Concentrations of progesterone,testosterone and estradiol-17β in the serum during the estrous cycle of Asian elephants (Elephas maximus)

The levels of progesterone, testosterone and estradiol-17β in serum samples from two female Asian elephants were measured for the period of 32 months from February 1987 to September 1989. Serum samples were collected weekly from unanesthetized elephants. Each elephant showed eight ovarian cycles in 32 months. Ovarian cycles, characterized by changes in concentrations of serum progesterone, averaged 16.8 ± 0.6 (mean ± SEM. n = 14) weeks in length. The changes in concentrations of testosterone in the serum showed a similar pattern to those of progesterone with a striking increase noted during the luteal phase. The highest levels of serum estradiol-17β were noted when progesterone levels showed low basal values. These results suggest that estradiol-17β may be an index of follicular maturation during the estrous cycle in Asian elephants, and that the ovaries of Asian elephants may produce testosterone in the luteal phase.     

Secretory Pattern of Inhibin During Estrous Cycle and Pregnancy in African (Loxodonta africana) and Asian (Elephas maximus) Elephants

The ovary of female elephants has multiple corpora lutea (CL) during the estrous cycle and gestation. The previous reports clearly demonstrated that inhibin was secreted from lutein cells as well as granulosa cells of antral follicles in cyclic Asian elephants. The aim of this study is to investigate the inhibin secretion during the pregnancy in African and Asian elephants. Two African elephants and two Asian elephants were subjected to this study. Circulating levels of immunoreactive (ir‐) inhibin and progesterone were measured by radioimmunoassay. Four pregnant periods of an African elephant and three pregnant periods of an Asian elephant were analyzed in this study. Circulating levels of ir‐inhibin started to increase at 1 or 2 week before the ovulation and reached the peak level 3 or 4 weeks earlier than progesterone during the estrous cycle in both African and Asian elephants. After last luteal phase, the serum levels of ir‐inhibin remained low throughout pregnancy in both an African and an Asian elephant. The mean levels of ir‐inhibin during the pregnancy were lower than the luteal phase in the estrous cycle despite high progesterone levels were maintained throughout the pregnancy. These results strongly suggest that CL secrete a large amount of progesterone but not inhibin during the pregnancy in elephants. Zoo Biol 31:511‐522, 2012. © 2011 Wiley Periodicals, Inc.     

Evaluation of progesterone and 20-oxo-progestagens in the plasma of Asian (Elephas maximus) and African (Loxodonta africana) elephants

The corpus luteum of African elephants produces high amounts of 5α-reduced progesterone metabolites (5α-pregnane-3,20-dione and 5-α-pregnane-3α-ol-20-one), whereas progesterone itself is quantitatively less important, and plasma levels of progesterone during the estrous cycle in elephants are considerably lower than those of other mammals. The objective of this study was to compare the concentration of progesterone in plasma of Asian and African elephants as determined by specific progesterone assays with those of total immunoreactive progestagens containing a 20-oxo-group (20-oxo-P). These metabolites were determined by an enzyme immunoassay using an antibody against 5-α-pregnane-3β-ol-20-one, 3HS:BSA. Plasma of non-pregnant Asian (n = 4) and African (n = 4) elephants was collected at weekly intervals for periods of 8–15 months and at random intervals during pregnancy in one Asian elephant. High-performance liquid chromatography separation of plasma samples of both species demonstrated that in the 20-oxo-P assay, 5α-pregnane-3,20-dione makes up ˜60% of the total immunoreactive material. The progesterone and 20-oxo-P values during the estrous cycle showed a parallel pattern and were significantly correlated (P < 0.001; Asian: r = 0.80; y = 3.76 × –0.10; African: r = 0.75; y = 2.66 × –0.08). Progesterone and 20-oxo-P values in Asian and African elephants were <0.15 ng/ml during the follicular phase (weeks –4 to 0) of the estrous cycle; progesterone values during the luteal phase (weeks 2–9) were 0.60 ± 0.03 and 0.53 ± 0.03 ng/ml, and the 20-oxo-P values were 2.19 ± 0.16 and 1.48 ± 0.12 ng/ml, respectively. The 20-oxo-P values of the pregnant animal, although slightly higher, were comparable to those of non-pregnant elephants during the luteal phase. Total immunoreactive 20-oxo-P values are about three times higher than those of progesterone during the luteal phase, and 5α-pregnane-3,20-dione is the major immunoreactive 20-oxo-P in the plasma of Asian and African elephants. Zoo Biol 16:403–413, 1997. © 1997 Wiley-Liss, Inc.     

