LH responses to single doses of exogenous GnRH in the Cape mole rat (Georychus capensis): the pituitary potential for opportunistic breeding

The Cape mole rat Georychus capensis is a solitary mole rat that inhabits the winter rainfall region of the Western Cape Province of South Africa. Circulating basal concentrations of luteinizing hormone (LH) were found to be significantly higher in the breeding season in both sexes. During both the breeding and non-breeding season, administration of exogenous gonadotropin-releasing hormone (GnRH) increased circulatory LH levels. The magnitude of the LH response to an overdose of exogenous GnRH both in and out of the breeding season in males and females was not significantly different. Typically, seasonally breeding species exhibit a down-regulation of the pituitary and reproductive functions out of the breeding season. It appears that there is no down-regulation of GnRH receptors at the level of the pituitary out of the breeding season, because the pituitary responds to an exogenous GnRH challenge equally both in and out of the breeding period. The Cape mole rat exhibits the potential for opportunistic breeding out of the breeding period, provided that environmental factors are favourable. This finding questions whether this mole rat is actually a seasonal breeder or whether reproduction is hindered by the ecological constraint of the lack of opportunities to burrow and find mates at certain times of the year.     

Ovarian growth and folliculogenesis in breeding and nonbreeding females of a social rodent,the Zambian common mole-rat,Cryptomys sp.

Zambian common mole-rats are subterranean rodents that live in families with only one female breeding. Her offspring remain in the parental nest and do not reproduce. Behavioral experiments (Burda, '95) demonstrated that their apparent “sterility” is based on incest avoidance and individual recognition of family members. To elucidate whether some kind of morphologically apparent ovarian suppression still takes place in daughters, ovaries of females of known age, weight, and reproductive histories were examined histologically and morphometrically. The body mass of old females (more than 3 years of age) begins to decrease, and the ovaries seem to begin to atrophy at the age of about 3–6 years. Ovaries in neonates exhibited primordial and primary follicles, sometimes clustered in nests. Ovaries of adult nonbreeding females expressed all stages of the follicular development up to tertiary follicles. Many unruptured luteinized follicles were present, but true corpora lutea as a morphological sign of ovulation were missing. Unruptured luteinized follicles also could be found (additionally to true corpora lutea) in ovaries of breeding females. The number of primordial follicles dropped rapidly during the first 2 years of age; the number of primary, secondary, and tertiary follicles was subject to individual variation; and there was no clear correlation with age or reproductive status. While a tendency to form accessory unruptured luteinized follicles may just reflect taxonomic affinities of bathyergids to hystricomorphs, the otherwise complete folliculogenesis in “sterile” daughters and the presence of unruptured luteinized follicles even in breeding females are further evidence that there is no hormonal suppression of the ovarial cycle. We suggest that ovulation in nonbreeding females is not actively suppressed by the breeding female, but instead that it is not released because the triggering mechanisms, most probably repeated copulation, are missing. J. Morphol. 237:33–41, 1998. © 1998 Wiley-Liss, Inc.     

LH responses to single doses of exogenous GnRH by social Mashona mole-rats: a continuum of socially induced infertility in the family Bathyergidae.

The Mashona mole-rat, Cryptomys darlingi, exhibits an extreme reproductive division of labour. Reproduction in the colony is restricted to a single breeding pair. The non-reproductive male and female colony members are restrained from sexual activity by being familiar and related to one another and the reproductive animals. Circulating basal concentrations of luteinizing hormone (LH) as well as LH levels measured in response to a single exogenous gonadotropin releasing hormone (GnRH) challenge are not significantly different between the reproductive and non-reproductive groups of either sex. Socially induced infertility in both non-reproductive males and females does not result from a reduced pituitary secretion of LH or decreased sensitivity to hypothalamic GnRH, but rather appears to result from an inhibition of reproductive behaviour in these obligate outbreeders. The African mole-rats exhibit a continuum of socially induced infertility with differing social species inhabiting regions of varying degrees of aridity. In this continuum a transition from a predominantly behavioural repression in a social mesic-adapted species through to complete physiological suppression lacking incest avoidance in an arid-adapted eusocial species occurs in this endemic African family of rodents.     

