Hypothalamic monoamine turnover in ring doves (streptopelia risoria), courting,incubating and brooding young

1. The turnovers of hypothalamic 5-hydroxytryptamine (5HT), dopamine (DA) and noradrenaline (NE) were measured in male and female ring doves (Streptopelia risoria) at three stages of the breeding cycle: courtship, 3 days after pairing; early incubation, 1–2 days after egg laying; and brooding, 1–3 days after the squabs had hatched.2. In both sexes plasma LH decreased progressively from courtship through incubation to brooding young. Crop sacs were fully developed in doves brooding young but not at other stages of the reproductive cycle, indicating increased concentrations of plasma prolactin.3. The turnovers of 5HT and DA in both sexes were significantly higher (P<0.001) in doves brooding young than in birds incubating eggs or nest building. The turnover of DA was higher in females than in males at the onset of incubation. The turnover of NE was lower (P > 0.01) in females at the onset of incubation than during courtship or brooding.4. Increased turnover of hypothalamic DA may be more closely related to brooding behaviour than to changes in prolactin or LH secretion.5. Increased hypothalamic 5HT turnover in brooding doves appears to be more directly related to crop sac development, and by inference increased prolactin secretion, than to depressed plasma LH concentrations.     

Concentrations of plasma prolactin and luteinizing hormone following nest deprivation and renesting in ring doves (Streptopelia risoria)

Ring doves with increased plasma prolactin and low plasma LH (Group A) or with low plasma prolactin and low plasma LH (Group B) which had been incubating sterile eggs for 12 or 18 days, respectively, had their nests and eggs removed for 3 days. Upon nest return, observations were made on the birds' readiness to renest and on changes in plasma prolactin and LH. Birds from Group A demonstrated a far greater tendency to resume incubation than birds from Group B. Nest deprivation resulted in a sharp fall in the concentration of plasma prolactin in birds which were deprived of their nests after 12 days of incubation (Group A). Following resumption of incubation no subsequent increase in the prolactin levels was observed in Group A or B. The concentration of plasma LH rose sharply after nest deprivation in both sexes of both groups and declined after return of the nests. Birds in Groups A and B which returned to their nests laid a new clutch of eggs while continuing to incubate. The total length of uninterrupted sitting following nest return was 20.9 +/- 0.48 days (n = 8). These results suggest that (1) once the mechanism responsible for the increase in plasma prolactin during incubation is disrupted, it cannot be reactivated unless the whole reproductive cycle is repeated. (2) The inhibition of LH secretion during incubation involves neural mechanisms which do not necessarily involve the anti-gonadotrophic action of prolactin.     

Prolactin and parental behavior in Adélie penguins: effects of absence from nest, incubation length, and nest failure

Adélie penguin (Pygoscelis adeliae) males and females, nesting in Antarctica, alternate attendance at the nest with absences of many days to forage at sea. We investigated the importance of tactile input from egg and chicks on prolactin levels by observing nest attendance patterns and obtaining blood samples (1) during the first nest exchange of the incubation stage, (2) from birds whose incubation period was artificially increased or decreased by about 10 days, and (3) from birds whose nests had failed. Prolactin levels in females after 8 to 11 days of absence from the breeding colony did not differ from those in incubating males and did not change after females resumed incubation. Moving eggs between nests resulted in nests in which chicks hatched after about 26, 36 (normal), or 46 days. Duration of incubation did not affect prolactin levels in the parents measured during incubation, at the pip stage, hatch stage, or early brood stage. Adults first left their chicks unguarded on about the same calendar date, regardless of chick age. However, chicks from long incubation nests averaged 8 days younger when they were left unguarded than chicks from control or short-incubation nests. In females, there was no effect of nest failure on prolactin levels. In males, prolactin levels were slightly lower after nest failure than in males tending nests. Testosterone was significantly higher in males after nest failure than in males still tending nests. Prolactin is elevated in Adélie penguins as part of the program of cyclical hormonal changes that accompany the lengthy reproductive season and is relatively independent of tactile input. Sustained prolactin secretion is probably required for the maintenance of parental behavior in offshore feeding species that must be absent from the nest for many days at a time.     

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.     