Short and long phases of progesterone secretion during the oestrous cycle of the African elephant (Loxodonta africana)

Serum samples were collected from 3 mature female African elephants once each week for 15-18 months. Circulating concentrations of progesterone, oestradiol and LH were determined by radioimmunoassay (RIA). The LH RIA was validated by demonstrating parallel cross-reaction with partly purified elephant LH pituitary fractions. Changing serum progesterone concentrations indicated an oestrous cycle length of 13.3 +/- 1.3 weeks (n = 11). The presumed luteal phase, characterized by elevated serum progesterone values, was 9.1 +/- 1.1 weeks (n = 11). Two abbreviated phases of progesterone in serum lasting 2-3 weeks were observed in 2 elephants, indicating short luteal phases. Oestradiol concentrations in serum were variable, with no clear pattern of secretion. More frequent blood samples were collected during periovulatory periods and 9 distinct LH peaks were detected; all were followed by rises in serum progesterone concentrations. Periovulatory changes in progesterone and LH in sera correlated with external signs of oestrus and mating behaviour.     

Social suppression of ovarian cyclicity in captive and wild colonies of naked mole-rats, Heterocephalus glaber

To investigate the endocrine cause of reproductive suppression in nonbreeding female naked mole-rats, animals from 35 colonies were studied in captivity. Urinary and plasma progesterone concentrations were elevated in pregnant females (urine: 10.0-148.4 ng/mg Cr, 27 samples from 8 females; plasma: 3.6-30.0 ng/ml, 5 samples from 5 females; Days 21-40 of pregnancy) and cyclic breeding females (urine: 0.5-97.8 ng/mg Cr, 146 samples from 7 females; plasma: less than 1.0-35.4 ng/ml, 25 samples from 7 females). The latter group showed cyclic patterns of urinary progesterone, indicating a mean ovarian cycle length of 34.4 +/- 1.6 days (mean +/- s.e.m.) with a follicular phase of 6.0 +/- 0.6 days and a luteal phase of 27.5 +/- 1.3 days (19 cycles from 9 breeding females). In non-breeding females urinary and plasma progesterone values were undetectable (urine: less than 0.5 ng/mg Cr, 232 samples from 64 females; plasma: less than 1.0 ng/ml, 7 samples from 6 females). Breeding females had higher (P less than 0.001) plasma LH concentrations (3.0 +/- 0.2 mi.u./ml, 73 samples from 24 females) than did non-breeding females (1.6 +/- 0.1 mi.u./ml, 57 samples from 44 females). Urinary and plasma progesterone concentrations in non-breeding females from wild colonies situated near Mtito Andei, Kenya, were either below the assay sensitivity limit (urine: less than 0.5 ng/mg Cr, 11 females from 2 colonies; plasma: less than 1.0 ng/ml, 25 females from 4 colonies), or very low (plasma: 1.6 +/- 0.6 ng/ml, 15 females from 4 colonies). In captivity, non-breeding females removed from their colonies (i.e. the dominant breeding female) and either paired directly with a non-breeding male (N = 2), or removed and housed singly for 6 weeks before pairing with a non-breeding male (N = 5) may develop a perforate vagina for the first time in as little as 7 days. Urinary progesterone concentrations rose above 2.0 ng/mg Cr (indicative of a luteal phase) for the first time 8.0 +/- 1.9 days after being separated. These results suggest that ovulation is suppressed in subordinate non-breeding female naked mole-rats in captive and wild colonies, and show that plasma LH concentrations are significantly lower in these non-breeding females. This reproductive block in non-breeding females is readily reversible if the social factors suppressing reproduction are removed.     

