Reproduction


The water buffalo has a reputation for being a sluggish breeder, but the average animal is so poorly fed that it's reproductive performance is unrepresentative of it's capabilities. Without reasonable nutrition the animals cannot reach puberty as early in life or reproduce as regularly as their physiology or genetic capabilities would normally allow.
With good management, adequately nourished buffaloes can reach puberty at about the same age as cattle, as early as 18 months of age in buffalo bulls. In northern Australia, Swamp females have conceived at 16 months of age43. In the herd at Punjab Agricultural University in Ludiana, India, 11 River buffalo heifers showed estrus at ages less than 18.5 months and a few came into heat when less than 15 months old44.
The water buffalo also can calve at an age comparable to that of cattle. At the Ain Shams University in Egypt a well fed Egyptian buffalo herd of several hundred animals has an average age at first calving of 27 months, 22 days45. Because of uncertainties, El Ashry and his colleagues believe that the body weight is a better indicator of sexual preparedness than age. These researchers at Ain Shamus University recommended mating heifers when they weigh 365 kg no matter what their age. Research at Punjub Agricultural University shows that heifers can be bred when they weigh over 270 kg and manifest estrus. Most animals in the Punjab Agricultural University River buffalo herd calved before 35 months, one at 28.3 months46. In one Venezuelan herd almost all heifers 20-24 months old were pregnant; virtually all calved before age 38 months, most by 30 months, and one at age 23 months47. In trials in Queensland, Australia and in Papua New Guinea it was noticed that female buffaloes (Swamp breed) came into estrus even while they were losing weight because of adequate nutrition, whereas cattle did not. Under these stressful conditions the buffalo calves also reached sexual maturity earlier and the buffaloes had a higher calving percentage and a shorter calving interval because they came back into estrus more quickly than cattle48. The age at first calving of more than 60 nutritionally poor buffaloes was 38 months in one herd and 45 months for Brahman cross cattle. Similar observations have been made in Florida. Trinidad, the Brazilian Amazon, Venezuela, and elsewhere. Although these are exceptions to the normal observations in Asia, where buffaloes seem to breed more slowly than cattle, they do demonstrate the buffalo's potential for improved breeding.
Unclean surroundings, poor nutrition, and poor management, causes a great number of early embryonic deaths and a high death rate among the calves; this also contributes to the buffalo's often low reproductive rate.

Female Reproduction


ESTRUS
Oestrus, estrus or heat is a specific period of reproductive function when the female becomes receptive. Estrus in buffaloes is manifested by changes in the reproductive system and in behavior. It usually lasts about 24 hours, but duration varies and may range from 11 to 72 hours. It occurs on an average of 21 day cycles. Determination of when a buffalo cow is in estrus is difficult because often the animal shows few external signs of "heat". This increases the chances of missing a cycle, especially for artificial insemination.
The intensity of estrus behavior in Egyptian and Indian buffaloes has been found to be much less than cows 48a. The usual weak symptoms of estrus in the normal breeding season (October to February in Asia and India) become still weaker during the hot months of summer. Unlike most of the buffalo breeds of tropical and temperate regions, the symptoms of estrus in Philippine buffaloes have been reported to be even more pronounced than cattle 48b. The problems of estrus detection in Swamp buffalo of Thailand has been described as one of the main constraints of the breeding program 48c.
Diurnal patterns of estrus behavior have been observed in most buffaloes. The Swamp usually breed at night 48d. In about 855 of Egyptian and Indian buffaloes signs of estrus have been recorded between 6 pm and 6 am 48e. However during monsoon, estrus often commences in the afternoon hours. In the Phillipines the management concept of night corrals has been used to increase reproductive performance.
Due to the high incidence of silent heat large numbers of buffaloes are left unbred. In Egyptian buffaloes 86% silent heat 48f has been reported. In Indian buffaloes, one study showed only 15% silent heat .
Work on estrus synchronisation using CIRD implants has been done in New Zealand.
ESTRUS DETECTION
Frequent urination is the most reliable symptom of estrus. The animals urinate small quantities every 4-6 minutes in some cases. Mucus vaginal discharge in quietly sitting animals can also be observed. Many buffaloes show clear or transparent cervical mucus at the onset of estrus which becomes cloudy and turbid to dirty translucent with the advancement of the estrus.
For the detection of estrus in buffaloes the application of a combination of aids has been suggested 48g. ESTRUS DURATION
In young Murrah buffaloes in India average estrus period has been observed to be 29 (24-72) hours 48h. Other studies have shown variations according to location, breed and season ...24 hrs 48iin the Philippines ...12-36 hrs in Egypt ...24-36 hrs in Bulgaria and 19.5 (with a range of 3 to 69 hrs. in Pakistan. The duration of estrus is usually slightly longer...and more regular in aged than in young buffaloes. An estrus duration of about 24 hours is usually used for practical purposes.
ESTRUS CYCLE
Similar to other bovines, the estrus cycle in buffaloes can be approximately divided into phases...
GESTATIONAL ESTRUS
The condition of gestational estrus or heat signs after a successful breeding has been observed in 6-18% of buffaloes (R). Animals which return to heat after service should always be examined for ovarian structure before rebreeding and if follicular status is vague, AI service should be placed outside the cervix rather than thru it into the uterine horn.
One important limiting factor influencing the productivity of buffaloes is the seasonality of breeding. This may be due to anoestrus or silent estrus during the non breeding season. Temperature, humidity, light periods and nutrition have all been mentioned as causes of this seasonality.
POSTPARTUM ESTRUS
The appearance of estrus signs following parturition is known as post partum estrus. The literature shows ranges of from 10 to 400 days. Again environment, nutrition and heat detection has significant influence on the length of this period. 30-60 days is considered an average return time. In Punjab, India, River buffaloes have been observed to come into estrus as early as 40 days after calving52.
ARTIFICIAL BREEDING
Based on work done at the University of Florida in the early 1980s some embryo transfer is also being done.
At the Philippine Carabao Center work on in-vitro maturation (IVM) and in-vitro fertilization (IVF) is in progress.

