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Postpartum
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Genital tract involution
Involution of the vulva
Softening of the vulva is usually observed a few days before parturition but maximum dilation occurs during birth. At the end of parturition, the vulva is very edematous and remains that way for about 2 to 3 weeks. In the dromedary, the size of the vulva returns to prepartum size 16 to 42 days after parturition (mean 31.7 ± 2.5 days).(218) Involution of the vulva is faster in primiparous than pluriparous animals.(218)
Postpartum vaginal discharge (lochia) is minimal in the camelidae and only a small amount of reddish fluid is noticed during the first few days after parturition. In normal cases, this discharge becomes thicker and very infrequent within one or two weeks.(77, 183, 218)
Involution of the uterus
Uterine involution is rapid in the female camelidae due to the microcotyledonary and diffuse nature of the placentation in these species which does not cause a great loss of uterine tissue. In the dromedary, the involution of the uterus is completed as early as 20 days.(77, 183) The major decrease in size is observed during the first 10 to 15 days postpartum. Some authors have reported, however, that size of the uterus does not return to normal until 31 to 54 days postpartum (40 ± 2.1 days).(218) This process is completed in 94% of the cases between 30-50 days and in 6.7% per cent after 50 days of parturition. Primiparous females have a significantly more rapid involution than pluriparous animals.(218) The cervix remains open during the first 20 days of the postpartum period.(183) Even after complete involution, the left uterine horn tends to increase in size with each new pregnancy. It measures 80-105 mm, 100 -115 mm, and 110-140 mm in females with 2, 4, and 6-7 pregnancies respectively.(277) Involution of the uterus seems to be faster in the Bactrian camel where the uterus is completely involuted and readily palpated by 15 days postpartum.(69, 70)
In the alpaca, a rapid change in the weight of the uterus is observed during the first 10 days postpartum.(121, 232, 248) Uterine weight decreases from its original 883 g at 24 hours to one third of that at 5 days and one fifth at 10 days postpartum.(248) The diameter of the left horn decreases from 15 cm at delivery to 3.5 and 3 cm at 10 and 20 days postpartum, respectively.(248)
Postpartum ovarian activity
Corpus luteum regression
The corpus luteum of pregnancy regresses rapidly during the first week postpartum and then slowly after that to form a corpus albicans. Many of the corpora albicantia can be observed on the surface of the ovary of old pluriparous females as well-circumscribed, spherical, white, hard structures protruding from the ovary and giving it its characteristic multilobulated. irregular appearance (cf. Anatomy of the ovaries). Their diameter varies between 15 and 18 mm early after calving and then decreases to 4-8 mm later.(98, 100, 249) The consistency of corpora albicantia is hard and fibrous. Their color is gray-white but sometimes yellowish-white. In cross section, a thick, grayish-white, tough, fibrous capsule of 2-3 mm is seen surrounding a central zone of light-to dark-brown tissue. Regression of the corpus luteum probably starts just before parturition under the effect of massive secretion of PGF2α which causes luteolysis and degeneration of the luteal tissue.(103) Greatly-thickened, fibrous capsule and trabeculae together with excessive fibrosis of the tortuous arterioles, indicates advanced regression.
In the alpaca, corpus luteum size decreases rapidly during the first 10 days postpartum and then slowly between 10 and 20 days postpartum.(248)
Follicular activity
The time of resumption of ovarian function and follicular activity following parturition in camelidae is highly variable and depends on the species and rearing conditions. In the dromedary, postpartum anestrus was reported to last anywhere from 15 days to one year (Table 15).(95, 135, 158, 168, 211, 260, 264, 274)
Interval from parturition to estrus | Remark | Reference |
---|---|---|
14 to 25 days | 274 | |
39 to 42 days | based on estradiol level ( > 80 pg/ml) | 95 |
25 to 30 days | well-fed animals | 5 |
4.5 to 10 months | 110 | |
40 days | Stall-fed suckling females | 272 |
25 days | weaned females | P.O. |
23 days | 200 | |
21 to 30 days | 270 | |
14 to 30 days | 261, 262 | |
40 ± 2,1 days | 98 | |
8 months to one year | suckling females | 183 |
4.5 to 10 months | weaned females | 98, 99 |
P.O. = Personal observation
Discrepancy in duration of postpartum anestrus is probably due to several factors including lactation, sucking, nutritional status, and method of evaluation. According to some authors, estrus detection is not a reliable method for evaluation of postpartum activity because the first estrus is usually "silent".(92 262) Also, it appears that the timing of postpartum estrus and its intensity depend on the time of year.(92 262) Early and late in the breeding season, intensity of estrus is lower, and it lasts longer, than in the middle.(94, 95) According to our observations, the major factors affecting length of postpartum anestrus in the dromedary are nutritional status and suckling. In well-fed females, ovarian activity as assessed by ultrasonography is evident in 70 to 80% of the females by 30 days postpartum and conception rate is normal at 45 days. Also, all females that lose their offspring or are weaned show a mature follicle within 10 to 12 days. Similar observations were reported in the Bactrian camel with ovarian activity resuming only 3 to 12 days after weaning or losing the newborn(69) while ovarian activity in suckling females (follicles of 10 to 14 mm or high estrogen) is observed 35 to 60 days after parturition.(35, 69)
In the llama and alpaca, ovarian activity resumes within the first week after parturition.(11, 52, 55, 56, 138, 232, 236, 248) In llamas the interval from parturition to resumption of ovarian follicular activity is about 5 days and follicles can be ready to ovulate by 10 days after parturition. Successful breeding of animals can occur as early as Day 15 postpartum. In alpacas, ovarian follicles are less than 3 mm in diameter 24 hours after parturition and increase steadily during the next 10 days.
