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Macropus Eugenii J The effect of pregnancy on the interval between one oestrus and the next in the tammar wallaby, Macropus eugenii J. C. Merchant Division of Wildlife Research, CSIRO, P.O. Box 84, Lyneham, A.C.T. 2602, Australia Summary. In females mated to vasectomized males the interval between oestrous periods was 30\m=.\6\m=+-\1\m=.\17(s.d.) days but when mated to intact males and pregnancy intervened, it was 29\m=.\4\m=+-\1\m=.\26days (P < 0\m=.\025).After removal of pouch young, females carrying diapausing blastocysts gave birth 26\m=.\2days later and came into oestrus 26\m=.\4\m=+-\0\m=.\57(s.d.) days later. When post-partum fertilization was prevented, removal of pouch young was followed by oestrus 30\m=.\4\m=+-\0\m=.\99days later (P < 0\m=.\0005).These results indicate an influence of the conceptus upon the oestrous cycle of Macropus eugenii. Introduction It is well known that the young marsupial exerts a powerful influence on the mother after birth, but in a review of marsupial reproduction Sharman (1970) points out that it has long been thought that it has no effect on the mother before birth and that the placenta has no endocrine function. However, the placenta of the tammar wallaby exerts a morphogenetic effect on the endometrium (Renfree, 1972), possibly as a result of the indirect evidence that the yolk sac placenta of the quokka and the tammar in vitro has the capacity to convert pregnenolone to progesterone (Bradshaw, McDonald, Hahnel & Heller, 1975; Renfree & Heap, 1977). Since gestation in these species is accommodated within the duration of one oestrous cycle it seemed probable that the endocrine function of the placenta is limited to effects on the immediate environment of the uterus but may not affect ovarian events, particularly the life-span of the corpus luteum or the next oestrus and ovulation. Indeed, Tyndale-Biscoe, Hearn & Renfree (1974) proposed that the control of pregnancy and the oestrous cycle in macropodid marsupials may be independent of each other except for the initial part of the cycle when both are controlled by the young corpus luteum. In the agile wallaby, Macropus agilis, which has a reproductive pattern similar to that of the tammar, M. eugenii (Group 2 of Sharman, Calaby & Poole, 1966) it was observed that the interval between two consecutive oestrous periods was significantly shorter when a pregnancy intervened than it was in unmated females (Merchant, 1976). Since the data were not obtained from sequential observations on the same females, individual differences or the effect of copulation could have obscured the effect of pregnancy itself. The present experiments were undertaken to test the hypothesis that some aspect of pregnancy can hasten the onset of oestrus in the tammar wallaby. Materials and Methods All the animals used in the experiments came from a breeding colony of tammar wallabies, held in CSIRO enclosures at Canberra, A.C.T., and derived from animals caught on Kangaroo Island, South Australia. They were kept outdoors in two large pens grassed with Phalaris tuberosa and were given a supplementary feed mix of ground lucerne, hay and oats. Downloaded from Bioscientifica.com at 09/28/2021 10:41:14AM via free access AU females were at least 2 years old and had bred before the start of the experiments. The males were adults of proven fertility, 2 of which were vasectomized 1 month before the experiments. The experiments were carried out in series over 2 breeding seasons, early January to late July (Sharman & Berger, 1969), in 1976 (Series 1) and 1977 (Series 2). Series 1 Eleven females that were known to have been with males during a post-partum oestrus and could be expected to be carrying a diapausing blastocyst had their young removed from the pouch (RPY) to initiate reproductive activity culminating in birth and post-partum oestrus (Text- fig, la). This provided the first set of intervals—RPY to post-partum oestrus—in which there was an intervening pregnancy. At post-partum oestrus, however, the females were allowed access only to vasectomized males, thus preventing fertilization and aiding diagnosis of the day of oestrus. When the pouch young were removed 8-15 days later, therefore, there was no diapausing blastocyst present to be activated and thus no intervening pregnancy before the next oestrus occurred. RPY |o RPY · (a) U il I I Jan. Feb. Mar. Apr. May Jun. 1976 lo RPY O · I· <C) _ -h J_I_I_I Jan. Feb. Mar. Apr. May Jun. 1.977 Text-fig. 1. Diagram of the experimental procedure in females of (a) Series 1, (b) Series 2, i.e. those which