Animal Reproduction Science 124 (2011) 194–199

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Animal Reproduction Science

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Reproductive cycles of buffalo夽

B.M.A.O. Perera ∗
Department of Farm Animal Production and Health, Faculty of Veterinary Medicine & Animal Science, University of Peradeniya, Peradeniya, Sri Lanka

a r t i c l e i n f o a b s t r a c t

Article history: The domestic water buffalo (Bubalus bubalis) has an important role in the agricultural econ-
Available online 24 September 2010 omy of many developing countries in Asia, providing milk, meat and draught power. It is
also used in some Mediterranean and Latin American countries as a source of milk and
Keywords: meat for specialized markets. Although the buffalo can adapt to harsh environments and
Buffalo live poor quality forage, reproductive efficiency is often compromised by such conditions,
Reproduction
resulting in late sexual maturity, long postpartum anoestrus, poor expression of oestrus,
Oestrus cycle
poor conception rates and long calving intervals. The age at puberty is influenced by geno-
Fertility
type, nutrition, management and climate, and under favourable conditions occurs at 15–18
months in river buffalo and 21–24 months in swamp buffalo. The ovaries are smaller than in
cattle and contain fewer primordial follicles. Buffalo are capable of breeding throughout the
year, but in many countries a seasonal pattern of ovarian activity occurs. This is attributed in
tropical regions to changes in rainfall resulting in feed availability or to temperature stress
resulting 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 greater
variation than observed in cattle. The signs of oestrus in buffalo are less overt than in cattle
and homosexual behaviour between females is rare. The duration of oestrus is 5–27 h, with
ovulation occurring 24–48 h (mean 34 h) after the onset of oestrus. The hormonal changes
occurring 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 waves
during an oestrous cycle varies from one to three and influences the length of the luteal
phase 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, while
swamp types with free suckling do so at 60–90 days. However, in many farming systems
prolonged postpartum anoestrus is a major problem, and the causes include poor nutrition
and 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 in
cattle, using various combinations of prostaglandins, progesterone releasing devices, GnRH
and eCG, but success rate is poor when treatment is done during the periods of marginal
breeding activity or seasonal anoestrus.
© 2010 Elsevier B.V. All rights reserved.

1. Introduction

The domestic water buffalo (Bubalus bubalis) is an

important livestock resource in many countries of Asia,
the Mediterranean region and Latin America. The world
夽 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.
∗ Corresponding author. http://faostat.fao.org/), of which 96% are in Asia. Water buf-
E-mail address: [email protected]. falo are classified in to two main ‘types’: the river type

0378-4320/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.anireprosci.2010.08.022
 B.M.A.O. Perera / Animal Reproduction Science 124 (2011) 194–199 195

located in South Asia and the swamp type spread across systems. In Europe, buffalo of the Mediterranean type pre-
the South-East Asian region (Cockrill, 1974). The Mediter- 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 (Jainudeen and Hafez, The reproductive physiology and endocrinology of
1993; Barile, 2005). This is manifested mainly as late matu- domestic buffalo and the comparative aspects with cat-
rity, long postpartum anoestrous intervals, poor expression tle were comprehensively reviewed by Dobson and
of oestrus, poor conception rates (CR) and long calving Kamonpatana (1986), and subsequent advances including
intervals. However, studies in Sri Lanka (Perera et al., 1987), those relating to the application of reproductive technolo-
Pakistan (Usmani et al., 1990) and Brazil (Vale, 1996) show gies were reviewed by Madan et al. (1996), Zicarelli (1997a)
that fertility can be highly acceptable, provided genotypes and Perera (1999). 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-
aged and fed properly. duction and production was published by FAO (Borghese,
2005) 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
different farming systems Presicce (2007) and Perera (2008). The present review deals
with 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
different regions of the world (Cockrill, 1974; Perera et brief overview of methods for controlling the reproductive
al., 2005) and are determined by several interacting fac- cycle.
tors that include climate (tropical or temperate, humid or
arid), location (rural, peri-urban or urban), cropping sys- 2. Puberty and the mature reproductive organs
tems (rain-fed or irrigated, annual or perennial crops), type
of operation (small or large farm, subsistence or commer- 2.1. Puberty
cial), and primary purpose for buffalo production and/or
management (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. from 18 to 46 months (Jainudeen and Hafez, 1993). The
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 (Borghese, 2005) shows that under favourable conditions
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- disadvantage.
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 protect
against 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 (Danell,
The production systems are mostly extensive or semi- 1987) 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
fallow fields and communal lands. when inactive and 4 g when active, with fewer tertiary fol-
The majority of buffalo in North Africa and the Middle licles (Danell, 1987; Zicarelli, 1997a). When palpated per
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, pus luteum (Jainudeen et al., 1983). The corpus luteum
for specialized production of milk and beef under intensive is smaller than that in cattle, often does not protrude
196 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 (Kaur and
identification of ovarian structures by rectal palpation in Arora, 1982; Nanda et al., 2003).
buffalo more difficult than in cattle (Perera et al., 1987; El- The duration of oestrus is similar in river and swamp
Wishy, 2007). 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 (Baruselli et al., (mean 14 h) after the end of oestrus (Kanai et al., 1990;
1997; Brito et al., 2002; Yindee et al., 2010). Perera, 1999). In hot climates duration of oestrus tends to
be shorter and signs of oestrus may be exhibited only dur-
2.3. Tubular tract ing the night or early morning. In Italian buffalo different
durations of oestrus have been observed (Zicarelli, 1997a)
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,
(Perera et al., 1987). The cervical canal is narrower than 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 with oestrus and ovulation

