Reproductive cycles of buffalo
Department of Farm Animal Production and Health, Faculty of Veterinary Medicine & Animal Science, University of Peradeniya, Peradeniya, Sri Lanka
The domestic water buffalo (Bubalus bubalis) has an important role in the agricultural econ-
omy of many developing countries in Asia, providing milk, meat and draught power. It isalso used in some Mediterranean and Latin American countries as a source of milk and
meat for specialized markets. Although the buffalo can adapt to harsh environments and
live poor quality forage, reproductive efﬁciency is often compromised by such conditions,
resulting in late sexual maturity, long postpartum anoestrus, poor expression of oestrus,
poor conception rates and long calving intervals. The age at puberty is inﬂuenced by geno-
type, nutrition, management and climate, and under favourable conditions occurs at 15–18months in river buffalo and 21–24 months in swamp buffalo. The ovaries are smaller than incattle and contain fewer primordial follicles. Buffalo are capable of breeding throughout theyear, but in many countries a seasonal pattern of ovarian activity occurs. This is attributed intropical regions to changes in rainfall resulting in feed availability or to temperature stressresulting in elevated prolactin secretion, and in temperate regions to changes in photope-riod and melatonin secretion. The mean length of the oestrous cycle is 21 days, with greatervariation than observed in cattle. The signs of oestrus in buffalo are less overt than in cattleand homosexual behaviour between females is rare. The duration of oestrus is 5–27 h, withovulation occurring 24–48 h (mean 34 h) after the onset of oestrus. The hormonal changesoccurring in peripheral circulation are similar to those observed in cattle, but the peak con-centrations of progesterone and oestradiol-17␤ are less. The number of follicular wavesduring an oestrous cycle varies from one to three and inﬂuences the length of the lutealphase as well as the inter-ovulatory interval. Under optimal conditions, dairy types man-aged with limited or no suckling resume oestrus cyclicity by 30–60 days after calving, whileswamp types with free suckling do so at 60–90 days. However, in many farming systemsprolonged postpartum anoestrus is a major problem, and the causes include poor nutritionand body condition, and stress due to harsh climates and improper management. Synchro-nization of time or induction of oestrus can be done using the same regimens as applied incattle, using various combinations of prostaglandins, progesterone releasing devices, GnRHand eCG, but success rate is poor when treatment is done during the periods of marginalbreeding activity or seasonal anoestrus.
2010 Elsevier B.V. All rights reserved.
The domestic water buffalo (Bubalus bubalis) is an
important livestock resource in many countries of Asia,the Mediterranean region and Latin America. The world
ଝ This paper is part of the special issue entitled: Reproductive Cycles of
population of buffalo is estimated to be 172 million (FAO:
Animals, Guest Edited by Michael G. Diskin and Alexander Evans.
falo are classiﬁed in to two main ‘types’: the river type
0378-4320/$ – see front matter 2010 Elsevier B.V. All rights reserved.
B.M.A.O. Perera / Animal Reproduction Science 124 (2011) 194–199
located in South Asia and the swamp type spread across
systems. In Europe, buffalo of the Mediterranean type pre-
dominate and are kept on large commercial farms under
ranean buffalo, which some consider to be a third type, is
modern intensive systems for milk and meat production. In
derived from the river type. The main breeds of dairy buf-
the Latin American region river, swamp and cross-bred buf-
falo belong to the river type and include the Murrah, Surti,
falo are produced, with production systems varying from
Jafarabadi and Nili-Ravi. The swamp type has no special-
extensive beef production, through dual-purpose systems
ized breeds but selective breeding in some countries has
to intensively managed herds for milk production.
resulted in populations with characteristic features.