Plasma prolactin concentrations in breeding pied flycatchers (Ficedula hypoleuca) with an experimentally prolonged brooding period

This experiment explored the stimulatory effect of brooding newly hatched young on plasma prolactin concentration in male and female pied flycatchers, Ficedula hypoleuca. By exchanging offspring between nestboxes, one group of parents was exposed to 1- to 3-day-old nestlings for 12 days. Females in this group continued night-time brooding during these 12 days. The females, but not the males (who do not participate in brooding nestlings), after 6 days had higher plasma prolactin concentrations than control birds which had shown a 50% reduction in night-time brooding at this stage. By Day 12, however, prolactin concentration had decreased to the same level as in control birds, even though these females were still brooding 3-day-old nestlings. Female flycatchers given 3-day-old nestlings on the day their own eggs hatched showed an earlier reduction in night-time brooding and an earlier decrease in plasma prolactin than did control birds. An early decrease in prolactin was also seen in the males of this experimental group.     

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.     

Environmental regulation of the reproductive system in a flexibly breeding Sonoran Desert bird, the Rufous-winged Sparrow, Aimophila carpalis

We investigated reproductive regulation in male Rufous-winged Sparrows, Aimophila carpalis, a Sonoran Desert passerine that breeds after irregular summer rains. Field and captive data demonstrate that increased photoperiod stimulates testicular development in March and maintains it until early September. Free-living birds caught in July and placed on captive long days (16L: 8D) maintained developed testes for up to 7 months, and free-living birds caught in September, during testicular regression, redeveloped testes when placed on captive long days, indicating that these birds were still photosensitive. Captive birds on long days maintained testicular development when exposed to temperatures mimicking those occurring during regression in free-living birds. In free-living birds, testicular development was observed during spring and summer, but unless this was associated with rainfall, breeding (indicated by juveniles) did not occur. Large increases in plasma luteinizing hormone (LH) in free-living males were correlated with heavy rainfall in July/August, when the birds bred, and in November, when they did not breed. In captive birds, plasma LH concentrations were unresponsive to photoperiodic changes, but may have responded to social cues. Plasma prolactin concentrations were directly correlated with photoperiod in free-living birds, but an effect of photoperiod on prolactin secretion was not seen in captive birds. It is concluded that male Rufous-winged Sparrows use long photoperiods to stimulate and maintain testicular development, but exposure to long photoperiods does not terminate breeding by inducing absolute photorefractoriness. The specific timing of reproductive behaviors is apparently determined by elevated plasma LH coinciding with long day stimulated gonad development.     

Passive immunization against chicken vasoactive intestinal polypeptide suppresses plasma prolactin and crop sac development in incubating ring doves

Passive immunization of incubating ring doves with daily injections of sheep anti-chicken vasoactive intestinal polypeptide (cVIP) serum prevented the proliferation of crop sac tissue observed in control doves given nonimmune serum. Daily injections of anti-cVIP serum did not prevent crop sac development in nonbreeding doves simultaneously treated with ovine prolactin. The concentrations of plasma prolactin were significantly depressed in birds given anti-cVIP serum although this effect became progressively less pronounced during the course of the 7- or 14-day treatment periods. Body weights and weights of regressed reproductive organs were unaffected by treatment with anti-cVIP serum and did not differ significantly from control birds. Doves showing a decreased prolactin response to anti-cVIP serum treatment developed an immune response to sheep serum which may have immunoneutralized the administered antibody. Concentrations of plasma LH were not consistently affected by anti-cVIP serum administration and were low throughout the study. The depression in plasma LH normally seen in females after their young hatch was not observed in females treated with anti-cVIP serum. No effect of treatment was observed upon the birds' incubation behavior or in their readiness to feed and brood their young. These results suggest that in the ring dove, VIP is the physiological prolactin-releasing factor responsible for stimulating prolactin secretion and consequently the development of the crop sac, during incubation. They further indicate that increased concentrations of plasma prolactin may not be essential for gonadal regression or the maintenance of full incubation and brooding behavior in ring doves under laboratory conditions.     

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.     