Survey of the reproductive cyclicity status of Asian and African elephants in North America

The Asian and African elephant populations in North America are not self‐sustaining, and reproductive rates remain low. One problem identified from routine progestagen analyses is that some elephant females do not exhibit normal ovarian cycles. To better understand the extent of this problem, the Elephant TAG/SSP conducted a survey to determine the reproductive status of the captive population based on hormone and ultrasound evaluations. The survey response rates for facilities with Asian and African elephants were 81% and 71%, respectively, for the studbook populations, and nearly 100% for the SSP facilities. Of the elephants surveyed, 49% of Asian and 62% of African elephant females were being monitored for ovarian cyclicity via serum or urinary progestagen analyses on a weekly basis. Of these, 14% of Asian and 29% of African elephants either were not cycling at all or exhibited irregular cycles. For both species, ovarian inactivity was more prevalent in the older age categories (>30 years); however, acyclicity was found in all age groups of African elephants. Fewer elephant females (～30%) had been examined by transrectal ultrasound to assess reproductive‐tract integrity, and corresponding hormonal data were available for about three‐quarters of these females. Within this subset, most (～75%) cycling females had normal reproductive‐tract morphologies, whereas at least 70% of noncycling females exhibited some type of ovarian or uterine pathology. In summary, the survey results suggest that ovarian inactivity is a significant reproductive problem for elephants held in zoos, especially African elephants. To increase the fecundity of captive elephants, females should be bred at a young age, before reproductive pathologies occur. However, a significant number of older Asian elephants are still not being reproductively monitored. More significantly, many prime reproductive‐age (10–30 years) African females are not being monitored. This lack of information makes it difficult to determine what factors affect the reproductive health of elephants, and to develop mitigating treatments to reinitiate reproductive cyclicity. Zoo Biol 23:309–321, 2004. © 2004 Wiley‐Liss, Inc.     

Estrous synchrony in a group of African elephants (Loxodonta africana) under human care

Synchrony of estrous, and consequently of conception and birth of young, may be of adaptive significance for certain mammals. Among the species in which estrous synchrony has been suspected several times are elephants, but clear evidence is still missing. We determined estrous cycles of African elephants (Loxodonta africana) (n=4) at the Vienna Zoo, Austria, between June 2003 and January 2006 by measuring serum progesterone levels from weekly blood samples. Except for the dominant female when she was intensively lactating, all animals showed clear cycles or progesterone release with a mean period of 105.3+/-15.37 days. For most of the study period, estrous cycles were asynchronous between females. However, after re-occurrence of the progesterone cycle in the dominant female following the first period of lactation, all four females showed high synchrony of progesterone release over the two subsequent cycles. Large changes in individual period lengths indicated that synchronization was due to the adjustment of cycle length in subdominants to that of the dominant female. We used a bootstrap procedure, based on resampling measured times of progesterone peaks, to determine if this apparent synchrony could have been caused by chance alone. This statistical analysis indicated that between-individual variances of the timing of progesterone peaks were much smaller that to be expected by chance (P=0.009). This finding represents the first evidence for estrous synchrony between elephants. We discuss various hypotheses to explain the biological function of cycle synchrony in elephants.     

Fecal 20-oxo-pregnane concentrations in free-ranging African elephants (Loxodonta africana) treated with porcine zona pellucida vaccine