The buffalo has been traditionally regarded as a poor
1.2. Previous reviews on buffalo reproduction
breeder due to having poor fertility in the majority of con-ditions under which they are raised (
The reproductive physiology and endocrinology of
domestic buffalo and the comparative aspects with cat-
rity, long postpartum anoestrous intervals, poor expression
of oestrus, poor conception rates (CR) and long calving
intervals. However, studies in Sri Lanka (
those relating to the application of reproductive technolo-
that fertility can be highly acceptable, provided genotypes
and More recently, a collection of papers
are matched to the environment and the animals are man-
containing in-depth reviews on several aspects of repro-
duction and production was published by FAO and the current status of knowledge on reproduc-
1.1. Role of the buffalo in livestock production in
tion and reproductive technologies was summarized by
present review dealswith the normal reproductive cycle of the buffalo, with
Systems of buffalo production vary widely through the
emphasis on some unique features in this species, and a
brief overview of methods for controlling the reproductive
and are determined by several interacting fac-
tors that include climate (tropical or temperate, humid orarid), location (rural, peri-urban or urban), cropping sys-
2. Puberty and the mature reproductive organs
tems (rain-fed or irrigated, annual or perennial crops), typeof operation (small or large farm, subsistence or commer-
cial), and primary purpose for buffalo production and/ormanagement (milk, meat, draught or mixed).
Buffalo heifers usually attain puberty when they reach
In South Asia, buffalo are predominantly of the river
about 55–60% of their adult body weight, but the age at
type and produced mainly for milk and meat production
which they attain puberty can be highly variable, ranging
by small-holders in subsistence or semi-commercial farms.
Intensive systems of management are practiced in irrigated
factors that inﬂuence this are genotype, nutrition, man-
and rain-fed cropping areas, while extensive systems are
agement, social environment, climate, year or season of
common in semi-arid and arid regions where free grazing
birth and diseases. A review of studies from many countries
is available on communal lands. The latter systems often
include use of buffalo as draught animals. Peri-urban sys-
river type buffalo exhibit ﬁrst oestrus at 15–18 months of
tems are found around major cities, where large herds are
age, while the swamp type do so at 21–24 months. The body
raised purely for milk production under intensive condi-
weight at which puberty is attained is strongly inﬂuenced
tions, with cows having greater production yielding 15–20 l
by genotype and is around 200–300 kg for the swamp type
of milk per day at peak lactation being brought in from
and 250–400 kg for the river type. Although buffalo attain
other areas soon after calving, and disposed after 200–300
puberty later than cattle, they have a longer reproductive
days, when the yield drops to an uneconomical amounts.
life, which tends to compensate for this early economic
In East and South-East Asia, the swamp type predom-
inates and is reared mainly for draught power and meat,on small farms with integrated crop-livestock farming sys-
2.2. Ovaries, follicles and corpora lutea
tems. The buffalo also serves as a capital asset to protectagainst economic risks such as crop failure, and features
The ovaries of post-pubertal buffalo heifers have a reser-
in religious and cultural events in some communities.
voir of only 10,000–20,000 primordial follicles (
The production systems are mostly extensive or semi-
compared with over 100,000 in cattle. The mature
intensive, with free or tethered grazing in home gardens,
ovaries are smaller than in cattle, weighing around 2.5 g
when inactive and 4 g when active, with fewer tertiary fol-
The majority of buffalo in North Africa and the Middle
East are of the river or Mediterranean type and are con-
rectum, mature follicles in swamp buffalo rarely exceed
centrated around the Nile delta. These animals are kept
8 mm in diameter, tend to protrude from the surface of
mainly by small-holders for producing milk and eventu-
the ovary and can be mistaken as an early developing cor-
ally beef. A few commercial operations exist in urban areas,
for specialized production of milk and beef under intensive
is smaller than that in cattle, often does not protrude
B.M.A.O. Perera / Animal Reproduction Science 124 (2011) 194–199
markedly from the surface of the ovary and sometimes
adverse environmental conditions, nutrition and irregular-
lacks a clear crown. These characteristics make accurate
ities in secretion of ovarian steroid hormones (
identiﬁcation of ovarian structures by rectal palpation in
The duration of oestrus is similar in river and swamp
Wishy, Ultrasonic imaging indicates mature follicles
buffalo, varying between 5 and 27 h, and ovulation occurs
range in size from 1.3 to 1.6 cm in diameter and mature cor-
about 24–48 h (mean 34 h) after onset of oestrus, or 6–21 h
pora lutea from 1.2 to 1.7 cm in diameter (
In hot climates duration of oestrus tends tobe shorter and signs of oestrus may be exhibited only dur-
ing the night or early morning. In Italian buffalo differentdurations of oestrus have been observed
The uterus and cervix of buffalo are smaller than those
and categorized as short (<12 h), medium (13–24 h), long
in cattle and the non-pregnant tract lies wholly within the
(24–48 h) and very long (>48 h). In the short and medium
pelvic cavity with the uterus usually being tightly coiled
oestrous cycles ovulation occurred after the end of oestrus,
around 6–72 h and 24–60 h after the onset of oestrus,
in cattle and is more difﬁcult to negotiate during artiﬁcial
respectively. In some of the long and very long oestrous
insemination (AI), especially in heifers.