External stimuli affecting incubation behavior and prolactin secretion in the duck (Anas platyrhynchos)

The importance of general environmental, including visual and tactile, stimuli on behavior and prolactin secretion during the incubation phase of reproduction in the duck (Anas platyrhynchos) was investigated. Nest occupancy rapidly increased at the end of egg laying and marked the initiation of incubation. Two recesses from the nest each day were synchronized to dawn and dusk; the median occurrence was 0.23 hr after dawn and 1.17 hr before dusk. Mean recess length was 36.1 +/- 1.9 min at dawn and 40.5 +/- 2.1 min at dusk. Plasma prolactin concentrations during incubation, 25.8 +/- 2.3 ng/ml, decreased to baseline levels, 10.8 +/- 1.9 ng/ml, within 24 hr after nestbox removal. The withdrawal of tactile, but not visual, stimuli of the clutch during incubation by either anesthesia or denervation of the incubation patch caused significant decreases in prolactin plasma concentrations within 24 hr. Prolactin plasma concentrations decreased rapidly at the end of incubation in ducks which successfully hatched young as well as in unsuccessful incubators. Temperature manipulations of the clutch, either above or below normal, caused decreases in plasma prolactin concentrations in parallel with temperature modification.     

Double nesting of the Red-legged Partridge Alectoris rufa

Some Red-legged Partridge females lay two clutches in separate nests, one immediately after the other, which are incubated separately by the male and female of the pair. Pairs remained together during the laying of both clutches so that there were delays between the end of laving and incubation at the first nests. Incubation of the two nests began at approximately the same time but discrepancies of up to ten days occurred. Males incubated first clutches and females usually the second, but probably the first if the second was destroyed during laying. The proportion of surviving eggs which hatched was similar in first and second clutches but declined if the delay between laying and hatching was exceptionally long.
Yearling females began laying late and few seemed to attempt two clutches compared with older females of which 60–80% showed the double nesting habit. A model predicting reproductive success for both sexes in relation to the rate of nest predation during laying, suggested that attempting two clutches rather than one would be disadvantageous at high predation rates. Females would produce more young if their mates incubated their first clutch immediately it was complete rather than accompanying them during the laying of the second. However, males may benefit by this delay, even though it exposes the nest to predators for a longer time, because they are able to guard their mate and prevent other males from mating with her and fertilizing eggs of her second clutch.     

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.     

鸽乳的生物学功能及其生成调控

鸽(Columba livia)是少数几种能分泌营养液哺育雏鸟的鸟类之一。孵化期的亲鸽嗉囊壁逐渐增厚,当雏鸽被孵出,亲鸽嗉囊会产生鸽乳(crop milk)以哺育雏鸽。鸽乳的营养成分及其生物学功能与哺乳动物的乳汁相似,其产生过程受催乳素的调节。在催乳素作用下,嗉囊上皮细胞快速增殖脱落形成鸽乳,该过程可能与膜联蛋白Icp35(AnxIcp35)等关键基因的转录以及JAK/STAT和Wnt等信号通路的激活有关。本文对鸽乳的主要成分、生物学功能和泌乳过程中嗉囊组织学变化进行了介绍,对鸽乳生成过程中特异的基因变化和分子调控机制进行了总结,以期为后续的相关研究工作提供有益的参考。     

Nest size affects clutch size and the start of incubation in magpies: an experimental study

Nest size has been suggested to be a sexually selected traitindicating parental ability of both males and females. To testwhether a female's reproductive decisions (e.g., clutch sizeand starting incubation) change in relation to experimentalmanipulation of nest size, as would be predicted if nest sizeis a sexually selected signal reflecting the male's parental quality, we manipulated nest size in a population of monogamousmagpies before laying by adding or removing about 20 cm oflarge sticks in the roof of magpie nests. On the one hand,we found that clutch size of reduced nests was smaller thanthat of control or enlarged nests. Moreover, clutch size was significantly related to nest size after manipulation, whichindicates that females adjust clutch size to the final sizeof the nest, nest size thereby being a good candidate for asexually selected trait. On the other hand, number of eggshatched during the first day is hypothesized to be related to the expected available resources during nestling growth, andsubsequent nestlings hatched are likely to die due to broodreduction if resources are not sufficient to raise well-developednestlings. Nest size is hypothesized to inform females abouta male's willingness to invest in reproduction, and we foundthat in broods of experimentally reduced nests, females startedto incubate earlier in the laying sequence than they did inbroods of control or enlarged nests. Moreover, in experimentallyreduced nests, fewer nestlings hatched during the first day,and the difference in body mass between the first and the fourthnestling hatched increased. This result is in accordance withthe hypothesis that the female's decision of when to start incubationin the laying sequence is mediated by nest size, a sexuallyselected trait signaling parental quality. We discuss alternativeexplanations for the results such as the possibility that nestsof different treatments may differ in their thermoregulationproperties or in their protection against predators.     