MALE REPRODUCTION


In general the buffalo bull is very similar to other bovines. The volume of buffalo semen ranges from 1.5 to 5.0 ml per ejaculate and semen quality is influenced by many factors. Most bulls are put into service between 18 and 24 months although environment, management and nutrition significantly influence their sexual activity. Young bulls less than 24 months, average volume (1.91 ml), initial mortality (22.8%), count of live spermatozoa (46.4%) and density of spermatozoa (107.14x10(6) per ml) indicate the poor quality of semen. This improved significantly in bulls of 2-8 years. Training of buffalo bulls for semen collection usually starts at 20-24 months.
Loss of potency has been observed by 6-7 years but many bulls remain fertile beyond 9 years however the largest period of usefulness is between 3 and 7 years 52a.A good bull can serve about 100 buffalo cows per year in natural service. In many parts of the world artificial insemination is practiced, however much of the population depends upon natural service by free living buffalo bulls of improved breed (mostly Murrah), stray nondescript bulls or entire working buffalo males. Distance of AI centers, frequent closure, and awkward times of estrus are the usual problems. Usually one buffalo bull operates in 2-4 villages having 200-300 breedable females in most parts of Nepal and India. These bulls serve a good number of animals during their active life. Some bulls are seen covering cows even after 10 years although with more enthuasism than effect.
OPTIMUM USE
In theory, the quantity and quality of semen collected successfully up to an average 8.75 ejaculates daily is almost identical. The calculated daily spermatozoa production capacity of bulls has been found to be 2.3-2.5 billion 52b. However, in practice this is not possible due to decreased libido and natural endurance problems. Thus the use of buffalo bulls for natural breeding or semen collection should be organized in a systematic manner for optimum utilization. Buffalo bulls should probably be used not more than 75 times a year....however in most areas about 75% of the services are required during 4 months of the peak breeding season. Breeding bulls should be used 3 times per week during the peak breedng season. Each mating should comprise two services with about a 15 minute interval.
ARTIFICIAL INSEMINATION
This technique for breeding buffaloes can be used for progeny improvement programs, optimum usage of good breeding bulls and ease of transportation to remote locations. AI was started in the early forties at Allahabad Agricultural Institute in India, and the first buffalo calf was born on 21 August, 1943 52c.Since the semen of each ejaculate is multiplied through the use of suitable extenders for the breeding of larger numbers of females it is essential to use semen of high quality. This increases the probability of high conception and the introduction of desired traits in the progeny.
SEMEN CHARACTERISTICS
In practice maximum rate of collection is on alternate days during the peak breeding season and twice weekly during the rest of the year.
INITIAL MOTILITY
For semen evaluation the determination of initial or mass motility of undiluted semen is an important quick test. The initial motility is either judged by the movements and waves (swirls) in a drop of semen under the microscope on a 0 to 5 scale (sometimes 0 to 10) or by counting the total and non-motile spermatozoa with the help of a hemocytometer. The former gives a rough estimate and the later provides actual percentages of motile sperm in the semen. The average mass motility of Murrah semen has been reported to be 3 to 4. Good cooling arrangements significantly improve the motility 52d.
