STUDIES ON INDIGENOUS ZEBU C A W . 1 . REPRODUCTIVE PATTERN UNDER TRADITIONAL MANAGEMENT

On-farm trials to determine the re~roductive p at tern. birth weight. , growth rate, age at puberty, age a t first calving, duration of postpartum anoestrus and gestation lenrrth ofindigenous zebucattle were conducted at three dry zone locations, namely Anamaduwa, Bandagiriya and Kandalama. Calving was seasonal with 66% calvings occurring during the months of October to January which coincided with North-East monsoon rains. The mean calving interval was 380.0 + 36.6 days with majorityofcalvinginte~als (81%) being less than 365 days. Mean birth weights of calves a t Anamaduwa and Bandagiriya were 15.8 f 2.5 kg and 15.6 k2.6 kg respectively. Average prepubertal growth rate of heifers was 107 5 208 g per day. Average ages at puberty at Anamaduwa, Bandagiriya and Kandalama were 845.1 f 138.8, 870.9 f 92.8 and 854.7 f 85.4 days respectively. Average length of gestation was 281 + 5 days. The ages a t first calving at Anamaduwa and Kandalama were 1248.1 f 130.2 and 1172.5 f 57.4 days respectively. Postpartum involution of the uterus was completed by 24 + 5 days. Lengths from calving to fust elevation of plasma progesterone concentration in trial I and I1 at Anamaduwa were 41.7f 39.3 and 62.1 f 27.1 days respectively, and the values for Bandagiriya and Kandalama, respectively were 94.5 f 16.6 and 64.2f 27.3 days. Majority of animals conceived a t the first postpartum oestrus. The length from calving to first elevation of plasma progesterone, calving to first observed oestrus and calving to conception intervals a t Bandagiriya were greater (p<0.05) than those at Anamaduwa and Kandalama. This may be due to free suckling by the calves at Bandagiriya as opposed to limited suckling practised a t the other two locations. Results of this s&dy suggest that indigen& zebu cattle under the dry zone management and enviimonmental conditions reproduce at a high degree of efficiency following slow sexual maturity.


INTRODUCTION
Cattle indigenous to Sri Lanka are of Bos indicus type ("Zebu cattle") and hence called indigenous Zebu.They form the majority of the island's cattle population of 1.7 million1 and are largely concentrated in the dry zone.They are managed extensively a s large herds along with indigenous buffaloes with minimum i n p ~t s .~-~ This low-input operation brings modest income to the farmers, primarily from sale of milk and animals for meat.They appear to be well adapted to the marginal environmental conditions which prevail in the dry zone and form the basis of a very sustainable mixed crop-livestock agricultural production system.Indigenous cattle are characterized by small body size and stature and by relatively poop development of hump and dewlap as compared with most other tropical ~a t t l e .~ Although the origin of the native cattle in Sri Lanka is not certain, fragmentary evidence suggests that they may be the dependants of a primitive herd which a t one time populated South Asia.Isolation in the island and absence of large scale importation of other types of cattle resulted in the perpetuation of these cattle in the ~o u n t r y .~ Apart from very early reports6J the indigenous Zebu cattle have not been subjected to much scientific investigation.
The objective of this study was to determine the reproductive characteristics such as calving pattern, weight a t birth, prepubertal growth rate, ages a t puberty and first calving, gestation length and calving interval of the indigenous Zebu cattle maintained under the traditional dry zone management conditions.

METHODS AND MATERIALS
Locations: Three locations in the d r y zone, viz.Anamaduwa,Bandagiriya and Kandalama were selected for the study.At all threelocations, males and females were grazed together on communal lands and provided with night paddocks.The rainfall was seasonal a t all three locations with precipitation ranging f?om 875 to 1875 mm per annum.Cows were milked once a day in the morning in the presence of the calfto provide a stimulus for milklet down.At Anamaduwa and Kandalama the calves were left behind in the paddock while the adults were taken for grazing, while a t Bandagiriya they were sent with the adults ta the grazing land and separated only in the evening.
Prepubertal Period: Birth weight measurements were obtained from calves from Anamaduwa (n=33) and Bandagiriya (n=17) using a weighing balance with a sensitivity of 0.5 kg.Growth rate and age at puberty were studiedin heifers aged 12-16 months a t Anamaduwa (n=10), 20-31 months a t Bandagiriya (n=19) and 25-30 months a t Kandalama (n=15).Body weight was measured monthly using a weigh band.Blood samples for plasma progesterone assay were collected ten days apart every two months a t Anamaduwa and ten days apart every month a t Bandagiriya and Kandalama.