Effect of nutrition on the LH response to calf removal and GnRH

Ten primiparous crossbred cows were assigned to two dietary groups at calving. One group received 120% and the other group received 80% of the National Research Council (NRC) recommended allowance of dietary energy for primiparous cows. At 60 days postpartum, calves were removed from their dams. Blood samples were collected from the cows at 15-min intervals for 8 hr beginning at the time of calf removal and again 24 hr, 48 hr and 72 hr after calf removal. At 72 hr after calf removal, all cows were given 200 ug GnRH intravenously. At calf removal, serum LH concentrations were higher (P<0.01) for cows on 120% (0.9 +/- 0.03 ng/ml) compared to cows on 80% (0.5 +/- 0.03 ng/ml) of recommendations. Serum LH concentrations increased (1.6 +/- 0.1 ng/ml, P<0.01) by 24 hr in cows on the highenergy diet. In contrast, a similar increase was not observed in cows on the low-energy diet until 48 hr after calf removal (1.4 +/- 0.2 ng/ml, P<0.01). These contrasting patterns in serum LH concentrations resulted in a diet by time interaction (P<0.01). Serum LH concentrations increased in both dietary energy groups following GnRH injection, but the response was greater (P<0.01) in cows on the low-energy diet compared to the cows fed the high-energy diet. These results indicate that inadequate dietary energy delays the LH response to calf removal and increases the LH response to exogenous GnRH.     

LH responses of female naked mole-rats, Heterocephalus glaber, to single and multiple doses of exogenous GnRH

To investigate possible differential pituitary secretion of LH in breeding and non-breeding female naked mole-rats, the LH responses to administration of exogenous GnRH were measured in 55 females from 20 captive colonies. Single doses of 0.1, 0.5 or 1.0 micrograms GnRH produced a significant rise in plasma LH concentrations 20 min after s.c. injection in breeding and non-breeding females at all doses (P less than 0.001). While at the highest dose of 1.0 microgram there was no difference in the LH response between breeding and non-breeding females, as the dose was lowered there was a progressive decline in the LH response in non-breeding females such that, at the 0.1 microgram dose, GnRH produced only a small, but significant, increase in plasma LH (1.3 +/- 0.2 to 2.9 +/- 0.5 mi.u./ml, N = 5) compared with breeding females (3.4 +/- 0.8 to 9.6 +/- 2.0 mi.u./ml, N = 6). The LH responses of the latter were not significantly reduced at the lower doses of GnRH. The apparent lack of sensitivity to low doses of exogenous GnRH in non-breeding females was reversed by 4 consecutive 1-h injections of 0.1 microgram, which produced a rise in LH from 1.2 +/- 0.2 to 9.0 +/- 0.2 mi.u./ml (N = 4), comparable to that of breeding females given a single injection of 0.1 microgram GnRH. These results suggest that the anterior pituitary in non-breeding female naked mole-rats is less sensitive to low doses of exogenous GnRH than in breeding females, possibly due to a lack of priming by endogenous GnRH. Therefore, the socially-induced block to ovulation in non-breeding female naked mole-rats may be due to inhibition of hypothalamic GnRH secretion.     

Continuous infusion of GnRH reduces the LH response to an intravenous GnRH injection but does not inhibit endogenous LH secretion in cows