Because of overpopulation of African elephants in South Africa and the consequent threat to biodiversity, the need for a method of population control has become evident. In this regard, the potential use of the porcine zona pellucida (pZP) vaccine as an effective means for population control is explored. While potential effects of pZP treatment on social behavior of African elephants have been investigated, no examination of the influence of pZP vaccination on the endocrine correlates in treated females has been undertaken. In this study, ovarian activity of free-ranging, pZP-treated African elephant females was monitored noninvasively for 1 yr at Thornybush Private Nature Reserve, South Africa, by measuring fecal 5α-pregnan-3β-ol-20-on concentrations via enzyme immunoassay. A total of 719 fecal samples from 19 individuals were collected over the study period, averaging 38 samples collected per individual (minimum, maximum: 16, 52). Simultaneously, behavioral observations were made to record the occurrence of estrous behavior for comparison. Each elephant under study showed 5α-pregnan-3β-ol-20-on concentrations rising above baseline at some period during the study indicating luteal activity. Average 5α-pregnan-3β-ol-20-on concentrations were 1.61 ± 0.46 μg/g (mean ± SD). Within sampled females, 42.9% exhibited estrous cycles within the range reported for captive African elephants, 14.3% had irregular cycles, and 42.9% did not appear to be cycling. Average estrous cycle duration was 14.72 ± 0.85 wk. Estrous behavior coincided with the onset of the luteal phase and a subsequent rise in 5α-pregnan-3β-ol-20-on concentrations. Average 5α-pregnan-3β-ol-20-on levels positively correlated with rainfall. No association between average individual 5α-pregnan-3β-ol-20-on concentrations or cyclicity status with age or parity were detected. Earlier determination of efficacy was established via fecal hormone analysis with no pregnancies determined 22 mo post-treatment and onward. Results indicate the presence of ovarian activity amongst pZP-treated female African elephants in 2 yr after initial immunization. Further study should now be aimed toward investigating the long-term effects of pZP vaccination on the reproductive function of female African elephants.     

Seasonal effects on the endocrine pattern of semi-captive female Asian elephants (Elephas maximus): timing of the anovulatory luteinizing hormone surge determines the length of the estrous cycle

Better breeding strategies for captive Asian elephants in range countries are needed to increase populations; this requires a thorough understanding of their reproductive physiology and factors affecting ovarian activity. Weekly blood samples were collected for 3.9 years from 22 semi-captive female Asian elephants in Thai elephant camps to characterize LH and progestin patterns throughout the estrous cycle. The duration of the estrous cycle was 14.6+/-0.2 weeks (mean+/-S.E.M.; n=71), with follicular and luteal phases of 6.1+/-0.2 and 8.5+/-0.2 weeks, respectively. Season had no significant effect on the overall length of the estrous cycle. However, follicular and luteal phase lengths varied among seasons and were negatively correlated (r=-0.658; P<0.01). During the follicular phase, the interval between the decrease in progestin concentrations to baseline and the anovulatory LH (anLH) surge varied in duration (average 25.9+/-2.0 days, range 7-41, n=23), and was longer in the rainy season (33.4+/-1.8 days, n=10) than in both the winter (22.2+/-4.5 days, n=5; P<0.05) and summer (18.9+/-2.6 days, n=8; P<0.05). By contrast, the interval between the anLH and ovulatory LH (ovLH) surge was more consistent (19.0+/-0.1 days, range 18-20, n=14). Thus, seasonal variation in estrous cycle characteristics were mediated by endocrine events during the early follicular phase, specifically related to timing of the anLH surge. Overall reproductive hormone patterns in Thai camp elephants were not markedly different from those in western zoos. However, this study was the first to more closely examine how timing of the LH surges impacted estrous cycle length in Asian elephants. These findings, and the ability to monitor reproductive hormones in range countries (and potentially in the field), should improve breeding management of captive and semi-wild elephants.     

Hormone secretion in the asian elephant (Elephas maximus): characterization of ovulatory and anovulatory luteinizing hormone surges.