Postpartum period and gestation length:
Cows were selected a t calving from Anamaduwa (study I, n=10; study 11, n=12), Bandagiriya (n=18), and Kandalama (n=18).Their reproductive tracts were palpated rectally a t seven days postcalving and blood samples were collected weekly until the next pregnancy was confirmed by palpation.Animals in study I a t Anamaduwa were also palpated a t weekly intervals from 7 days postpartum for ovarian size and structures, and changes in uterine size and position.At Anamaduwa 18 animals were monitored for duration of pregnancy by blood sampling a t monthly intervals.

~b l v i n ~ p a t & r n :
. ~t Anamaduwa, 15 farmers Were instructed to record calving dates over 24 months.They were visited a t monthly intervals to verify and collect the data.
Blood Sampling: Blood samples were obtained by jugular venepuncture using heparinized vacutainers and were kept on ice until centrifugation a t 3000 rpm for ten min.Plasma was stored a t +5OC until transported to the laboratory where it was stored at -20°C until assayed for plasma progesterone by radioimmunoassay (RIA).
Progesterone assay: Plasma was subjected to RIAusing a solid-phase coated-tube method employing 12610dingprogesterone as tracer (Joint FAOIIAEA Division of Nuclear Techniques in Agriculture, Vienna).The assay has previously been validated in our l a b o r a t ~r y .~ The sensitivity of the assay was 0.1 nglml and the intra and inter-assay coefficients ofvariation were 9.7% and 13.7%, respectively.

Statistical analysis:
The estimates of various parameters were tested for differences among locations with Student's 't' test and ANOVA.

RESULTS
Prepubertal Period: The mean birth weight, growth rate and age a t puberty in each trial are given in Table 1.The mean birth weights were 15.8 +2.5 kg a t Anamaduwa and 15.6 * 2.6 kg a t Bandagiriya (no significant difference, p>0.05).
The average prepubertal growth rate of heifers was 107+208 glday, with fluctuations related to the time of the year (Fig. 1).There was greater loss of weight during the months of August to October 1990, coinciding with a severe drought.The average body weight at first oestrus was 147 + 23 kg.The average age at puberty, as determined by the first elevation of plasma progesterone with or without observed oestrus, at Anamaduwa, Bandagiriya and Kandalama were 845 + 138,870 * 93 and854 + 85 days,respectively (p>0.05).
The ages at observed .oestruswere 972 + 124, 907 + 113 and 936 + 108 days, respectively for the three locations.Seventy percent ofthe animals at Anamaduwa, 58% ofthe animals at Bandagiriya and 60% ofthe animals at Kandalama showed short duration of elevated plasma progesterone concentrations prior to the observed oestrus periods.The age at first calving for Anamaduwa, Bandagiriya and Kandalama were 1248 + 130,1159e63 and 1172 +57 days respectively.
Gestation Length: The average length of gestation was 281 k 5 days with a range of 276 to 291 days.The mean plasma concentration ofprogesterone was 3.0 * 1.2 nglml during gestation.
Postpartum Period: The involution of the uterus was completed by 24 + 5 days (range 18 to 34).The ovaries were very small and were estimated by palpation to be 10 to 15 mm long, 5 to 15 mm wide and 5 to 10 mm high.The follicillar development was hardly detectable and the palpation of a corpus-luteum was possible only in 62% of the cases where plasma progesterone was elevated.
Selected postpartum plasma progesterone profiles are shown in Figure 2. The intervals from parturition to first elevation of plasma progesterone, to first observed oestrus, and to conception are given Table 2.In the majority of animals progesterone elevations were preceded by behavioural oestrus, and high proportion of animals conceived at the first oestrus.The intervals from calving to first elevation of progesterone, first observed oestrus and conception at Bandagiriya were greater than those at the other two locations (p<0.05).The percentage of animals which remained anoestrus over a prolonged period was highest at Bandagiriya.
Calvingpattern and Calving intervals: Atotal of696 calvings were recorded.The majority of calvings (66%) occurred during the months of October to January, with the peak in December (Figure 3).The calving season commenced with the onset of the North-East monsoon rains.The monthly frequency of calvings was correlated with the rainfall (r=0.85).The average calving interval was 338 + 36 days with a range of 298 to 515 days (n=254); 81% of calving intervals were less than 365 days.

