Seasonal changes in the reproductive organs and plasma and pituitary hormone content of the male Bennett's wallaby (Macropus rufogriseus vufogriseus)

In a number of species of seasonally breeding marsupial, the male is fertile throughout the year but there is a marked seasonal change in weight of the accessory sexual glands. In this study, body weight, prostate, epididymis and testis weights and plasma concentrations of testosterone, LH and prolactin and pituitary content of LH and prolactin were determined in male Bennett's wallabies shot at 1–2 month intervals over a period of 17 months. There was a highly significant increase in prostate weight which was coincident with the breeding season for this species. A small but significant increase in testis weight was also observed but epididymis weight remained unchanged. Plasma testosterone concentrations were significantly increased at a time coincident with the increase in prostate weight. Plasma prolactin and LH concentrations were low in most animals and remained unchanged during the study. In contrast, pituitary prolactin and LH contents showed highly significant changes, with increasing and peak hormone content preceding maximum prostate weight and plasma testosterone concentrations by several months. While these latter results suggest a role for prolactin and LH in the seasonal control of the reproductive organs in the male wallaby, a more intensive study of the pattern of secretion of these hormones and possibly more sensitive hormone assays are required to understand their relative roles in regulating the annual cycle of prostate growth.     

Reproductive endocrinology of the Wandering Albatross Diomedea exulans in relation to biennial breeding and deferred sexual maturity

The reproductive endocrinology of the Wandering Albatross Diomedea exulans was studied at South Georgia to investigate the potential endocrine correlates of biennial breeding and of the acquisition of sexual maturity. Gonads of breeding birds and of known-age immature birds of both sexes were examined by laparoscopy throughout the period that they were at the nest site. Blood samples, subsequently analysed to determine concentrations of luteinizing hormone (LH), prolactin, progesterone, testosterone and oestradiol-17/i, were obtained from samples of breeding birds of both sexes at regular intervals from first arrival until the chicks fledged nearly a year later. Before laying in December, breeding birds had mature testes and ovarian follicles and high concentrations of LH, prolactin and sex steroids. Gonadal regression and a rapid drop in hormone levels (except for LH in females) occurred in early incubation (January). Testes (and follicles to a lesser extent) enlarged in mid-incubation, coinciding with high levels of LH and increases in prolactin and testosterone. Gonads finally regressed completely near hatching time. LH, prolactin and testosterone remained at low levels throughout chick rearing (April to November), but females had several periods of active progesterone and oestradiol secretion, and progesterone was detectable in males only late in the chick-rearing period. Although some changes in hormone levels are difficult to explain, the patterns are fairly typical of temperate birds. The persistence of progesterone secretion in both female breeders and non-breeding ‘immature’ birds is viewed as part of a mechanism inhibiting an ovary from becoming vitellogenic. Although testis size and testosterone concentrations increased with age in immature males (of ages 4–10 years), birds of 5 years and older are probably physiologically mature, even though breeding does not start until they are 7 years of age and only half an age group has bred by an age of 11 years. Immature females (of age 4–7 years) had undeveloped follicles, very low oestradiol concentrations but high progesterone levels, providing further support for the role of this hormone in inhibiting gonadotropin secretion. The condition of the female is therefore probably decisive in determining when a pair first attempts to breed but it is unknown what factors initiate normal ovarian development.     

Seasonal variations of LH, prolactin, androstenedione, testosterone and testicular FSH binding in the male blue fox (Alopex lagopus)

The seasonal changes in testicular weight in the blue fox were associated with considerable variations in plasma concentrations of LH, prolactin, androstenedione and testosterone and in FSH-binding capacity of the testis. An increase in LH secretion and a 5-fold increase in FSH-binding capacity were observed during December and January, as testis weight increased rapidly. LH levels fell during March when testicular weight was maximal. Plasma androgen concentrations reached their peak values in the second half of March (androstenedione: 0.9 +/- 0.1 ng/ml: testosterone: 3.6 +/- 0.6 ng/ml). A small temporary increase in LH was seen in May and June after the breeding season as testicular weight declined rapidly before levels returned to the basal state (0.5-7 ng/ml) that lasted until December. There were clear seasonal variations in the androgenic response of the testis to LH challenge. Plasma prolactin concentrations (2-3 ng/ml) were basal from August until the end of March when levels rose steadily to reach peak values (up to 13 ng/ml) in May and June just before maximum daylength and temperature. The circannual variations in plasma prolactin after castration were indistinguishable from those in intact animals, but LH concentrations were higher than normal for at least 1 year after castration.     