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- (Dobson and Kamonpatana, 1986; Perera et al., 1987; Singh
toperiod is relatively constant, annual changes in rainfall et al., 2001). 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 nancy (Perera and Abeyratne, 1979; Jainudeen et al., 1983;
the onset of monsoonal rains, followed by conceptions that Roy and Prakash, 2009). For assessing cyclic ovarian activ-
result in a peak calving season 10 months later (Perera et al., ity and for early diagnosis of non-pregnancy, progesterone
1987). 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 (Kaur and Arora, tive of the presence of luteal function and those below
1982) and the Amazon region of Brazil (Vale et al., 1990). 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 clusive.
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- atively less than that in cattle (Avenell et al., 1985; Kanai
terone secretion (Roy and Prakash, 2007). In temperate et al., 1990; Roy and Prakash, 2009). Although this has
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-
and mediated by melatonin secretion (Zicarelli, 1997b; lation (Zicarelli, 1997a; Borghese, 2005).
Borghese, 2005). The effects of these and other factors on The data available on the concentrations of
postpartum ovarian activity are discussed in Section 5 of gonadotrophic hormones in blood of buffalo indicate
the present review. that the temporal patterns of both LH and FSH are basically
similar to those in cattle (Avenell et al., 1985; Kanai et al.,
4. Oestrous cycles 1990), with a preovulatory LH surge occurring on the day
of 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
oestrus to ovulation has been observed in animals that conceived
to AI and those that did not, being 25 ± 13 and 46 ± 18 h,
The duration of the oestrous cycle in buffalo is similar respectively (Moioli et al., 1998).
to that in cattle, ranging from 17 to 26 days with a mean A study in which timing of oestrous cycles among Mur-
of around 21 days (Jainudeen and Hafez, 1993). However, rah buffalo was synchronised (Singh et al., 2001) peak
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) 194–199 197

For buffalo with two-waves of ovarian follicular develop-

ment during oestrous cycles, the growth rate and diameter
of the largest follicle was significantly smaller in heifers
than cows for both the first follicular wave (1.3 compared
to 1.7 mm per day and 10.5 compared to 13.3 mm, respec-
tively) and the second follicular wave (1.0 compared to
1.3 mm per day and 11.0 compared to 13.8 mm).
A recent study on suckled Thai swamp buffalo (Yindee
et al., 2010) has shown that the first postpartum ovula-
tion was followed by a short oestrous cycle (10.2 ± 0.38
days) in 84% of the animals, but the prevalence of these
decreased thereafter. The mean diameters of ovulatory fol-
Fig. 1. 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 of
an 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
secretion in buffalo. 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 4.4. Changes in reproductive organs
depicted in Fig. 1.
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
oestrous cycle the vestibular mucosa (Danell, 1987; Kanai et al., 1990).
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
buffalo has been reviewed by Manik et al. (2002) and is 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. down (Perera et al., 1977) or with the urine (Kanai et al.,
Studies on Murrah buffalo in Brazil (Baruselli et al., 1990). Thus in buffalo that are housed or tethered, detec-
1997) 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 mucus.
0 = ovulation) in all categories of animals, while the sec-
ond wave emerged on days 10.8 and 9.3 for the animals 4.5. Oestrous behaviour
with 2 and 3 wave cycles, respectively, and the third wave
emerged 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- rare (Danell, 1987; Perera, 1987; Vale et al., 1990). In the
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 (Barkawi urine, but these are not consistently exhibited by all ani-
et al., 2009). 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
and mixed parity cows (Presicce, 2004). Following an 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-waves
of 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. pleted in 25–35 days after calving (Jainudeen and Hafez,
198 B.M.A.O. Perera / Animal Reproduction Science 124 (2011) 194–199

1993; Perera et al., 1987). The stimulus of suckling shortens procedures (Zicarelli et al., 1997; Brito et al., 2002)
involution time (Usmani et al., 1990). 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 improved fertility (De Rensis and López-Gatius, 2007).
management conditions (Dobson and Kamonpatana, 1986; A review of studies using various treatment pro-
Jainudeen and Hafez, 1993). Under optimal conditions buf- tocols (Borghese, 2005) shows that CR achieved with
falo resume anoestrus by 30–90 days, but factors such as prostaglandins either alone or in combination with GnRH
poor nutrition and body condition (Baruselli et al., 2001), ranged from 7 to 56%, while CR following the use of a pro-
suckling management (Perera et al., 1987; Usmani et al., gesterone releasing devices either alone or in combination
1990) and climate (Nanda et al., 2003), which also influ- 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 of 33–60% (Baruselli et al., 1999; Paul and Prakash, 2005).
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
150–200 days (Perera et al., 1987). 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
buffalo by El-Wishy (2007) showed that it is a major cause 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) treatment at implant removal (De Rensis and López-Gatius,
and cessation of oestrous cyclicity occurred after the first or 2007).
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 (Perera, 2008): (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-
(Nanda et al., 2003). 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 (Mohan et al., be having oestrous cycles.
1990). 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 (Singh et al., 2006).
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
cycles of buffalo”.
by calves and alleviating heat stress by permitting wal-
lowing or use of water sprinklers (Perera et al., 2005). The
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