MORPHOLOGY
A normal sperm of the buffalo is constituted of 3 distinct parts ...head, mid piece and tail. It resembles a balloon with an attached string or a tadpole. The head of the buffalo spermatozoan is more rectangular than that of cattle. Head length, breadth, and shape can be affected by extenders season, number of ejaculates and handling of the ejaculate. MOTALITY
This is determined by counting the percentage of motile sperm in semen diluted to contain about 20 million sperm per ml. The percentage of motility in fresh diluted semen of nondescript buffalo bulls was 67, which gradually decreased after storage at 4-6C. The motility was 40 after 5 days and 30 after 8 days of storage in egg yolk glucose bicarbonate extender 52e. A wide variation in motility, from 45% to 85% has been reported for the buffalo bull In the same sample ...65% in fresh whole semen, 75% in fresh diluted semen and 62% in diluted semen frozen in ministraws and then thawed was observed.
LIVE/DEAD COUNT
The percentage of live and dead sperm is determined by counting of slides prepared by a differential staining technique. The nigrosine eosin stain is widely used for differential staining of buffalo spermatozoa. A wide variation in the ratio of live and dead sperm of buffalo semen has been found. Again age, breed, frequency of collection, season, extender, temperature and handling all influence the percentage. Averages of 75% to 85% live in fresh collections are expected but 40% is commonly seen.
ABNORMAL SPERM
A large variety of abnormalities in the structure of sperms have been reported. The ratio of normal and abnormal is used as a parameter for the determination of semen quality. 2% to 20 % is seen with about 6% being average.
EXTENDERS
The development of efficient extenders for the preservation of live spermatozoa and their fertility for extended time requires knowledge of detailed chemical composition of spermatozoa and seminal plasma. A significantly lower initial content of fructose in the semen of the buffalo was observed 52f. However, later work has indicated that this is probably a handling defect. Chloride content is higher 52g and citric acid lower. Several specialized extenders for buffalo semen have been developed.
FREEZING
The life span of unprocessed spermatozoa in whole warm semen is limited to a few hours The high temperature has been found unsuitable and the rate of deterioration is rapid due to increased metabolic processes. Cooling can extend this life for several days... however if the temperature becomes too low crystallization of water occurs within the cells and they can be damaged. Therefore for freezing semen needs to be extended in a suitable medium which is little affected by the freezing process. Glycerol has been found to protect the sperm considerably from the freezing process. Deep freezing of buffalo semen was started in India in 1955. Limited research has been done on freezing buffalo semen and even after 50 years of research it is not in wide use nor of equal quality as cattle semen. About 80% recovery of motile spermatozoa on thawing of deep frozen buffalo semen in egg yolk glycine extender has been reported 52h. Pregnancy percentages from the insemination of thawed semen preserved deep frozen for several months have been reported at 40% to 70%. Reasonable quality deep frozen semen is now commercially available in some areas and it's use is spreading. Overall conception rates of 70-80 percent are reported. It is estimated that some 100,000 buffaloes are now being artificially inseminated50.