Short days induce premature reproductive maturation in juvenile starlings, Sturnus vulgaris

Castration of juvenile and photorefractory adult starlings caused no immediate increase in circulating concentrations of LH. In castrated juveniles and adults exposed to natural changes in daylength, plasma LH increased between mid-October and mid-November, although the increase was more rapid in adults. In castrated photorefractory adults, plasma LH increased 3-5 weeks after transfer to artificial short days (8L:16D). In castrated juvenile starlings plasma LH increased 4-6 weeks after transfer to 8L:16D, irrespective of the age of the birds. Birds as young as 17 weeks had high LH concentrations. These results suggest that the reproductive system of juvenile starlings is in the same state as that of photorefractory adults, and therefore the activation of the hypothalamo-pituitary axis for the first time in juveniles is analagous to the termination of photorefractoriness in adults.     

Modulation of gonadotrophin and prolacrin secretion by daylength and breeding behaviour in free-living starlings, Sturnus vulgaris

Plasma levels of luteinizing hormone (LH), follicle-stimulating hormone (FSH) and prolactin were measured by radioimmunoassay in plasma samples collected from free-living starlings, Sturnus vulgaris , trapped in nest-boxes. By leaving some nest-boxes undisturbed, and repeatedly destroying nests in others, birds from a single-brooded population were trapped whilst nest-building, incubating or feeding nestlings, at different times throughout the normal breeding season. In both males and females trapped whilst nest-building, plasma LH and prolactin levels increased progressively from mid March until late May. In females sampled during incubation, LH and FSH levels were high throughout May but decreased in early June. Prolactin levels were highest in late May. In both males and females trapped during mid May, LH levels were highest in these birds which were nest-building at this time and lowest in those feeding nestlings, FSH did not change significantly, and prolactin was low in those birds which were nest-building and high in those incubating or feeding nestlings. In female starlings from a double-brooded population, levels of LH and FSH were much lower whilst feeding the second brood than when feeding the first brood, whereas prolactin levels were similar. The results suggest that incubation and feeding behaviour inhibit the increase in LH secretion caused by increasing daylength, but stimulate prolactin secretion in excess of that caused by increasing daylength.     

Effect of cyclosporin treatment on luteinizing hormone and prolactin.

The effect of cyclosporin A (CsA) treatment on LH and prolactin was investigated. Chronic daily administration of an immunosuppressive dose of CsA (1.5 mg/100g bw) increased serum LH concentrations and pituitary gland LH content. CsA treatment also resulted in increased serum testosterone. Immunosuppressive doses of CsA had no effect on serum prolactin or pituitary gland prolactin content. Acute administration of low doses of 0.12, 1.2, 12 and 120ug CsA/100g bw had no effect on serum LH or prolactin. These results suggest that administration of immunosuppressive doses of CsA alters serum and pituitary LH and serum testosterone but not prolactin.     

Maturation of the pituitary-gonadal system in the male rat

Serum FSH and testosterone concentrations reached maximum levels between 35 and 45 days of age, which coincided with the appearance of mature spermatozoa in the majority of seminiferous tubules. Spermatozoa were not observed in sections of the urethra until the age of 46 days. Serum LH concentrations were low (5-6 ng/ml) before Day 25, became highly variable (12-57 ng/ml) between Days 25 and 53 and remained consistently above 35 ng/ml thereafter. Serum prolactin levels rose significantly between 30 and 43 days of age. Maximum prolactin levels coincided with the start of accelerated growth in the prostate and seminal vesicle glands. Testicular weights relative to body weight reached a plateau by 35 days of age, while relative pituitary and adrenal weights decreased throughout the study period. It is suggested that spermatogenesis is not complete until FSH and testosterone reach maximum levels, while prolactin may be involved in the stimulation of accessory sex organ growth. The pronounced variation in serum LH concentrations during the maturation period may reflect a progressive change in the sensitivity of the hypothalamic-pituitary axis to the negative feedback of gonadal steroids.     