In the elephant, two distinct LH surges occur 3 wk apart during the nonluteal phase of the estrous cycle, but only the second surge (ovLH) induces ovulation. The function of the first, anovulatory surge (anLH) is unknown, nor is it clear what regulates the timing of these two surges. To further study this observation in the Asian elephant, serum concentrations of LH, FSH, progesterone, inhibin, estradiol, and prolactin were quantified throughout the estrous cycle to establish temporal hormonal relationships. To examine long-term dynamics of hormone secretion, analyses were conducted in weekly blood samples collected from 3 Asian elephants for up to 3 yr. To determine whether differences existed in secretory patterns between the anLH and ovLH surges, daily blood samples were analyzed from 21 nonluteal-phase periods from 7 Asian elephants. During the nonluteal phase, serum LH was elevated for 1-2 days during anLH and ovLH surges with no differences in peak concentration between the two surges. The anLH surge occurred 19.9+/-1.2 days after the end of the luteal phase and was followed by the ovLH surge 20.8+/-0.5 days later. Serum FSH concentrations were highest at the beginning of the nonluteal phase and gradually declined to nadir concentrations within 4 days of the ovLH surge. FSH remained low until after the ovLH surge and then increased during the luteal phase. Serum inhibin concentrations were negatively correlated with FSH during the nonluteal phase (r = -0.53). Concentrations of estradiol and prolactin fluctuated throughout the estrous cycle with no discernible patterns evident. In sum, there were no clear differences in associated hormone secretory patterns between the anLH and ovLH surge. However, elevated FSH at the beginning of the nonluteal phase may be important for follicle recruitment, with the first anLH surge acting to complete the follicle selection process before ovulation.     

Urinary progesterone in free-ranging red howler monkeys (Alouatta seniculus): preliminary observations of the estrous cycle and gestation

The goals of this study were to develop and validate a radioimmunoassay (RIA) for measurement of unconjugated progesterone (P) concentrations in the urine of red howler monkeys (Alouatta seniculus) and to use urinary P profiles to characterize the reproductive cycle of this species. Analysis of P profiles from two females provided a preliminary estimate of the length of the estrous cycle (mean days +/- S.E.M. = 29.5 +/- 1.5; n = 2), and indicated that one female red howler copulated throughout two apparent estrous cycles. Urinary P concentrations during two confirmed pregnancies (211.8 +/- 29.7 ng P/ml) were higher (P < 0.05) than during the luteal phase (77.4 +/- 10.6 ng P/ml; n = 4) of the cycle.     

Investigation of individual and group variability in estrous cycle characteristics in female Asian elephants (Elephas maximus) at the Oregon Zoo

Evaluating ovarian cycle activity through longitudinal progestagen monitoring is important for optimizing breeding management of captive elephants and understanding impact of life events (births, deaths, and transfers) on reproductive function. This study summarized serum progestagen profiles for eight Asian mainland elephants (Elephas maximus indicus) and one Bornean elephant (E. maximus borneensis) at the Oregon Zoo over a 20-yr interval, and represents the longest longitudinal dataset evaluated to date. Estrous cycle characteristics were more varied than previously reported for this species, with an overall duration of 12 to 19 wk, luteal phase duration of 4 to 15 wk, and follicular phase duration of 2 to 12 wk. In general, there was more cycle variability across than within individual elephants. Compared with other elephants in the group, the Borneo female exhibited consistently longer cycle lengths, higher progestagen concentrations, and greater cycle variability; however, it is not known if this represents a subspecies or an individual difference. Cycle durations did not appear to change over time or with age, and the first pubertal cycle was similar to subsequent cycles. Variability in duration of the follicular phase was greater than that of the luteal phase. In addition, there was a significant negative relationship between luteal and follicular phase durations, suggesting a possible regulatory role of the follicular phase in maintaining a relatively consistent cycle duration within individuals. Overall, we found these elephants to be highly resilient in that major life events (births, deaths, and changes in herd structure) had minimal effect on cycle dynamics over time. In conclusion, the higher range in cycle phase characteristics is likely because of the larger number of elephants studied and longer duration of longitudinal monitoring, and may be more representative of the captive population as a whole. Furthermore, identification of significant interanimal variability suggests that understanding the complexities of herd reproductive characteristics could facilitate development of more effective institution-specific breeding management strategies.     