Plasma LH concentrations were monitored in 6 Hereford X Friesian suckled cows at about 80 days post partum, before and during a 14-day period of continuous s.c. infusion of GnRH (20 micrograms/h). Blood samples were collected at 10-min intervals on Days -2, -1, 1, 2, 3, 4, 7, 10, 13 and 14 (Day 1 = start of infusion). Plasma LH concentrations rose from mean pretreatment levels of 1.3 +/- 0.20 ng/ml to a maximum of 17.1 +/- 3.09 ng/ml within the first 8 h of GnRH infusion, but returned to pretreatment levels by Day 2 or 3. In 4/6 animals, the initial increase was of a magnitude characteristic of the preovulatory LH surge. In all animals, an i.v. injection of 10 micrograms GnRH, given before the start and again on the 14th day of continuous infusion, induced an increase in LH concentrations but the increase to the second injection was significantly (P less than 0.01) less (mean max. conc. 6.4 +/- 0.76 and 2.3 +/- 0.19 ng/ml). Mean LH concentrations (1.0 +/- 0.08, 1.1 +/- 0.08 and 0.9 +/- 0.06 ng/ml) and LH episode frequencies (3.3,4.3 and 3.2 episodes/6 h) did not differ significantly on Days -2,7 and 13. However, the mean amplitude of LH episodes was significantly lower (P less than 0.05) on Day 13 (1.3 +/- 0.10 ng/ml) than on Day -2 (1.8 +/- 0.16 ng/ml). Therefore, although the elevation in plasma LH concentrations that occurs in response to continuous administration of GnRH is short-lived and LH levels return to pre-infusion values within 48 h of the start of infusion, these results show that the pituitary is still capable of responding to exogenous GnRH, although the LH response to an i.v. bolus injection of GnRH is reduced. In addition, this change in pituitary sensitivity is not fully reflected in endogenous patterns of episodic LH secretion.     

Validation of radioimmunoassays for LH and FSH in the sooty mangabey (Cercocebus atys): Characterization of LH and the response to GnRH

Heterologous radioimmunoassays (RIA) for macaque LH and FSH were validated for the measurement of these hormones in the sooty mangabey and mangabey pituitary LH was characterized relative to rhesus monkey LH. Dilutions of a pituitary mangabey extract and a partially purified preparation of mangabey LH ran parallel to a rhesus monkey standard (LER 1909-2) in the ovine-ovine (o-o) LH assay but showed some deviation from parallelism in the rhesus monkey FSH assay. The LH potency of the mangabey extract and standard were six and 190 times more potent, respectively, than LER 1909-2 in the LH RIA. Mangabey LH was estimated to have a molecular weight of 40,000–42,000 daltons vs 35,000–38,000 daltons for rhesus LH on Sephadex G-100 chromatography. Plasma levels of radioimmunoreactive LH, FSH, and testosterone were assayed before and after a bolus administration of 25, 50, or 100 μg synthetic go-nadotropin releasing hormone (GnRH) to adult male mangabeys. A significant increase in serum levels of LH was seen within 30 min with levels more than fourfold higher than the basal level of LH after administration of 100 μg GnRH. However, no consistent increases in plasma FSH values were detected. The integrated mean LH response above preinjection levels following 25, 50, or 100 μg GnRH was dose related. Serum levels of testosterone were also elevated after administration of GnRH, but peak concentrations of testosterone lagged behind peak levels of LH by approximately 30 min. These studies indicate that the heterologous RIAs may be used for measuring gonadotropins in the mangabey and that the male mangabey is apparently more sensitive to GnRH than the rhesus monkey.     

LH response to GnRH infusion in postpartum, fall-lambing ewes subjected to prepartum energy restrictions

Fall-lambing western range ewes were fed either a high-or low-energy ration the last two months of gestation and were fed protein and energy in excess postpartum. GnRH was infused for 10 hours on day 5 or 26 postpartum. Blood samples were collected at 20-minute intervals during infusion and radioimmunoassayed for LH. Net weight change from day 60 prepartum to day 1 postpartum was 0.67 +/- 1 kg vs. -6.9 +/- 1.5 kg (mean +/- SE) for the high-and low-energy groups, respectively (P < 0.05). Nutritional treatment had no effect on LH response at either day 5 or day 26 postpartum, althoughthe day 26 LH response to GnRH was greater (P < 0.05) than the day 5 LH response. LH response was greater (P < 0.05) in ewes that gained weight prepartum vs. ewes that lost weight prepartum, but only in ewes infused on day 5 postpartum.     