Endocrine correlates of parental care in an Antarctic winter breeding seabird, the emperor penguin, Aptenodytes forsteri

Plasma levels of luteinizing hormone (LH) and prolactin associated with parental behavior were measured in the Antarctic winter breeding emperor penguin, Aptenodytes forsteri. Males exclusively incubate the egg while females exclusively brood the nonhomeothermic young; both sexes alternate in rearing the homeothermic young. Birds were sampled on arrival from the sea through egg laying, incubation, and brooding. All parent birds lost their chicks at the end of the brooding period due to harsh weather but sampling continued. In females, LH titers dropped after egg laying but levels were restored when the birds returned from the sea to brood the chicks and were not depressed by high prolactin levels. Plasma prolactin remained low in males captured on arrival and kept until the free-living males finished incubation. In breeders, prolactin secretion increased during the prelaying period when day length decreased. Prolactin levels stayed elevated in males during incubation and in brooding females returning after a 2-month absence at sea. Prolactin values were higher in brooding females than in males ending incubation or returning in late brooding. These levels did not drop after chick loss, and the sexual difference in prolactin values was maintained after breeding failure. In emperor penguins, increased prolactin secretion appears to be triggered around the time of egg laying and continues, driven by an endogenous mechanism, through incubation and brooding until rearing is completed. Prolactin secretion independent of external stimuli may have evolved in pelagic seabirds to maintain parental care despite long absences at sea from the breeding colony.     

Patterns of breeding behaviour and plasma levels of hormones in a free-living population of Pied flycatchers, Ficedula hypoleuca

Plasma levels of luteinizing hormone (LH) and steroid hormones were measured by radioimmunoassay in a series of plasma samples collected from a free-living population of Pied flycatchers, Ficedula hypoleuca. Both males and females have similar circulating levels of LH and testosterone when they arrive in the breeding area. However, as territories and pair bonds are established, and as courtship and nest-building ensue, the plasma levels of testosterone in males become elevated over those of females, although a lesser peak also occurs in females. As the egg laying phase and incubation begin, plasma levels of LH and testosterone decline sharply in both sexes. On the other hand polyterritorial males, that establish secondary territories, maintain higher levels of LH and testosterone than males remaining in the home-territory, at least until they return to the home-territory, usually at the middle part of the incubation period. By this time polyterritorial males also have basal levels of these hormones. Curiously, females on home-territories show a small, but significant, rise of circulating LH and testosterone at the time the males return.
In males, plasma levels of corticosterone are elevated from the time of arrival on the breeding grounds until the end of the nest-building period. Thereafter corticosterone levels decline in males staying on the home-territory but remain elevated in polyterritorial males until the latter return to the home-territory. In contrast, circulating levels of corticosterone in females are only elevated for brief period during egg laying.     

Effects of removal of chicks from hens on concentrations of prolactin, luteinizing hormone and oestradiol in plasma of brooding Gifujidori hens.

Gifujidori hens were allowed to repeat a breeding cycle in one season. In the first breeding cycle the duration of the brooding (raising chicks) stage was limited to 3 weeks, whereas in the second breeding cycle it was limited to 1 week by removing all chicks from mother hens. In the first breeding cycle, plasma prolactin (PRL) was high during the incubation period, but rapidly decreased on the day of hatching and reached minimum values about 1 week after hatching. In contrast, plasma luteinizing hormone (LH) concentrations were low during the incubation period, but after hatching they gradually increased and reached peak values immediately after removal of chicks. Concentrations of oestradiol in plasma were low in the incubation and brooding stages but increased significantly immediately after removal of chicks. In the second breeding cycle, changes in PRL and LH concentrations were similar to those observed in the first breeding cycle except that even greater increases in plasma LH and oestradiol concentrations were observed one week after hatching when the chicks were removed. These results suggest that coexistence of newly hatched chicks may suppress LH secretion from the pituitary of the hen in the natural breeding cycle.     