Effects of hyperprolactinemia on the control of luteinizing hormone and follicle-stimulating hormone secretion in the male rat

Experiments were conducted to determine the effects of acute hyperprolactinemia (hyperPRL) on the control of luteinizing hormone and follicle-stimulating hormone secretion in male rats. Exposure to elevated levels of prolactin from the time of castration (1 mg ovine prolactin 2 X daily) greatly attenuated the post-castration rise in LH observed 3 days after castration. By 7 days after castration, LH concentrations in the prolactin-treated animals approached the levels observed in control animals. HyperPRL had no effect on the postcastration rise in FSH. Pituitary responsiveness to gonadotropin hormone-releasing hormone (GnRH), as assessed by LH responses to an i.v. bolus of 25 ng GnRH, was only minimally effected by hperPRL at 3 and 7 days postcastration. LH responses were similar at all time points after GnRH in control and prolactin-treated animals, except for the peak LH responses, which were significantly smaller in the prolactin-treated animals. The effects of hyperPRL were examined further by exposing hemipituitaries in vitro from male rats to 6-min pulses of GnRH (5 ng/ml) every 30 min for 4 h. HyperPRL had no effect on basal LH release in vitro, on GnRH-stimulated LH release, or on pituitary LH concentrations in hemipituitaries from animals that were intact, 3 days postcastration, or 7 days postcastration. However, net GnRH-stimulated release of FSH was significantly higher by pituitaries from hyperprolactinemic, castrated males. To assess indirectly the effects of hyperPRL on GnRH release, males were subjected to electrical stimulation of the arcuate nucleus/median eminence (ARC/ME) 3 days postcastration. The presence of elevated levels of prolactin not only suppressed basal LH secretion but reduced the LH responses to electrical stimulation by 50% when compared to the LH responses in control castrated males. These results suggest that acute hyperPRL suppresses LH secretion but not FSH secretion. Although pituitary responsiveness is somewhat attenuated in hyperprolactinemic males, as assessed in vivo, it is normal when pituitaries are exposed to adequate amounts of GnRH in vitro. Thus, the effects of hyperPRL on pituitary responsiveness appear to be minimal, especially if the pituitary is exposed to an adequate GnRH stimulus. The suppression of basal LH secretion in vivo most likely reflects inadequate endogenous GnRH secretion. The greatly reduced LH responses after electrical stimulation in hyperprolactinemic males exposed to prolactin suggest further that hyperPRL suppresses GnRH secretion.     

Mate-sharing costs in polygynous Northern Lapwings Vanellus vanellus

In this study bigamous female Northern Lapwings Vanellus vanellus received significantly less incubation relief from their males than monogamous females. On average, monogamous males spent 34.3% of their time incubating and bigamous males 29.9%. Bigamous males divided their effort between their nests, incubating on average 9.4% on primary nests and 20.5% on secondary nests. Bigamous females compensated for the lack of male relief. Primary females incubated for 71.8% of their time, secondary females for 64.2%, while monogamous females spent 52.7% of their time incubating. As a result, there was no significant difference in total nest attentiveness among nests of different status. Primary and secondary females received equivalent incubation relief from the male. Bigamous males increased their contribution to incubation significantly as the season progressed. A bigamous male's distribution of incubation relief between his females was unrelated to female body mass, or to the degree of asynchrony between primary and secondary females in arrival and laying. Incubation time was significantly, negatively, correlated with total nest attentiveness. Monogamous females spent most time, secondary females spent an intermediate time, and primary females spent the least time on maintenance behaviour (foraging, comfort behaviour, inactivity). No significant differences were found in hatching success among females of different mating status. However, the ratio of unhatched to hatched eggs (i.e. the eggs that remained in the nest at the time of hatching) differed significantly: secondary females hatched a smaller proportion of their eggs than monogamous and primary females.     