DISCUSSION
This is the first study on characterising the reproductive functions of indigenous zebu animals in their traditional management systems.Previous studies have shown that zebu cattle form the base ofthe cattle farmingin the dry z ~n e .~. ~ These animals are managed extensively as large herds on communal grazing lands with minimum inputs.The body weight at birth was ib agreement with the values reported for indigenous zebu maintained under experimental c~hditions.~The prepubertal growth rate was very low, with fluctuations reiated to the time of the year.In comparison, birth weight and growth.rateperformances of indigenous zebu are much lower than those of indigenous buff aloe^.^The lower birth weights may be due to continuous inbreeding of the animals in close localities over centuries.1°Further, this may have been due to poor rrutrition ofthe mother during gestation, which often extends over into the drier months of the year.Another reason for this low average growth rate may be the wide fluctuations which occur due to adverse environmental conditions.In this study it was shown that the animals tend to lose weight during the drier months of the year, which usually extend from July to September.The negative effects of environmental stresses such as high ambient temperature and high relative humidity coupled with poor quality of herbage on prepubertal growth has been demonstrated.ll The age at which puberty occurred in animals in this study ranged from 22 to 36 months with an average of 32 months.These values were much higher compared to those reported for Bos taurus cattle but similar to many values estimated from age at first calving for many native breeds of cattle in Asia and Africa, and for Brahman cattle in arid zones ofAmerica.12The delay in reaching puberty may be due to the low birth weight and slow growth rate during the prepubertal period; the latter could be due to a combination of low genetic potential for growth and poor nutrition during the major part of theyear.Studies have shown that the age at puberty in Zebu cattle could be advanced by improving nutrition which also resulted in faster growth rates.13 The first oestrous cycle of many heifers was preceded by a short elevation of progesterone.This may be due to a short cycle prior to normal oestrus or due to luteinization of a preovulatory follicle before onset of regular oestrous cycles.Similar observations were noted by others in Bos taurus cattle."The majority of animals became pregnant at the first oestrous cycle.The age at first calving of indigenous animals was in close agreement G t h previous reports6 but higher than that reported in Bos taurus cattle.14The gestation length of indigenous zebu cattle was similar to that of Bos taurus cattleI2 but less than that reported for buffaloes.' 6   Calving interval of indigenous zebu cattle ranged from 298 to 515 days suggesting that fertility following calving is subjected to high variation.Further, cumulative relative frequency showed that about 81% of total calving intervals were within the range of 298 to 365 days.This variation is conceivably due to the effect of time of calving on the postpartum reproduction.In this study it was shown that the calvings were seasonal with the majority occurring during the North-East monsoon as also observed in indigenousg and exotic buffaloes16 in Sri Lanka.These studies concluded that the seasonality of the calvings was due to entrainment of postpartum ovarian activity by seasonal availability of pasture and fodder which is governed by rairifall.Hencethose animals which calve early in the calving season will resume oestrus activity postpartum earlier than the animals which calve late in the calving season.The latter usually experienced prolonged postpartum anoestrus periods as they go into the drier months.The seasonal calving pattern and intercalvingintervals were in agreement with the previous reports on indigenous ~a t t l e .~, ~, ~ The fact that a very high percentage of zebu cattle have intercalving intervals of less than one year suggests that their fertility following calving is very high contrary to the general belief.These short calving intervals are achieved despite the free suckling stimulus.The low milk production of these animals and their small body size may not demand a high plane of nutrition postpartum; this may be the reason for early return to oestrus following calving.The age at calving and calving interval have been shown to be dependent heavily on environmental and management factors since the heritability and repeatability of these traits are lowJ6 Uterine involution was completed within an average of 24 days postpartum; this is relatively shorter than reported in other types of cattle17J8 and buff aloe^.'