Reproductive endocrinology of free living Pied flycatchers (Ficedula hypoleuca): prolactin and FSH secretion in relation to incubation and clutch size

Plasma levels of prolactin and FSH in free-living Pied flycatchers were studied in relation to breeding stage, clutch size and diurnal variations. In the female, the concentration of prolactin started to increase during the egg-laying and reached maximal values at the end of the incubation period. After hatching prolactin levels decreased. Male Pied flycatchers showed an increase in plasma levels of prolactin at the time when they returned to their home-territories, i.e. at the end of the incubation period. Males still occupying secondary territories at this time had low levels of prolactin. In the males, as in the females, plasma levels of prolactin decreased after hatching.
Male Pied flycatchers showed at all times higher plasma levels of FSH than did females. In both sexes FSH levels were high during the early part of the breeding season and started to decrease after the egg-laying period. After hatching time plasma levels of FSH were below the sensitivity of the assay. No differences in FSH were found between males occupying secondary territories and males in home-territories.
In incubating females FSH, but not prolactin, showed a distinct unimodal daily cycle.
No differences were found in plasma levels of prolactin and FSH between females incubating different sized clutches.     

Pituitary luteinizing hormone and prolactin messenger ribonucleic acid levels are inversely related in laying and incubating turkey hens.

The relationships of prolactin (PRL) and LH messenger (m) RNA to serum and pituitary content were determined for turkey hens at different phases of the reproductive cycle. In the nonphotostimulated, reproductively inactive hen, serum and pituitary PRL content and pituitary PRL mRNA levels were low. All three PRL values rose after photostimulation and peaked during the incubation phase. Relative to nonphotostimulated hens, hyperprolactinemic incubating hens showed 220-, 11-, and 57-fold increases in serum PRL, pituitary PRL content, and pituitary PRL mRNA levels, respectively. These peak levels declined 80-, 3-, and 6-fold, respectively, in photorefractory hens. In contrast to PRL levels, serum LH, pituitary LH, and pituitary LH beta-subunit mRNA levels did not change as dramatically. Serum LH showed no significant changes for the different reproductive phases. Pituitary LH peaked after photostimulation and declined to its lowest level in incubating hens. Pituitary LH-beta mRNA abundance was highest in photostimulated and laying hens and lowest in incubating and photorefractory hens. These results demonstrate that the abundance of LH-beta and PRL mRNA shows an inverse relationship in photostimulated/laying and incubating turkey hens.     

Quantification of prolactin messenger ribonucleic acid, pituitary content and plasma levels of prolactin, and detection of immunoreactive isoforms of prolactin in pituitaries from turkey embryos during ontogeny.

The content of prolactin mRNA as well as total prolactin content and type of isoforms of prolactin were measured in single pituitary glands from turkey embryos and poults. Levels of mRNA and pituitary content of prolactin remained low until 5 days before hatching, while plasma concentrations remained low until 2 days before hatching. Levels of prolactin mRNA then increased until the day of hatch, stayed stable during the 3 first days of age, and significantly increased until 2 wk of age. Similar changes were observed in pituitary content and plasma levels of prolactin. Two immunoreactive bands of apparent molecular masses of 24 and 27 kDa, corresponding to the nonglycosylated and glycosylated form of prolactin, respectively, were visualized on Western blots. In pituitary glands from embryos at 22 days of incubation, 31.5% of the protein was glycosylated, whereas in embryos at 27 days of incubation and poults at 1 and 7 days of age, 48.6%, 48.0%, and 56. 0% of prolactin was glycosylated, respectively. The results indicate that the increases in the synthesis and the release of prolactin occur mainly around and after the time of hatching in the turkey embryo. Higher percentages of glycosylated isoforms were associated with increasing levels of total prolactin in the pituitary gland. Thus, the synthesis of prolactin and its post-translational modifications may be important factors involved in the physiologic changes occurring around the time of hatching.     

Effect of neonatal testosterone and oestradiol treatment on the development of the hypothalamo-hypophysial axis in the female rat.

