Reproductive cycles of buffalo☆
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 population of buffalo is estimated to be 172 million (FAO: http://faostat.fao.org/), of which 96% are in Asia. Water buffalo are classified in to two main ‘types’: the river type located in South Asia and the swamp type spread across the South-East Asian region (Cockrill, 1974). The Mediterranean buffalo, which some consider to be a third type, is derived from the river type. The main breeds of dairy buffalo belong to the river type and include the Murrah, Surti, Jafarabadi and Nili-Ravi. The swamp type has no specialized breeds but selective breeding in some countries has resulted in populations with characteristic features.
The buffalo has been traditionally regarded as a poor breeder due to having poor fertility in the majority of conditions under which they are raised (Jainudeen and Hafez, 1993, Barile, 2005). This is manifested mainly as late maturity, long postpartum anoestrous intervals, poor expression of oestrus, poor conception rates (CR) and long calving intervals. However, studies in Sri Lanka (Perera et al., 1987), Pakistan (Usmani et al., 1990) and Brazil (Vale, 1996) show that fertility can be highly acceptable, provided genotypes are matched to the environment and the animals are managed and fed properly.
Systems of buffalo production vary widely through the different regions of the world (Cockrill, 1974, Perera et al., 2005) and are determined by several interacting factors that include climate (tropical or temperate, humid or arid), location (rural, peri-urban or urban), cropping systems (rain-fed or irrigated, annual or perennial crops), type of operation (small or large farm, subsistence or commercial), and primary purpose for buffalo production and/or management (milk, meat, draught or mixed).
In South Asia, buffalo are predominantly of the river type and produced mainly for milk and meat production by small-holders in subsistence or semi-commercial farms. Intensive systems of management are practiced in irrigated and rain-fed cropping areas, while extensive systems are common in semi-arid and arid regions where free grazing is available on communal lands. The latter systems often include use of buffalo as draught animals. Peri-urban systems are found around major cities, where large herds are raised purely for milk production under intensive conditions, with cows having greater production yielding 15–20 l of milk per day at peak lactation being brought in from other areas soon after calving, and disposed after 200–300 days, when the yield drops to an uneconomical amounts.
In East and South-East Asia, the swamp type predominates and is reared mainly for draught power and meat, on small farms with integrated crop-livestock farming systems. The buffalo also serves as a capital asset to protect against economic risks such as crop failure, and features in religious and cultural events in some communities. The production systems are mostly extensive or semi-intensive, with free or tethered grazing in home gardens, fallow fields and communal lands.
The majority of buffalo in North Africa and the Middle East are of the river or Mediterranean type and are concentrated around the Nile delta. These animals are kept mainly by small-holders for producing milk and eventually beef. A few commercial operations exist in urban areas, for specialized production of milk and beef under intensive systems. In Europe, buffalo of the Mediterranean type predominate and are kept on large commercial farms under modern intensive systems for milk and meat production. In the Latin American region river, swamp and cross-bred buffalo are produced, with production systems varying from extensive beef production, through dual-purpose systems to intensively managed herds for milk production.
The reproductive physiology and endocrinology of domestic buffalo and the comparative aspects with cattle were comprehensively reviewed by Dobson and Kamonpatana (1986), and subsequent advances including those relating to the application of reproductive technologies were reviewed by Madan et al. (1996), Zicarelli (1997a) and Perera (1999). More recently, a collection of papers containing in-depth reviews on several aspects of reproduction and production was published by FAO (Borghese, 2005) and the current status of knowledge on reproduction and reproductive technologies was summarized by Presicce (2007) and Perera (2008). The present review deals with the normal reproductive cycle of the buffalo, with emphasis on some unique features in this species, and a brief overview of methods for controlling the reproductive cycle.
Section snippets
Puberty
Buffalo heifers usually attain puberty when they reach about 55–60% of their adult body weight, but the age at which they attain puberty can be highly variable, ranging from 18 to 46 months (Jainudeen and Hafez, 1993). The factors that influence this are genotype, nutrition, management, social environment, climate, year or season of birth and diseases. A review of studies from many countries (Borghese, 2005) shows that under favourable conditions river type buffalo exhibit first oestrus at
Reproductive patterns and the influence of nutrition, ambient temperature and photoperiod
Buffalo are polyoestrous and are capable of breeding throughout the year. However, in many countries a seasonal pattern of breeding activity, and consequently calving, has been observed. In tropical locations where photoperiod is relatively constant, annual changes in rainfall appear to influence oestrous cyclicity, with availability and quality of herbage related to this cyclical reproductive pattern. In the dry zone of Sri Lanka, buffalo kept under free grazing commence ovarian activity some
Duration and variability of the oestrous cycle and oestrus
The duration of the oestrous cycle in buffalo is similar to that in cattle, ranging from 17 to 26 days with a mean of around 21 days (Jainudeen and Hafez, 1993). However, there is greater variability of oestrous cycle length in buffalo, with a greater incidence of both abnormally short and long oestrous cycles, attributed to various factors including adverse environmental conditions, nutrition and irregularities in secretion of ovarian steroid hormones (Kaur and Arora, 1982, Nanda et al., 2003).
Postpartum reproductive events
As in cattle, uterine involution in buffalo is usually completed in 25–35 days after calving (Jainudeen and Hafez, 1993, Perera et al., 1987). The stimulus of suckling shortens involution time (Usmani et al., 1990).
The period of postpartum anoestrus or anoestrus is usually longer in buffalo than in cattle under comparative management conditions (Dobson and Kamonpatana, 1986, Jainudeen and Hafez, 1993). Under optimal conditions buffalo resume anoestrus by 30–90 days, but factors such as poor
Control of the oestrous cycle with exogenous hormones
Early studies on synchronization of time of oestrus in buffalo were based on those developed for cattle, aimed at either inducing premature luteolysis using prostaglandins or prolonging the luteal phase using progestagens (Perera, 1987). However, recent knowledge on the effect of ovarian follicular wave dynamics on the outcome of these procedures (Zicarelli et al., 1997, Brito et al., 2002) have prompted studies aimed at manipulating follicular development to achieve greater oestrous synchrony
Conflict of interest statement
The author has no financial or personal relationship with other people or organisations that could inappropriately influence or bias the paper entitled “Reproductive cycles of buffalo”.
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This paper is part of the special issue entitled: Reproductive Cycles of Animals, Guest Edited by Michael G. Diskin and Alexander Evans.