaculated of
the techniques used are adopted maturation of follicular oocytes' Maturation
COC's is generally accomplished by
semen. Most of incubation
f rom work in other sPecles'
at 38.5"C ,nO" i SX CO, tension with high
humidity in tissue culture medium supplemented
Methods of oocYte retrieval been with hormones, serum, pyruvate and
antibiotics'
Two metfrods of oocyte retrieval have unde^r such
using Sixty two per cent llama oocytes
reported in camelids' Th e in vivo methods incubation reached metaphase ll after 36 hours
laproscopy (Tibary and .1n:i":i'^,t,nnt)
pick up of culture whereas for dromedary,
coc's matured
transvaginal ultrasound guided ovum in vitro under nearty similar conditions the
maturation
(OPU) (Brogliatti et al, 1996) and the lt appears that
complexes rate was 47% (Bou et a:,1993).
methods using collection of oocyte iong.r time for'oocyte maturation is required for
(Tibary and
after overiect6my or post mortem (Del camelids co.put"d to other species
such as the
Anouassi, t OSZl,'slaughter house material
recoy"t.y f rom cow (36 hours vs 24 hours)' ;'.
Cr.po et al, lsigz ano t994) and et al' 1999)'
dead f emale camels (Purohit recovery SPerm PreParation
Amongst the in vivo methods surgical Del
through a tnitial attempts of Bou et al(1993) and
of mature follicles from the ovaries
(Tibary Campo et ai(1 99i, 1994) have been marginally
laparoscopy is described for dromedaries rates successful' Using epididymal sperm'
the former
the recovery *ill sperm
and Anouassi, 1997). HOwever, workers insemlnated oocytes caffeine
are described to be very low as
the follicles .
and
easily' Laproscopic capacitated in medium containing
become f r,ugit" and bleed in the BSA for 2 hours. After 6-7 hours' the oocytes
follicutar as[iration can be accomplished best were transferred to TCM-1 99 containing
FCS and
yet reported' Th.e
llama ano alpada but not is sodium pyruvate' A fertilisation rate of 43% with
in vivo oocyte collection technique described embryos developing to the B-16 cell stage in vitro
transvaginal ultrasound guided follicular workers washed sperm
Anouassi' 1997)' Under was reported. The latter
aspiration liiOary and
June 1 ege l7
Journalof Camel Practice and Research
(Anouassi and Ali, 199
Skidmore et al, 1996
1996). SimilarlY in the I
doses of GnRH (0.5-1
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": ;
c hen s H H, B u r t D w an d
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