Plasma levels of prolactin and gonadal steroids in relation to multiple-brooding and renesting in free-living populations of the song sparrow, Melospiza melodia

Plasma levels of testosterone (T), estradiol (E2), and prolactin were measured in samples collected from free-living song sparrows, Melospiza melodia. In males, plasma levels of T were elevated early in the season when territories were established and when females laid the first clutch of eggs. Thereafter, T levels declined and remained low throughout the remainder of the breeding cycle. However, if the first brood was lost to a predator, or by experimental removal of the nest, plasma levels of T increased as renesting, to replace the clutch, occurred. Circulating levels of prolactin in males began to rise during the egg-laying stage of the first brood, reached a maximum toward the end of the incubation stage, remained elevated until breeding was terminated, and then declined throughout the moulting stage to basal values in October. Prolactin levels remained high throughout the breeding season irrespective of whether a brood was raised successfully or whether the nest was lost and renesting occurred. In females, plasma levels of E2 were elevated prior to the egg-laying stage for each brood as is typical of multiple-brooded species. However, prolactin titers rose dramatically during egg-laying for the first clutch (slightly higher than in males) and were maximal by onset of incubation. Only females of this species incubate, although males do feed young. As in males, plasma prolactin in females remained high between broods and during experimentally induced renesting, and then declined to basal by the end of the moult stage in October. These data suggest that there are no differences in the temporal patterns of prolactin concentrations in blood between multiple-brooding and renesting. In a separate experiment, captive male song sparrows were transferred from a short day to artificial long days (18L 6D) and a control group was maintained on 9L 15D. In the long-day group, prolactin levels rose abruptly over the first 20 days, as the testes developed, and remained high well into postnuptial moult after the gonads had regressed. Prolactin remained basal in the control group. These data suggest that the temporal pattern of circulating prolactin levels throughout the breeding season is regulated at least partly by changing photoperiod. However, nonphotoperiodic factors are also important since photoperiodically induced increases in prolactin are significantly less than those seen in free-living individuals. These differences may be related to parental behavior.     

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.     

A neuroendocrine model for prolactin as the key mediator of seasonal breeding in birds under long- and short-day photoperiods

Seasonal breeding is associated with sequential increases in plasma luteinizing hormone (LH) and prolactin in the short-day breeding emu, and in long-day breeding birds that terminate breeding by the development of reproductive photorefractoriness. A model of the avian neuroendocrine photoperiodic reproductive response is proposed, incorporating a role for prolactin, to account for neuroendocrine mechanisms controlling both long- and short-day breeding. The breeding season terminates after circulating concentrations of prolactin increase above a critical threshold to depress gonadotropin releasing hormone (GnRH) neuronal and gonadotrope (LH) activity. Subsequently, photorefractoriness develops for prolactin secretion and for LH secretion, independently of high plasma prolactin. The breeding season in the emu is advanced compared with long-day breeders, because after photorefractiness for both LH and prolactin secretion is dissipated, plasma concentrations of both hormones increase to maximum values while days are still short.     

Don'T put all your eggs in one nest: spread them and cut time at risk

Abstract In many egg-laying animals, some females spread their clutch among several nests. The fitness effects of this reproductive tactic are obscure. Using mathematical modeling and field observations, we analyze an unexplored benefit of egg spreading in brood parasitic and other breeding systems: reduced time at risk for offspring. If a clutch takes many days to lay until incubation and embryo development starts after the last egg, by spreading her eggs a parasitic female can reduce offspring time in the vulnerable nest at risk of predation or other destruction. The model suggests that she can achieve much of this benefit by spreading her eggs among a few nests, even if her total clutch is large. Field data from goldeneye ducks Bucephala clangula show that egg spreading enables a fecund female to lay a clutch that is much larger than average without increasing offspring time at risk in a nest. This advantage increases with female condition (fecundity) and can markedly raise female reproductive success. These results help explain the puzzle of nesting parasites in some precocial birds, which lay eggs in the nests of other females before laying eggs in their own nest. Risk reduction by egg spreading may also play a role in the evolution of other breeding systems and taxa-for instance, polyandry with male parental care in some birds and fishes.