cycles ovulation occurred before the end of oestrus.
3. Reproductive patterns and the inﬂuence of
4.2. Reproductive hormones and temporal relationships
nutrition, ambient temperature and photoperiod
Buffalo are polyoestrous and are capable of breed-
The changes in concentrations of progesterone in blood
ing throughout the year. However, in many countries a
and milk during the oestrous cycle are similar to those
seasonal pattern of breeding activity, and consequently
in cattle, but the peak concentration is relatively less
calving, has been observed. In tropical locations where pho-
toperiod is relatively constant, annual changes in rainfall
Progesterone concentration in fat-free milk
appear to inﬂuence oestrous cyclicity, with availability and
is usually below 1 nmol/l (∼0.3 ng/ml) during the follic-
quality of herbage related to this cyclical reproductive pat-
ular phase of the oestrous cycle and ranges from 3 to
tern. In the dry zone of Sri Lanka, buffalo kept under free
12 nmol/l (∼1–4 ng/ml) during the luteal phase and preg-
grazing commence ovarian activity some 2–3 months after
the onset of monsoonal rains, followed by conceptions that
result in a peak calving season 10 months later (
ity and for early diagnosis of non-pregnancy, progesterone
Similar effects of climate and nutrition on reproduc-
concentrations above 3 nmol/l are considered to be indica-
tive patterns have been observed in India
tive of the presence of luteal function and those below
1 nmol/l are considered indicative of the absence of luteal
Heat stress during the hot summer months in India is
function, with intermediate values being considered incon-
a major cause of anoestrus in buffalo and is associated
with elevated blood concentrations of prolactin, which
The concentration of oestradiol-17␤ in blood during the
is thought to inﬂuence ovarian activity as well as cause
follicular phase of the oestrous cycle also appears to be rel-
sub-fertility and repeat breeding by decreasing proges-
regions such as Italy, however, where buffalo are fed with
been suggested as a possible reason for the lesser intensity
a constant balanced diet, a distinct seasonal reproductive
of oestrus exhibited by buffalos, studies on Italian buf-
pattern is also found, and the inference from a series of
falo have shown no differences in the endocrine proﬁles
studies is that seasonality is inﬂuenced by photoperiod
of those with overt and oestrus without an associated ovu-
The effects of these and other factors on
postpartum ovarian activity are discussed in Section
gonadotrophic hormones in blood of buffalo indicate
that the temporal patterns of both LH and FSH are basicallysimilar to those in cattle
4. Oestrous cycles
with a preovulatory LH surge occurring on the dayof oestrus and lasting 7–12 h. In Italian buffalo a signiﬁcant
4.1. Duration and variability of the oestrous cycle and
difference in the interval from peak concentration of LH
to ovulation has been observed in animals that conceivedto AI and those that did not, being 25 ± 13 and 46 ± 18 h,
The duration of the oestrous cycle in buffalo is similar
to that in cattle, ranging from 17 to 26 days with a mean
A study in which timing of oestrous cycles among Mur-
there is greater variability of oestrous cycle length in buf-
concentrations of LH and FSH of 38 and 24 ng/ml were
falo, with a greater incidence of both abnormally short and
detected, respectively, during oestrus. Concentrations of
long oestrous cycles, attributed to various factors including
FSH in the plasma showed three mid-cycle elevations
B.M.A.O. Perera / Animal Reproduction Science 124 (2011) 194–199
For buffalo with two-waves of ovarian follicular develop-ment during oestrous cycles, the growth rate and diameterof the largest follicle was signiﬁcantly smaller in heifersthan cows for both the ﬁrst follicular wave (1.3 comparedto 1.7 mm per day and 10.5 compared to 13.3 mm, respec-tively) and the second follicular wave (1.0 compared to1.3 mm per day and 11.0 compared to 13.8 mm).