Behavioral indices of estrus in a group of captive African elephants (Loxodonta africana)

This study investigated behavioral signals of estrus by systematically monitoring the interactions of one male with four female African elephants housed in a naturalistic outdoor enclosure at Disney's Animal Kingdom over a period of 11 months. We measured changes in five spatial behaviors and 22 tactile‐contact behaviors, as well as changes in serum progestagen and LH concentrations, across three ovarian cycles for each female. Two females did not cycle during the study. Three different phases of the ovarian cycle were identified: mid luteal, anovulatory follicular, ovulatory follicular. The male followed more and carried out more genital inspections, flehmen, and trunk‐to‐mouth behaviors toward cycling females during their ovulatory phase. Genital inspections by the male peaked above baseline levels on the day of an LH surge, and up to 9 days before, in both cycling females and, thus, might be a useful behavioral index of estrus. The male also carried out more genital inspections, flehmen, and trunk touches to the back leg toward ovulatory cycling than noncycling females. Overall, our results indicated that: 1) a single subadult African elephant male could discriminate two females in the ovulatory phase of their cycle (i.e., during the 3 weeks preceding ovulation) from the mid luteal phase; 2) the male also discriminated two cycling females in the ovulatory and anovulatory follicular phases from two noncycling females; 3) two females in the ovulatory phase of the cycle displayed a greater variety of tactile‐contact behavior toward the male compared to the other cycle phases. Zoo Biol 0:1–19, 2005. © 2005 Wiley‐Liss, Inc.     

Reproductive inhibition in the Cape porcupine, Hystrix africaeaustralis

Sexually mature female Cape porcupines kept under natural conditions of illumination and temperature did not conceive while housed within their natal groups. Before removal from their natal groups the sexually mature offspring copulated and experienced cyclic ovarian activity, but conception occurred only 70-120 days after dispersal. Mean oestrous cycle length of these females (36.9 +/- 11.5 days; n = 34) was similar to that of breeding females (33.0 +/- 11.64 days; n = 16), but mean peak plasma progesterone concentration (6.45 +/- 6.03 ng/ml; n = 34) was significantly (P less than 0.01) lower than that of cyclic breeding females (13.58 +/- 6.98 ng/ml; n = 16). Mean progesterone concentration at oestrus in non-breeding females (0.72 +/- 0.45 ng/ml; n = 34) was also significantly (P less than 0.01) lower than that of non-pregnant breeding females (4.21 +/- 2.44 ng/ml; n = 16). Reproductive inhibition within natal groups, in which only one female reproduces, therefore cannot be ascribed to a failure to copulate, but may be due to some factor inhibiting full expression of luteal activity or affecting ovulation.     

Results of the third reproductive assessment survey of North American Asian (Elephas maximus) and African (Loxodonta africana) female elephants

A written survey assessed reproductive status of female Asian and African elephants in AZA/SSP facilities in 2008, and data were compared to surveys conducted in 2002 and 2005. Results showed that ovarian acyclicity rates across the surveys remained unchanged for Asian (13.3, 10.9 and 11.1%) and African (22.1, 31.2 and 30.5%) elephants, respectively (P > 0.05), but were higher overall for African compared to Asian elephants (P < 0.05). In 2008, the percentages of Asian and African elephants with irregular cycles (14.3 and 15.8%) and irregular + no cycles (25.4 and 46.4%) was similar to 2005 (7.6 and 11.8%; 18.5 and 43.0%), but were increased compared to 2002 (2.6 and 5.2%; 16.0 and 27.3%), respectively (P < 0.05). For both species, ovarian acyclicity increased with age (P < 0.05). Reproductive tract pathologies did not account for the majority of acyclicity, although rates were higher in noncycling females (P < 0.05). Bull presence was associated with increased cyclicity rates (P < 0.05) for Asian (92.5 vs. 58.3%) and African (64.9 vs. 57.8%) elephants compared to females at facilities with no male, respectively. Cyclicity rates were higher for Asian (86.8 vs. 65.2%) and African (67.9 vs. 56.7%) elephants managed in free compared to protected contact programs (P < 0.05), respectively. Geographical facility location had no effect on cyclicity (P > 0.05). In summary, incidence of ovarian cycle problems continues to predominantly affect African elephants. Although percentages of acyclicity did not increase between 2005 and 2008, 42.2% Asian and 30.2% African females were no longer being hormonally monitored; thus, reproductive cycle abnormalities could be worse than current data suggest.     