^Faster involution of the uterus could be due to its smaller size in indigenous Zebu cattle and the suckling stimulus provided by the young.The effect of milking and suckling on uterine involution has been shown to be due to more frequent and higher levels of oxytocin secretion.18 Ovaries of indigenous Zebu cattle were small and their dimensions were similar to those reported for indigenous buff aloe^^^+'^ but were smaller than those ofBos taurus cattle.l7 Due to the small ovaries, it was relatively difficult to assess follicular development rectally.
Postpartum ovarian activity, as measured by plasma progesterone concentration, commenced between 42 to 94 days in the majority of cows.In some animals, the first oestrus was followed by a short elevation of progesterone.A similar phenomenon has been reported in Bos indicus cattle elsewhere,20 Bos taurus cattle 21 and in buffaloes.lgThis could arise from a luteinized follicle21 or a short-lived corpus l ~t e u m .~~ The cows at Anamaduwa and Kandalama resumed oestrus activity early in the postpartum period compared to those a t Bandagiriya.Further, at Bandagiriya a higher percentage of animals remained in anoestrus over an extended period.The cows at Bandagiriya were subjected to ad-libitum suckling during the day time compared to those at Anamaduwa and Kandalama.Thus it is likely that the suckling stimulus and the cow-calf interaction in this group delayed the resumption of postpartum ovarian activity.This could prolong the calving to conception interval, as has been shown elsewhere in Zebu23 and taurine cattle.24Despite the suckling stimulus the majority of animals resumed oestrous activity within 100 days postpartum and conceived at the first oestrus.This enables many cows to calve once a year.The high fertility found in this study is in agreement with the findings in other countries among native cattle of Bos indicus type.25 Tropical cattle and buffaloes in general have been considered as animals with variable calving intervals ranging from 12 to 24 months in many countries.25This wide variation undoubtedly suggests that the reproductive function ofthe tropical Zebu cattle and buffaloes could be affected to a large extent by management and nutritional factors.This indeed has been demonstrated by numerous studies and the major determinants were suckling, poor nutrition and season of c a l ~i n g .~~, ~~ The present findings corroborate previous conclusions that the interactions between genotype and environment are major determinants of the efficiency of reproduction.
It can therefore be concluded that the indigenous Zebu cattle of Sri Lanka, managed under the traditional rural dry zone system with minimum inputs for feed, labour and health care are able to reproduce with a high degree of efficiency following very slow sexual maturity.This high fertility of Zebu cattle under dry zone conditions appear to be the basis for a very sustainable cattle farming system i n the dry zone.However, as the pressure on land increases, intensification will be inevitable and therefore further studies on minimizing suckling intensity and selection for traits with high heritability such as growth rate and milk yield may be very appropriate to optimize the milk and meat production from this valuable animal.
Corresponding author.Parts of this work were presented a t the final research coordination meeting of the F A O M A coordinated research programme on "strengthening research on animal reproduction and disease diagnosis in Asia through the application of immunoassay techniques", 1993, and the third annual congress of the Postgraduate Institute of Agriculture, University of Peradeniya, 1993.

Figure 1 :
Figure 1: Body weights andgrowth rates duringpmpubertal periodandaround puberty of indigenous zebu heifers in the dry zone.

Table 2 :b
Estimates of postpartum reproductive parameters of indigenous zebu cattle.First observed postpartum oestrus (days) Calving to conception interval (days) Percentage of animals with progesterone elevation prior to observed oestrus Percentage of animals conceived at the first postpartum oestrus Percentage of animals remained in anoestrus Means in the same row with different superscripts differ

Figure 2 :
Figure 2: Selected plasma progesterone profiles in postpartum cows.

Figure 3 :
Figure 3: Calving pattern of indigenous zebu cattle in the dry zone.