A recent study on suckled Thai swamp buffalo
has shown that the ﬁrst postpartum ovula-tion was followed by a short oestrous cycle (10.2 ± 0.38days) in 84% of the animals, but the prevalence of thesedecreased thereafter. The mean diameters of ovulatory fol-
Schematic representation of the hormonal changes occurring in
licles increased between the ﬁrst and second ovulation
blood of buffalo during the oestrous cycle.
(13.50 ± 0.52–14.31 ± 0.38 mm), and the mean diameter ofan ovulatory follicle and that of the resulting corpus luteum
which corresponded to comparatively less inhibin and
were signiﬁcantly larger in those that became pregnant at
elevated oestradiol-17␤ concentrations during the same
that ovulation than in those that did not become pregnant.
period, leading to the conclusion that both inhibin and
Further studies are clearly warranted to examine the rea-
oestradiol-17␤ have a feed-back regulatory effect on FSH
sons for the above differences, and to determine whether
they inﬂuence the quality of the oocyte in the dominant
A schematic representation of the hormonal changes
follicle that is destined for ovulation.
occurring in blood of buffalo during the oestrous cycle,summarized from the literature reviewed above, is
Externally detectable physical changes around the time
4.3. Waves of ovarian follicular development during the
of oestrus include swelling of the vulva and reddening of
the vestibular mucosa (Vulval swelling results in obliteration of the horizontal
Ovarian follicular growth during the oestrous cycle in
wrinkles which are present on its external surface and
this, together with vestibular reddening, can be detected by
basically similar to that in cattle, characterized by waves
regular examination of individual animals under conﬁned
of follicular recruitment, growth and regression. How-
systems. Mucus secreted from the cervix during oestrus is
ever, interesting and as yet unexplained differences in the
less copious than in cattle, does not usually hang as strands
number of follicular waves per oestrous cycle and the char-
from the vulva, but tends to accumulate on the ﬂoor of the
acteristics of follicular growth and regression have been
vagina and be discharged either when the animal is lying
reported from buffalo in different countries.
Thus in buffalo that are housed or tethered, detec-
showed that the number of follicular waves during
tion of oestrus can be aided by examining the ﬂoor near the
an oestrous cycle was one in 3% of animals, two in 63%
rear of the animal each morning and evening for signs of
and three in 33%. The ﬁrst wave commenced on day 1 (day
0 = ovulation) in all categories of animals, while the sec-ond wave emerged on days 10.8 and 9.3 for the animals
with 2 and 3 wave cycles, respectively, and the third waveemerged on day 16.8 in the latter group. The oestrous cycles
A major difference between buffalo and cattle is that
with two and three follicular waves of follicular develop-
behavioural signs of oestrus are less overt than in the for-
ment differed in the mean length of the luteal phase (10.4
mer, with homosexual behaviour between females being
compared to 12.7 days) and the interval between ovula-
tions (22.3 compared to 24.5 days). A study on Egyptian
absence of a bull, the main behavioural signs are restless-
buffalo, however, indicated the majority of oestrous cycles
ness, bellowing and frequent voiding of small quantities of
(54%) had three waves of follicular development
urine, but these are not consistently exhibited by all ani-
mals. When a bull is present, however, the bull will show
Studies on Italian buffalo indicate differences in
increasing interest in a cow that is approaching oestrus,
ovarian follicular dynamics between nulliparous heifers
and the cow will stand to be mounted by the bull during
oestrus. During periods of greater ambient temperature the
oestrous synchronization protocol using progesterone and
duration of oestrus may be shorter and the oestrual signs
prostaglandin, heifers exhibited one, two or three waves
exhibited only during the night or early morning.
of ovarian follicular development during an oestrous cycle,with inter-oestrous intervals of 8–12, 20–26 and 25 days,
5. Postpartum reproductive events
respectively. In contrast, all parous cows had a two-wavesof ovarian follicular development during oestrous cycles
As in cattle, uterine involution in buffalo is usually com-
with an inter-oestrous interval ranging from 19 to 25 days.