The hypothalamic LH-RH content and the concentrations of pituitary and plasma LH were measured at various ages in female rats treated daily with 10 micrograms testosterone propionate or 10 micrograms oestradiol-17beta from birth to Day 15. Persistent vaginal oestrus was induced in all the treated rats. Both hormones significantly reduced the hypothalamic LH-RH content and pituitary and plasma LH concentrations. Hypothalamic LH-RH increased after cessation of treatment but pituitary LH did not return to normal levels. Plasma LH levels were significantly lower than those in control rats. It is concluded that testosterone propionate and oestradiol-17beta (1) have a direct negative feed-back influence on the hypothalamus in the neonatal female rat; (2) alter the normal pattern of plasma and pituitary LH in developing female rats; (3) prevent the cyclic secretion of plasma LH after maturity; and (4) probably cause a chronic impairment in the release of LH-RH.     

Changes in plasma gonadotrophin and prolactin concentrations following castration of the pony stallion

Concentrations of gonadotrophins and prolactin were recorded in pony stallions castrated during the early breeding season, to examine the regulatory role of the gonad at a time when testosterone has been postulated to exert positive feedback on LH secretion. Further, gonadotrophin concentrations in geldings are reported to return to values within the normal range of the entire stallion. In an attempt to characterize this species-specific reversal, the gonadotrophin concentrations of 6 male ponies castrated on 25 March were monitored for 4 months, and 4 stallions were used to generate control data. Blood samples were collected daily, from 3 d before to 10 d after castration (Day 0), and weekly thereafter until Day 122. The pituitary response to castration was immediate. Castration resulted in a previously unreported, dramatic (13-fold) but transient (3 d) surge in circulating concentrations of LH. Concentrations of LH and FSH increased in a logarithmically scaled (LH, R2 = 0.77; FSH, R2 = 0.93) manner over the subsequent 5 wk, during which temporal changes in concentrations of both hormones were strongly correlated (R2 = 0.97). The ratio of plasma gonadotrophin concentrations was consistent throughout (LH:FSH, 1.43 +/- 0.04). Maximal concentrations of LH (20.58 +/- 1.97 ng/mL, Day 34.8 +/- 3.2) were attained approximately 2 wk before the peak in FSH (16.99 +/- 1.97 ng/mL, Day 49.7 +/- 3.0). Plasma gonadotrophin concentrations exceeded those of entire stallions throughout the study. The equine testes inhibited LH secretion during the early breeding season, and no chronic decrease in plasma gonadotrophin concentrations was recorded. However, the LH surge evident for 3 d immediately afer castration, may be related to the dynamic seasonal interaction between gonadal steroids and the regulation of pituitary gonadotrophin release.     

Thyroidectomy results in termination of photorefractoriness in starlings (Sturnus vulgaris) kept in long daylengths

Four groups of 10 male starlings were transferred from short daylengths (8 h light/day) to long daylengths (18 h light/day), which caused the tests to develop rapidly to maximum size and then to decrease to minimal size as birds became photorefractory. Birds were surgically thyroidectomized at 8, 16 or 28 weeks. A fourth group was left intact. Testicular volume and plasma FSH and prolactin concentrations were measured. After 42 weeks all birds were castrated and plasma FSH was measured during the next 6 weeks. Testicular growth began in all thyroidectomized birds between 4 and 8 weeks after thyroidectomy. By 42 weeks, the testes of all thyroidectomized birds were large, whereas those of intact birds were still of minimal size. Plasma FSH concentrations remained low in all birds and plasma prolactin values, originally elevated by long daylengths, decreased at a similar rate in thyroidectomized and intact birds. After castration at 42 weeks, plasma FSH values increased rapidly in all thyroidectomized birds but remained low in non-thyroidectomized birds. The results demonstrate that thyroidectomy of photorefractory starlings does not induce immediate testicular growth but may initiate a process which eventually terminates photorefractoriness in a way similar to that caused by return to short daylengths.     