Non-invasive assessment of the reproductive cycle in free-ranging female African elephants (Loxodonta africana) treated with a gonadotropin-releasing hormone (GnRH) vaccine for inducing anoestrus

ABSTRACT: BACKGROUND: In southern Africa, various options to manage elephant populations are being considered. Immunocontraception is considered to be the most ethically acceptable and logistically feasible method for control of smaller and confined populations. In this regard, the use of gonadotropin-releasing hormone (GnRH) vaccine has not been investigated in female elephants, although it has been reported to be safe and effective in several domestic and wildlife species. The aims of this study were to monitor the oestrous cycles of free-ranging African elephant cows using faecal progestagen metabolites and to evaluate the efficacy of a GnRH vaccine to induce anoestrus in treated cows. METHODS: Between May 2009 - June 2010, luteal activity of 12 elephant cows was monitored non-invasively using an enzyme immunoassay detecting faecal 5alpha-reduced pregnanes (faecal progestagen metabolites, FPM) on a private game reserve in South Africa. No bulls of breeding age were present on the reserve prior to and for the duration of the study. After a 3-month control period, 8 randomly-selected females were treated twice with 600 micrograms of GnRH vaccine (Improvac[REGISTERED SIGN], Pfizer Animal Health, Sandton, South Africa) 5-7 weeks apart. Four of these females had been treated previously with the porcine zona pellucida (pZP) vaccine for four years (2004-2007). RESULTS: All 12 monitored females (8 treated and 4 controls) showed signs of luteal activity as evidenced by FPM concentrations exceeding individual baseline values more than once. A total of 16 oestrous cycles could be identified in 8 cows with four of these within the 13 to 17 weeks range previously reported for captive African elephants. According to the FPM concentrations the GnRH vaccine was unable to induce anoestrus in the treated cows. Overall FPM levels in samples collected during the wet season (mean 4.03 micrograms/gram dry faeces) were significantly higher (P<0.002) than the dry season (mean 2.59 micrograms/gram dry faeces). CONCLUSIONS: The GnRH vaccination protocol failed to induce anoestrus in the treated female elephants. These results indicate that irregular oestrous cycles occur amongst free-ranging elephants and are not restricted to elephants in captivity. The relationship between ecological conditions and endocrine activity were confirmed. Free-ranging female elephants were observed to not cycle continuously throughout the year in the absence of adult bulls.     

Interrelationship of serum testosterone, dominance and ovarian cyclicity status in female African elephants

The African elephant population in North American zoos is not self-sustaining, in part due to the prevalence of ovarian acyclicity. While little is known about the cause of this condition, earlier research has shown that females without cyclic corpus luteum (CL) function rank higher in the dominance hierarchy than females with cyclic CL function. The goal of this study was to measure longitudinal serum testosterone concentrations in captive female African elephants to determine if there is a relationship among serum testosterone concentrations, social dominance rank and ovarian cyclicity status. Weekly blood samples from 49 female African elephants (24 having and 25 not having cyclic CL function at 22 facilities) were collected over a 12-month period and analyzed for serum testosterone using an enzymeimmunoassay. A progesterone radioimmunoassay was used to quantify serum progestagen concentrations and categorize ovarian cyclicity status. The dominance hierarchy of individual elephants within each herd was assessed by a written temperament survey, which identified 19 dominant, 15 middle and 15 subordinate females. No clear patterns of serum testosterone secretion were observed in females with and without cyclic CL function. Furthermore, no significant relationships were found among serum testosterone concentrations, dominance rank, and ovarian cyclicity status. These data suggest that increased circulating testosterone concentrations are not associated with greater rates of ovarian acyclicity or dominance status in captive female African elephants.     