Pituitary luteinizing hormone responses to single doses of exogenous GnRH in female social Cape ground squirrels exhibiting low reproductive skew

The Cape ground squirrel Xerus inauris is unusual among social mammals as it exhibits a low reproductive skew, being a facultative plural breeder with not all females breeding within a group. We investigated pituitary function to assess whether there was reproductive inhibition at the level of the pituitary and potentially the hypothalamus in breeding and non-breeding female Cape ground squirrels. We did so during the summer and winter periods by measuring luteinizing hormone (LH) responses to single doses of 2 g exogenous gonadotropin-releasing hormone (GnRH) and physiological saline administered to 42 females from 11 colonies. Basal LH concentrations of females increased in response to the GnRH challenge. Basal plasma LH concentrations were greater during winter, when most oestrus events are observed. However, we found no differences in plasma LH concentrations between breeding and non-breeding females. We showed that the anterior pituitary of non-breeding female ground squirrels is no less sensitive to exogenously administered GnRH than that of breeding females. We therefore concluded that the pituitary is no more active in breeding than non-breeding females. The lack of differentiation in response to GnRH suggests that either non-breeding females have ovaries that are less sensitive to LH or that they refrain from sexual activity with males through an alternative mechanism of self-restraint.     

A mathematical model for LH release in response to continuous and pulsatile exposure of gonadotrophs to GnRH

In a previous study, a model was developed to investigate the release of luteinizing hormone (LH) from pituitary cells in response to a short pulse of gonadotropin-releasing hormone (GnRH). The model included: binding of GnRH to its receptor (R), dimerization and internalization of the hormone receptor complex, interaction with a G protein, production of inositol 1,4,5-trisphosphate (IP₃), release of calcium from the endoplasmic reticulum (ER), entrance of calcium into the cytosol via voltage gated membrane channels, pumping of calcium out of the cytosol via membrane and ER pumps, and release of LH. The extended model, presented in this paper, also includes the following physiologically important phenomena: desensitization of calcium channels; internalization of the dimerized receptors and recycling of some of the internalized receptors; an increase in G _q concentration near the plasma membrane in response to receptor dimerization; and basal rates of synthesis and degradation of the receptors. With suitable choices of the parameters, good agreement with a variety of experimental data of the LH release pattern in response to pulses of various durations, repetition rates, and concentrations of GnRH were obtained. The mathematical model allows us to assess the effects of internalization and desensitization on the shapes and time courses of LH response curves.     

Use of a GnRH agonist to prevent the endogenous LH surge and injection of exogenous LH to induce ovulation in heifers superstimulated with FSH: a new model for superovulation

A new protocol for superovulating cattle which allows for control of the timing of ovulation after superstimulation with FSH was developed. The preovulatory LH surge was blocked with the GnRH agonist deslorelin, and ovulation was induced by injection of LH. In Experiment 1, heifers (3-yr-old) were assigned to a control group (Group 1A, n = 4) or a group with deslorelin implants (Group 1B, n = 5). On Day -7, heifers in Group 1A received a progestagen CIDR-B((R))device, while heifers in Group 1B received a CIDR-B((R))device + deslorelin implants. Both groups were superstimulated with twice daily injections of FSH (Folltropin((R))-V): Day 0, 40 mg (80 mg total dose on Day 0); Day 1, 30 mg; Day 2, 20 mg; Day 3, 10 mg. On Day 2, heifers were given PGF (a.m.) and CIDR-B((R)) devices were removed (p.m.). Three heifers in Group 1A had a LH surge and ovulated, whereas neither of these events occurred in Group 1B (with deslorelin implants) heifers. In Experiment 2, heifers (3-yr-old) were assigned to 1 of 4 equal groups (n = 6). On Day -7, heifers in Group 2A received a norgestomet implant, while heifers in Groups 2B, 2C and 2D received norgestomet + deslorelin implants. Heifers were superstimulated with FSH starting on Day 0 as in Experiment 1. On Day 2, heifers were given PGF (a.m.) and norgestomet implants were removed (p.m.). Heifers in Groups 2B to 2D were given 25 mg LH (Lutropin((R))): Group 2B, Day 4 (a.m.); Group 2C, Day 4 (p.m.); Group 2D, Day 5 (a.m.). Heifers in Group 2A were inseminated at estrus and 12 and 24 h later, while heifers in Groups 2B to 2D were inseminated at the time of respective LH injection and 12 and 24 h later. Injection of LH induced ovulation in heifers in Groups 2B to 2D. Heifers in Group 2C had similar total ova and embryos (15.2 +/- 1.4) as heifers in Group 2A (11.0 +/- 2.8) but greater (P < 0.05) numbers than heifers in Group 2B (7.0 +/- 2.3) and Group 2D (6.3 +/- 2.0). The number of transferable embryos was similar for heifers in Group 2A (5.8 +/- 1.8) and Group 2C (7.3 +/- 2.1) but lower (P < 0.05) for heifers in Group 2B (1.2 +/- 0.8) and Group 2D (1.3 +/- 1.0). The new GnRH agonist-LH protocol does not require observation of estrus, and induces ovulation in superstimulated heifers that would not have an endogenous LH surge.     