gave birth at the end of the non-breeding season, and (c) Series 2, i.e. those which did not give birth at the end of the non-breeding season. RPY = removal of pouch young; = birth; O = oestrus with vasectomized male; · = oestrus with intact male; solid line = period of daily examination; hatched bar = pregnant cycle; hollow bar = non-pregnant cycle. Series 2 Twelve females, without pouch young, were examined daily from the end of the non-breeding season until birth and/or mating occurred: 6 gave birth to young derived from blastocysts carried over the non-breeding period (Text-fig. lb) and all 12 mated with vasectomized males. The pouch young of those females which gave birth were removed 8-15 days later to initiate the delayed cycle of reproduction. At the next oestrus they were again mated with vasectomized males and underwent non-pregnant cycles. At the end of this cycle they were mated to fertile males and underwent pregnant cycles. The 6 females which mated with a vasectomized male without giving birth (Text-fig. lc) then entered non-pregnant cycles, at the end of which they were mated to intact males and provided pregnant cycles. In order to minimize disturbance to the females they were kept together in groups of no more than 6 in the same pen throughout one series of experiments; pairs of intact or vasectomized males were introduced to or removed from the pens as required. Furthermore, examination of Downloaded from Bioscientifica.com at 09/28/2021 10:41:14AM via free access females was restricted to once per day and not begun until Day 24 after RPY, from which time birth and oestrus could be expected from previous knowledge of the reproductive cycle of this species (Berger, 1970). These examinations were continued on each of 4 days after mating. The day of birth ± 12 h was determined by examining the pouch for newborn young and the day of oestrus from examination of vaginal smears, using the methods of Poole & Catling (1974). A plug of coagulated semen is frequently found in the urogenital sinus and is the best evidence of copulation but spermatozoa and prostatic bodies in the vaginal smear provided additional confirmation. In Series 1 the onset of the cycle was fixed precisely by the time of RPY and the intervals to birth and oestrus were accurate to ±12 h. In Series 2, however, the initial oestrus was known only to an accuracy of ±12 h and so the intervals to birth and next oestrus were accurate to ±24 h. Results The results of the Series 1 experiments are presented in Table 1. The mean intervals from RPY to birth and RPY to post-partum oestrus were the same because in most animals birth and post- partum oestrus occurred within the same 24-h period. From other studies the actual interval from birth to onset of oestrus is known to be 10-16 h (Tyndale-Biscoe & Rodger, 1978). The interval from RPY to oestrus in the same females after being mated to vasectomized males was 4 days longer, the difference being highly significant (P < 0-0005). Table 1. Intervals (days) from removal of pouch young (RPY) to birth, to next oestrus and to the next oestrus after no pregnancy in female tammar wallabies Mated to intact male Mated to vasectomized male Animal RPY to birth RPY to next oestrus RPY to next oestrus 3945 26-5 26-5 30-5 3947 26-5 26-5 31-5 3955 27-5 27-5 31-5 3956 26-5 26-5 28-5 3975 25-5 26-5 30-5 4032 25-5 25-5 29-5 3960 26-5 26-5 31-5 3721 25-5 26-5 29-5 3892 26-5 26-5 30-5 4158 25-5 25-5 30-5 3825* 28-5 28-5 30-5,38-5,43-5 Overall 26-2 ± 0-67 f26-4±0-57 f30-4 ± 0-99 mean ±s.d. (N = 10) * Female excluded from analysis (see text). t Means used in one-tailed / test; observed t value 11 11 ; < 0-0005. · One female (No. 3825) was excluded from the analysis because she exhibited oestrous changes in the vaginal smear by Day 31 but did not mate then. However, she mated on Day 39 and Day 44 after RPY. As no other female was observed to mate other than on the day of oestrus, as determined by vaginal smear, the behaviour of this female cast doubt on when oestrus occurred. Nevertheless, whichever figure was used it would have been in excess of the interval for her pregnant cycle, 28-5 days, and if included would have exaggerated the difference between the two conditions. Downloaded from Bioscientifica.com at 09/28/2021 10:41:14AM via free access The results of Series 2 are presented in Table 2 and again show the close proximity of post- partum mating to birth. In this series the difference between the two intervals was less marked, being about 1 day, but it was statistically significant (P < 0-025).
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