The interval from mating to ovulation (2.0 ± 0.1 days) and growth rate of the preovulatory follicle (0.8 ± 0.2 mm/day) are not affected by lactation status in the llama.(11) However, the interval between two follicular waves is shorter by 2.5 ± 0.05 days in lactating compared to non-lactating females. In addition, the size of the dominant follicle is smaller in lactating that in non-lactating females (10.4 ± 0.2 vs. 11.7 ± 0.3 mm).(11) This shows that although no lactational anestrus is observed in this species, there is a definite effect on the ovarian function that could be exacerbated it nutritional requirements are not met.
Interval from parturition to conception
Interval from parturition to first acceptance of the male is variable in the dromedary and should not be used as a criteria to evaluate ovarian activity and postpartum fertility. We have observed accidental mating as early as 2 days after parturition. Normal conception cannot be expected until the uterus is completely involuted and ovarian function is restored. The time from parturition to the establishment of these conditions is not known with precision. However, we have obtained pregnancies in several females as early as 45 days postpartum. Others have reported no conception in females mated 12 and 40 days after parturition and suggest that low conception rate is to be expected if breeding occurs before 50 days postpartum because uterine mucosa has not yet returned to its normal status before this stage.(92, 98) Our preliminary studies on the histology of the uterus in the postpartum period show that uterine conditions are compatible with embryo survival and pregnancy as early as 26 days postpartum. Also, we have been able to obtain normal fertilization rates and collection of viable embryos in females bred 3 weeks after induction of abortion at 3 to 10 months of pregnancy. In the Bactrian camel, conception was obtained as early as 10 days postpartum but follicular development was induced by eCG (PMSG) treatment.(89, 275, 276)
In alpacas and llamas, females can accept males as early as 24 hours after parturition(206, 248) but normal conception rates are not achieved until 10 to 15 days after parturition.(52, 55, 248) The breeding observed at 24 hours after parturition is only behavioral and is never associated with an ovulation. The first postpartum ovulation is at 5 days with a pregnancy rate of 30%.(55, 248) In one study, all females were bred and ovulated by Day 17 or 18 postpartum in conjunction with a second ovarian follicular waved.(55, 56) The interval from parturition to breeding also has a significant effect on the incidence of ovulation and on embryo survival beyond day 40. The proportion of alpaca females ovulating at 10, 20 and 30 days postpartum was respectively 64.5%, 76.9%, and 83.9% with conception rates of 62.5%, 87.5%, and 90.4% respectively. However, the proportion of females that experienced pregnancy loss by day 40 was 24%, 11.4%, and 6.4% for breeding at 10. 20, and 30 days postpartum.(55)
Endocrinology of the postpartum
The endocrinology of the immediate postpartum period is characterized by a fall of progesterone due to regression of the corpus luteum of pregnancy. Some authors have reported a gradual fall during the first week postpartum(20) whereas others found that progesterone levels were negligible as early as 24 hours after parturition.(120, 149) Progesterone levels in the blood or urinary pregnanediol glucoronide remain low with a short intermittent increase until the females are bred. In this case, an increase is observed 3 to 4 days following breeding showing ovulation and formation of normal coipus luteum.(53, 56, 149)
Plasma estradiol and urinary estrone sulfate concentrations closely follow follicular activity in the postpartum period. In the llama, significant estrone sulfate levels compatible with the presence of a normal follicular wave with a dominant follicle were observed between 5 and 10 days postpartum.(56,149)
No studies are available on the profile of pituitary hormones during the postpartum period in camelidae. However, from the observations on ovarian function and steroid hormone profiles, one can assume that pituitary activity (FSH and LH secretion) are re-established very early during the postpartum period. Low LH surge following breeding in the first 10 days postpartum has been observed in the llama and alpaca. This could be due to low response of the pituitary gland to copulation and may be the cause of increased ovulation failure at this stage. Suckling does not seem to have a great impact on pituitary and ovarian function in the New World camelidae, but it does seem to be an important factor in the delay of resumption of activity in the dromedary. More detailed endocrinological studies on the effect of lactation and suckling on ovarian function are needed.
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