B.M.A.O. Perera / Animal Reproduction Science 124 (2011) 194–199
have prompted studies aimed at manipulating follicular
The period of postpartum anoestrus or anoestrus is usu-
development to achieve greater oestrous synchrony and
ally longer in buffalo than in cattle under comparative
A review of studies using various treatment pro-
falo resume anoestrus by 30–90 days, but factors such as
prostaglandins either alone or in combination with GnRH
ranged from 7 to 56%, while CR following the use of a pro-
gesterone releasing devices either alone or in combination
with eCG, and in some cases further supplemented with
ence nutrition through feed quality and availability, can
hCG or GnRH, ranged from 8 to 64%. The ‘Ovsynch’ protocol
delay this considerably. For example, indigenous buffalo in
(GnRH followed by prostaglandin 7 days later and a second
Sri Lanka raised under free grazing with abundant natural
GnRH 2 days later) has been used successfully in buffalo,
feed and calves allowed access to the dams for suckling
with synchronization of time of ovulation in 70–90% and CR
only once per day, resumed oestrous cyclicity by 30–60
days after calving, whereas those under harsher conditions
In most studies the success rate was lower when treat-
with free suckling by the calves remained anoestrual for
ment was done during the periods of low breeding activity
or during seasonal anoestrus, and various modiﬁed pro-
A review of the literature on postpartum anoestrus in
tocols have been tried to overcome these problems. The
two most effective approaches appear to be the Ovsynch
of infertility resulting in economic loss to buffalo breeders
protocol supplemented by administration of progesterone
in many countries. In Egypt, India and Pakistan only 34–49%
for 7 days between the ﬁrst GnRH and prostaglandin treat-
of animals showed oestrus during the ﬁrst 90 days after
ments, and progesterone based regimens of 10–14 days
calving, while 31–40% remained anoestrous for more than
with either GnRH or oestradiol treatment at the time of
150 days. The ﬁrst postpartum ovulation was frequently
progesterone implant insertion and prostaglandin plus eCG
followed by one or more short oestrous cycles (<18 days)
and cessation of oestrous cyclicity occurred after the ﬁrst or
second ovulation in about 25% of animals due to ovulatory
In addition to the type of protocol selected, the follow-
failure or prolonged luteal activity. In India, the incidence
ing factors must also be addressed when using any regimen
of postpartum anoestrus in buffalo herds can range from
in buffalo ((a) selection of animals that are in
20 to 80%, with the greatest incidence during hot summers
good body condition score and free from disease; (b) min-
imize stress during the treatment administration and AI,
LH secretion in buffalo remains low during the early
especially under tropical conditions, when animals may be
postpartum period and episodic pulses become detectable
herded together, tethered or moved to other locations; and
a few weeks before ovarian activity commences, with ani-
(c) where seasonal differences exist, scheduling treatments
mals that are subjected to better nutrition and restricted
for the more favourable periods or during the peak of the
suckling commencing episodic release of LH earlier than
breeding season when the majority of animals are likely to
those under poor nutrition or free suckling
The response of LH secretion to a challenge of exoge-nous GnRH at 25–35 days postpartum, measured as peak
Conﬂict of interest statement
concentration and area under the curve, is greater in non-suckled than suckled buffalo
The author has no ﬁnancial or personal relationship
Methods that are recommended for overcoming pro-
with other people or organisations that could inappropri-
longed postpartum anoestrus in buffalo include adequate
ately inﬂuence or bias the paper entitled “Reproductive
nutrition before and after calving, restricting the suckling
by calves and alleviating heat stress by permitting wal-lowing or use of water sprinklers Thepresence of buffalo bulls in a herd also has a biostimulatory
effect, reducing irregularities in oestrous cycle patterns and
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