Reproductive cycle of the elephant

The combination of a few factors, including poor captive reproduction, secession of importation from the wild and advances in hormone detection and ultrasonography, has contributed to the current knowledge on the elephant reproductive cycle. Several reproductive features in elephants differ markedly from other mammals. These include the urogenital tract anatomy, length and structure of the reproductive cycle, the formation of multiple corpora lutea and the type and secretion pattern of reproductive hormones. Being 13-18 weeks in length, the elephant estrous cycle is the longest amongst all studied non-seasonal mammals to date. Progesterone increases 1-3 days after ovulation, indicating the start of the luteal phase, which lasts 6-12 weeks. This is followed by a 4- to 6-week follicular phase that is concluded by two, precisely spaced and timed, LH surges. In general, the first, anovulatory LH surge occurs exactly 19-21 days before the second, ovulatory surge. Normally, a single follicle is ovulated. However, beside a corpus luteum (CL) forming on the site of ovulation, multiple accessory CLs can be found on the ovaries. Unlike many other species, the predominant progestagen secreted by luteal tissues is not progesterone, but rather its 5-alpha-reduced metabolites. The currently known aspects of the unique estrous cycle in Asian and African elephants, covering estrous behavior, circulating hormones, ultrasonography and anatomy of the reproductive organs as well as hormonal manipulation treatment possibilities, will be reviewed here.     

Secretory patterns of serum prolactin in Asian (Elephas maximus) and African (Loxodonta africana) elephants during different reproductive states: Comparison with concentrations in a noncycling African elephant

Serum prolactin was quantified in adult female Asian (Elephas maximus) and African (Loxodonta africana) elephants during various reproductive states and the profiles compared to that in a noncycling African elephant. In reproductively normal elephants, there was no effect of season, estrous cycle stage, or lactational status on quantitative or qualitative prolactin secretion (P > 0.05), nor where there any differences (P > 0.05) in overall prolactin concentrations between species. In pregnant elephants, prolactin concentrations remained at baseline for the first 4–6 months of gestation. Thereafter, concentrations during early pregnancy averaged ∼four-fold higher than those during the estrous cycle, increasing to ∼100-fold over baseline during mid- to late gestation in both species. In contrast to cycling elephants, prolactin concentrations in an African elephant exhibiting chronic anovulation (on the basis of an acyclic serum progesterone profile) and mild galactorrhea were consistently about five-fold higher (P < 0.05), suggesting she is hyperprolactinemic. Other endocrinological assesments confirmed the hypogonadal state of this female. Serum estradiol concentrations were consistently at or below detectable levels. Additionally, no preovulatory luteinizing hormone (LH) surges occurred in daily serum samples analyzed over a 12-month period. The pituitary was not totally refractory, however, and responded with a several-fold increase in serum LH concentration (peak, 3.07 ng/ml) over baseline (0.75 ng/ml) after i.v. injection of gonadotropin-releasing hormone. This study describes normal baseline serum prolactin values for Asian and African elephants and is the first to identify hyperprolactinemia as a possible cause of reproductive acyclicity and galactorrhea in an African elephant. Zoo Biol 16:149–159, 1997. © 1997 Wiley-Liss, Inc.     

Oestrous cycles and the breeding season of the Père David's deer hind (Elaphurus davidianus)

Reproductive cycles were studied in a group of tame Père David's deer hinds. The non-pregnant hind is seasonally polyoestrous and, in animals studied over 2 years, the breeding season began in early August (2 August +/- 3.3 days; s.e.m., N = 9) and ended in mid-December (18 December +/- 5.7 days; N = 8) and early January (6 January +/- 3.2 days; N = 11) in consecutive years. During the anoestrous period, plasma progesterone concentrations were low (0.2 +/- 0.01 ng/ml) or non-detectable. There was a small, transient increase in progesterone values before the onset of the first cycle of the breeding season. In daily samples taken during an oestrous cycle in which hinds were mated by a marked vasectomized stag, progesterone concentrations remained low (less than 0.5 ng/ml) for a period of about 6 days around the time of oestrus, showed a significant increase above oestrous levels by Day 4 (Day 0 = day of oestrus) and then continued to increase for 18 +/- 2.8 days to reach mean maximum luteal levels of 3.5 +/- 0.6 ng/ml. The plasma progesterone profiles from a number of animals indicated that marking of the hinds by the vasectomized stag did not occur at each ovulation during the breeding season and therefore an estimate of the cycle length could not be determined by this method. In the following year, detection of oestrus in 5 hinds was based on behavioural observations made in the absence of the stag. A total of 19 oestrous cycles with a mean length of 19.5 +/- 0.6 days was observed.(ABSTRACT TRUNCATED AT 250 WORDS)