Reproductive Indexes


PREGNANCY
The pregnancy is an important reproductive function ... and should be the objective of every reproductive program . Unfortunately focus is sometimes lost and the breeding or insemination numbers are stressed and used to evaluate livestock development programs. In buffaloes, development of the fetus occurs in the uterine cornua and involvement of the right cornua is much more common than the left 50a. Corpora lutea are almost always present in the ovary of the same side. Recto-vaginal palpation of the gravid uterus is the most common method for diagnosis of pregnancy in both buffaloes and cows. Diagnosis of pregnancy in the buffalo before 2-3 months of gestation is difficult, especially for those with little experience and practice. At many AI centers the non return of inseminated animals is considered as a measure of pregnancy .... which may be true due to large numbers of available entire males. Recently progesterone levels in blood, plasma or serum, and milk has been used for diagnosis at early stages of gestation. Hopefully, the use of these techniques will provide improved evaluation of buffalo reproductive programs.
In many areas, calving is seasonal. This seems to be largely due to changes in nutrition. It may also be caused by heat stress, in either males or females, which results in a low breeding rate during the hot seasons. However, when buffalo cows are well fed and cooled, they come into estrus and will breed in any season.
CONCEPTION RATES
A wide variation in the conception rate of buffaloes has been reported by different observers. Most values are based on non return rates of the inseminated buffaloes for reinsemination. Rates of 50-60% are average for natural service and half that for most evaluated AI field services. The color and consistency of cervical mucus has a significant influence on the conception rates. When mucus changes from transparent to turbid and the crystallization pattern is normal fern like, the conception rate is highest. Many natural matings take place at night and are therefore unobserved. In one set of pregnancy diagnoses in northern Australia, the buffalo's conception rate (81%) was higher than that of the Brahman crossbreeds (70%) they were with49. Similar figures have been recorded in Venezuela49a.
GESTATION
The length of time required to carry the result of conception to it's full term under normal physiological environment is known as gestation period.The water buffalo's gestation period is about one month longer and is more variable than that of cattle. Whereas cattle give birth after about 280 days(Angus, 279,Holstein, 279-280, Brown Swiss,286), buffaloes take 300-334 days (average 310) or roughly 10 months and 10 days. In Egypt, the average period according to a study of 424 gestations, is 316+/- 8 days. The development of the fetus in buffaloes is affected by many factors such as breed, type, parity, season, nutritional status, sex of calf and geographical location )51. Wide variation in the gestation period of River(300-315 days) and Swamp (320-340 days) buffalo has been observed )51a.
PARTURATION
Dilation of the cervix, expulsion of the fetus, and detachment of the placenta are all similar to other bovines. Involution of the uterus averages about 30-50 days.
PREGNANCY RATE
The definition of pregnancy rate is: detection of estrus (%) times conception rate (%). The detection of estrus can be defined as the number of cows in estrus per 21 days divided by number of cows in the breeding group. Pregnancy rate includes all cows in the breeding group whether or not insemination has occurred. It is the best reproductive managment tool available for buffalo but not widely used.
CALVING INTERVAL
The interval between the subsequent calvings is known as calving interval or intercalving period. It is a significant reproductive trait for the assessment of the life time production potential of buffaloes. Similar to other reproductive traits the calving interval is also widely variable. Range of 334-580 days with mean of about 465 days in River buffaloes and 350-800 days with a 553 mean in the Swamp. Only under uncommon circumstances can a buffalo cow produce a calf each year. In one herd of 800 cows in Venezuela the average female buffalo over 4 produces 2 calves every 3 years53. In response to a recent questionnaire, the majority of Indonesian farmers estimated that the calving rate was between 3 and 5 calves in 5 years54. A few claimed a calf a year, some only 1 or 2 calves in 5 years. In Florida it has been noted that some buffalo cows having just calved became pregnant more quickly than cattle, so that a calf may indeed be produced each year55. Calving intervals of 14-15 months have been reported in Egypt and Venezuela55a. Regular yearly breeding has been noted also in northern Australia56.
Preliminary results in northern Australia indicate that weaning can be carried out as late as 12 months of age without any effect on conception times of the buffalo dam.57
SERVICE PERIOD
The interval between calving and conception is known service period. The shorter the service period the longer would be productive life. 100 to 150 days is reported.
DRY PERIOD
The interval between the end of lactation and subsequent calving is called dry period. A minimum dry period of 60 days has been suggested as resting period which helps in rebuilding of body reserves and mammary tissue. Dry periods of 0 to 250 days have been reported with about 150 days average.
REPRODUCTIVE FAILURES
Anestrus...Anatomical, hereditary, functional and nutritional conditions either singly or in combination produce anestrus in buffaloes. It is highest during the hot months and has characterized the buffalo as a seasonal breeder. Small, hard inactive ovaries are the rule rather than the exception in most buffalo of the world. Perhaps as newer management methods are developed we will see more information and research on this important constraint to buffalo reproduction. The major cause of infertility in Asian buffaloes is subactive ovaries. Poor feeding, nutritional deficiencies and parasites are the main cause of ovarian inactivity and excessive follicular atresia with the failure of maturation and ovulation.
OVARY PATHOLOGY
Hypojunctions, cysts, persistent CL and encapsulation of ovaries are common observations in buffalo. One should consider that many of these reports are from the observation of collected genitalia in slaughter facilities.... from old animals at the end of their useful life.
Tubular Genitalia ....Salpingitis has been reported as much higher in buffalo than cattle(R). Cervicitis, metritis, pyometra and vaginitis have all been observed but again many of these reports are from slaughter animals. A high number of iatrogenic tubular genital problems occur in AI programs because of rough handling and poor technique.
The incidence of abortions, dystocia, retained placenta, and other parturition problems in buffaloes is similar to that in cattle. Twinning is very rare; probably no more than 0.01% of buffalo pregnancies produce twins.

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david j. ligda
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Last Updated:Monday, February 16, 1998 8:24:37 PM