Plasma prolactin, thyroxine, triiodothyronine, testosterone, and luteinizing hormone during a photoinduced reproductive cycle in mallard drakes

The temporal relationships between plasma concentrations of prolactin, thyroxine (T4) and triiodothyronine (T3) were determined in a group of six wild mallard drakes during the development and maintenance of long-day refractoriness after transfer from 6 h light: 18 h darkness (6L:18D) to 20L:4D for 24 weeks. As shown by changes in the plasma concentrations of luteinizing hormone (LH) and testosterone, the birds came into breeding condition and then became long-day refractory within 5 weeks of photostimulation. Long-day refractoriness was maintained for the remainder of the study. Plasma prolactin began to increase immediately after photostimulation, although not as fast as the increases in plasma LH and testosterone. The concentration of plasma T4 also increased after photostimulation but, as shown by decreased plasma LH and testosterone levels, only after the birds had become long-day refractory. The development of long-day refractoriness was thus directly correlated with an increased plasma prolactin and not with a change in plasma concentration of T4. Plasma T3 decreased after photostimulation but returned to prestimulation values as the birds became long-day refractory and remained stable for the remainder of the study. Concentrations of plasma T4 and prolactin returned to baseline values after about 15 weeks photostimulation showing that the long-term maintenance of long-day refractoriness is not directly related to continuously high plasma concentrations of either hormone.     

Thyroxine treatment prevents the development of photosensitivity in european starlings (Sturnus vulgaris)

Summary In starlings, the breeding season is terminated by a state of photorefractoriness. Birds remain completely reproductively inactive as long as long days are maintained, and only exposure to short days restores photosensitivity. Two experiments investigated the role of different doses of thyroxine in the development of photosensitivity in castrated starlings. First, photorefractory castrated male starlings were moved from long (18L:6D) to short (8L:16D) days, and received in the drinking water either 1 or 10 mg · 1^-1 thyroxine for the first 7 weeks of a 14-week observation period. Control birds regained photosensitivity after 5 weeks of short days, as signaled by a spontaneous increase in plasma LH, whereas the return to photosensitivity was delayed until weeks 7 and 9 in the 1- and 10-mg · 1^-1 thyroxine-treated birds, respectively. In the second experiment, the effect of different doses of thyroxine was explored at the level of the hypothalamic Gn-RH neurosecretory neurones. The acquisition of photosensitivity in control birds transferred from long to short days was characterized by a marked increase in hypothalamic Gn-RH content (while long-day controls maintained low Gn-RH content). Doses of 10 and 20 mg · 1^-1 of thyroxine completely prevented the return to photosensitivity, as seen through changes in either plasma LH concentrations or hypothalamic Gn-RH content, while a dose of 1 mg · 1^-1 allowed a partial recovery of photosensitivity, as hypothalamic Gn-RH content increased to an intermediate level and the spontaneous rise in plasma LH occurred slowly but steadily.Abbreviations Gn-RH gonadotrophin-releasing hormone - LH luteinizing hormone - LHRH-I luteinizing hormone releasing hormone     

Prolactin and LH release in response to LH-RH and TRH in ewes during dioestrus, pregnancy and post partum

With advancing pregnancy in the ewe there was a marked decline in plasma LH concentrations and pituitary LH-RH responsiveness (integrated LH release) and a marked increase in plasma prolactin values and pituitary TRH responsiveness (integrated prolactin release). In lactating ewes plasma LH levels and pituitary LH-RH responsiveness had returned to values found in the luteal phase of the normal cycle by 21 days post partum, whereas at 42 days post partum prolactin levels were still high. No interaction between TRH and LH-RH on prolactin and LH release in dioestrous ewes was detected. In non-pregnant ewes plasma prolactin levels were significantly higher in June than in January but TRH responsiveness was similar. It is concluded that, in sheep, pituitary LH secretion recovers more rapidly from the chronic negative feedback effect of oestrogens and progesterone in pregnancy than prolactin secretion recovers from the chronic positive feedback effects of oestrogens. This finding may be a contributory factor in the resistance to resumption of breeding activity.     

Effect of daylength on the rate of recovery of photosensitivity in male starlings (Sturnus vulgaris).

Four groups of castrated photorefractory starlings were transferred from a photoperiod of 18 h light/day (18 L) to photoperiods of 6 L, 8 L, 11 L or 12 L. A control group was kept on 18 L. Plasma concentrations of luteinizing hormone (LH) were low in all groups initially. The first significant increase in LH, which signals the recovery of photosensitivity, occurred after 4 weeks in the groups on 6 L and 8 L, after 8 weeks in the 11 L group and after 13 weeks in the 12 L group. There was no increase in the group on 18 L. The rate of recovery of photosensitivity is therefore inversely proportional to daylength, for daylengths between 8 L and 12 L.