Gonadotropin response to GnRH during sexual ontogeny in the common carp, Cyprinus carpio

This study was designed to reveal whether gonadotropic response to GnRH in the common carp (Cyprinus carpio) changes during sexual ontogeny and whether the response of FSHbeta and LHbeta subunits is uniform or differential. The study comprised fish at the following stages: juveniles (4-month-old females with primary oocytes and early spermatogenic males); maturing (9-month-old previtellogenic females and advanced spermatogenic males); and mature (16-month-old postvitellogenic females and spermiating males). Fish were injected with superactive salmon GnRH analogue (sGnRHa; 25 microg/kg) and blood was sampled 6, 12 and 24 h later for cGtH (LH) and sex steroid levels. Pituitaries were taken for determination of FSHbeta and LHbeta mRNA levels by slot-blot hybridization and for cGTH content in the same glands by radioimmunoassay (RIA). Values were compared with the levels prior to sGnRHa administration and with control fish sampled at the same intervals. Juvenile fish did not respond at all to sGnRHa. In maturing females, FSHbeta mRNA increased by >300%, while that of LHbeta increased by 200%. In maturing males, FSHbeta mRNA did not change and only a slight increase occurred in that of LHbeta. In 16-month-old postvitellogenic females, there was no response of FSHbeta mRNA, while that of LHbeta dramatically increased. In spermiating males of the same age, mRNA of both FSHbeta and LHbeta increased following sGnRHa injection. Immunoreactive cGtH was present in the pituitary and plasma of all fish examined, but in juveniles it did not change following sGnRHa injection. In maturing and mature fish of both genders, sGnRHa administration was followed by a marked increase in circulating cGtH, concomitant with a decrease in its pituitary content, indicating the limited amount of the hormone stored in the gland. In conclusion, the response of the gonadotropin subunit mRNAs in the common carp was found to be differential and dependent on the gender and the phase of sexual ontogeny.     

Pollination and reproductive biology of twelve species of neotropical Malpighiaceae: stigma morphology and its implications for the breeding system

BACKGROUND AND AIMS: This study on reproductive biology examines the stigmatic morphology of 12 Brazilian Malpighiaceae species with regard to their pollination and breeding system. METHODS: The species were studied in natural populations of a semi-deciduous forest fragment. Style tips were processed for observation by SEM and pollen-tube growth was analyzed under fluorescence microscopy. The breeding system was investigated by isolating flowers within waterproof bags. Floral visitors were recorded through notes and photographs. KEY RESULTS: Flowers are yellow, pink or white, protogynous, herkogamous and sometimes lack oil glands. While Banisteriopsis pubipetala has functional female flowers (with indehiscent anthers), 11 species present hermaphrodite flowers. Stigmas of these species may be terminal, with a slightly concave surface, or internal, consisting of a circular cavity with a large orifice, and are covered with a thin, impermeable cuticle that prevents pollen from adhering, hydrating, or germinating. Malpighiaceae have a special type of 'wet' stigma, where a secretion accumulates under the cuticle and is released by mechanical means-mainly rupture by pollinators. Even though six species show a certain degree of self-compatibility, four of them present a form of late-acting self-incompatibility, and the individual of B. pubipetala is agamospermous. Species of Centris, Epicharis and Monoeca bees pollinate these flowers, mainly collecting oil. Some Epicharis and Monoeca species collected pollen by vibration. Paratetrapedia and Tetrapedia bees are pollen and oil thieves. CONCLUSIONS: The Malpiguiaceae species studied are pollinator-dependent, as spontaneous self-pollination is limited by herkogamy, protogyny and the stigmatic cuticle. Both the oil- and pollen-collecting behaviours of the pollinators favour the rupture of the stigmatic cuticle and the deposition of pollen on or inside the stigmas. As fruit-set rates in natural conditions are low, population fragmentation may have limited the sexual reproduction of these species.     

The social structure and reproductive biology of the common mole-rat, Cryptomys h. hottentotus and remarks on the trends in reproduction and sociality in the family Bathyergidae

Cryptomys h. hottentotus occurs in mesic and semi-arid regions of South Africa. It lives in colonies ofup to 14 individuals, in which reproduction is limited to one ofthe largest males and the largest female in the colony.
Reproduction and details of colony size and the number of breeding animals in a colony are described for one complete and two incomplete field-captured colonies. One incomplete colony was kept in captivity for more than 3 years, whereas the complete colony was autopsied.
The social hierarchy of the common mole-rat is almost linear and has a value of 0.7 1 calculated from Landau's linearity index. Subordinate mole-rats weighed less than dominant mole-rats. The reproductive mole-rats are the dominant animals within the colony. The non-reproductive females rank lowest in the hierarchy.
Mating was not confined to a particular season, but occurred more frequently during the summer months. The alpha male initiated pre-copulatory behaviour. The gestation length is 55 to 66 days (n = 2). Two litters each of three pups were born to the colony during captivity. The pups were relatively altricial. They wandered out of the nest 5 days after birth, began to eat solids when 10 days old, and were fully weaned at 4 weeks. When 2 weeks old the pups began to spar with each other and with some of the adult colony members, but levels of aggression were never high. The pups did not disperse but were incorporated into the colony, this suggesting that C. h. hottentotus colonies are largely composed of related individuals.     

Effect of betamethasone treatment on luteal lifespan and the LH response to GnRH in dairy cows

Betamethasone (a synthetic glucocorticoid, 15 mg) was administered i.m. twice daily for 10 days to 4 regularly cycling dairy cows, beginning on Day 10 of the oestrous cycle. Luteal function, monitored by plasma progesterone, was extended by 7, 9, 19 and 20 days, respectively. Luteal function in the next cycle was normal. Endogenous cortisol values were suppressed for 14, 13, 34 and 27 days, respectively. Pituitary responsiveness to 20 micrograms GnRH was assessed by LH measurement on Days -1, +3 and +7 relative to the start of betamethasone treatment. There was a progressive decrease in peak LH concentrations after each GnRH challenge compared to control cows. Hourly measurements of PGF-2 alpha metabolite during the expected period of luteolysis failed to reveal normal increases. It is suggested that betamethasone caused prolonged luteal function, either by directly inhibiting PGF-2 alpha release, or by suppressing pituitary stimulation of follicular growth and hence lowering oestradiol concentrations, since it is known that PGF-2 alpha and oestradiol act synergistically to cause luteolysis.     

Relationship of age to seasonal levels of LH, FSH, prolactin and testosterone in male, white-tailed deer

Seasonal levels of LH, FSH, Prolactin (PRL) and testosterone (T) were determined in blood plasma of penned male white-tailed deer, ranging in age from 2 to 10 yr. Peak levels of T (observed around the rutting season in November) gradually increased until the 7th yr and then they began to decline slowly; a sharp decrease was registered in the 10th yr. Peak levels of PRL (measured in June) steadily increased until the 6th yr and then dropped rapidly in the 8-9-yr-old group. Peak concentrations of FSH (observed during September-October) rose gradually until the 6th yr, decreased in 8-yr-olds and then increased again in the 9th and 10th yr of life. On the other hand LH maxima (occurring during July-September) were rising until the 4th yr and then remained steady until the 6th yr. LH peaks in the 8th and 9th yr were more than 50% higher than that of the 4-6th yr. These data indicate that increasing peak levels coincide with approaching "prime male age" around 5-7 yr. In senior bucks (9-10 yr) decreasing gonadal function may be the sign of diminished responsiveness to pituitary hormones since gonadotrophins are elevated to the "castrate-type" levels.