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.     

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.     

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.     

Predicting reproductive success from hormone concentrations in the common tern (<Emphasis Type="Italic">Sterna hirundo</Emphasis>) while considering food abundance

In birds, reproductive success is mainly a function of skill or environmental conditions, but it can also be linked to hormone concentrations due to their effect on behavior and individual decisions made during reproduction. For example, a high prolactin concentration is required to express parental behaviors such as incubation or guarding and feeding the young. Corticosterone level, on the other hand, is related to energy allocation or stress and foraging or provisioning effort. In this study, we measured individual baseline prolactin and corticosterone between 2006 and 2012 in breeding common terns (Sterna hirundo) using blood-sucking bugs. Reproductive parameters as well as prey abundance on a local and a wider scale were also determined during this period. Baseline prolactin and corticosterone varied significantly between years, as did breeding success. At the individual level, prolactin was positively and corticosterone was negatively linked to herring and sprat abundance. At the population level, we also found a negative link between corticosterone and prey abundance, probably reflecting overall foraging conditions. High prolactin during incubation was mainly predictive of increased hatching success, potentially by supporting more constant incubation and nest-guarding behavior. It was also positively linked to a lesser extent with fledging success, which could indicate a high feeding rate of young. Corticosterone concentration was positively related to high breeding success, which may be due to increased foraging activity and feeding of young. In general, our study shows that baseline prolactin and corticosterone levels during incubation can predict reproductive success, despite the presence of an interval between sampling and hatching or fledging of young.     

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.     

Endocrine changes in the incubating and brooding turkey hen

Turkey hens were allowed to incubate eggs and to hatch and rear young. Plasma prolactin (Prl) levels increased prior to the start of continuous incubation and rose sharply as incubation progressed to reach a peak of 1178.2 +/- 221.8 ng/ml (mean +/- SEM) just before hatching. Prl levels then fell precipitously before the hens left the nest, and returned to preincubation levels (36.8 +/- 3.4 ng/ml) by the time the poults were 2 weeks old. These results show that the high plasma concentrations of Prl found during incubation are not initiated or maintained only by the stimulus of nesting. We suggest that the decline in Prl levels at the end of incubation could be related to the pipping and hatching of eggs, and the consequent shift to maternal behavior. Plasma growth hormone (GH) levels were significantly increased in hens which were brooding poults, but not in hens incubating eggs. An elevenfold, 1-day increase in plasma GH was observed immediately after the hens left the nests. Mean plasma GH levels rose from 12.0 +/- 4.7 ng/ml on the day that the hens left the nests to 133.0 +/- 32.0 ng/ml on the following day, and then declined to 23.1 +/- 9.6 ng/ml after an additional day. There were no significant changes in plasma thyroxine levels during laying, incubation and brooding. Plasma glucose concentration was significantly depressed during incubation.     

Sex-specific patterns in baseline corticosterone and body condition changes in breeding Red-footed Boobies Sula sula

Plasma levels of corticosterone often mirror changes in body condition and parental effort. In seabirds, the brooding of young chicks is often paralleled by a marked decline in adult body condition. This may reflect the trade-off between the needs of the chick to be fed regularly and brooded, and that of the adult to spend enough time at sea to restore its body reserves and find food for the chick. In this paper, we describe changes in body condition and baseline corticosterone levels in a tropical seabird, the Red-footed Booby Sula sula , in which the chick is continuously brooded for 5–6 weeks. Body condition did not decline during the brooding period. Female body condition remained stable throughout breeding whereas male body condition declined significantly during the late chick-rearing period. Baseline corticosterone levels were low during the prelaying phase, increased significantly during incubation and were highest during the brooding period. Later in the chick-rearing period, baseline corticosterone levels decreased markedly in females, but stayed elevated in males. There was no correlation between baseline corticosterone and body condition. Because of the chick's slow growth and reduced daily need for food, brooding in Red-footed Boobies may not require a marked increase in parental effort, and the rise in corticosterone levels is probably more a reflection of slight changes in foraging activity. Red-footed Booby males are 15% smaller than females and we suggest that the decline in male body condition during the late chick-rearing period is likely to result from higher energetic flight costs or a lower foraging efficiency imposed by a smaller body size.     

Serum prolactin levels and crop-sac development in ring doves during a breeding cycle

Using a turkey prolactin radioimmunoassay, the serum prolactin levels of male and female ring doves (Streptopelia risoria) during the breeding cycle were measured and their circulating prolactin levels were compared to crop-sac weight on a within-bird basis. During the early phase of the incubation period, crop weight showed a delayed response to prolactin stimulation and there was no correlation; during the midincubation period when prolactin and crop-sac weight were increasing, there was a strong positive correlation; around hatching, when prolactin was at its peak, there was no correlation. There were again strong positive correlations at later samples, when squabs were developing and both prolactin and crop-sac weight were declining. Thus, it appears that the correlation between the circulating prolactin level and crop-sac development depends on the stage of the breeding cycle. While males and females showed similar pattern of circulating prolactin during the period of incubation and parental care, only females consistently showed a postovulatory rise of prolactin. These results were discussed in the context of the role of prolactin in the breeding cycle.     

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.     

Blood profile of pigeons (Columba livia) during growth and breeding

Changes in the blood profile of domestic pigeons (Columba livia) were studied during growth and breeding cycle. Counts of erythrocytes and leucocytes, and values of mean corpuscular volume (MCV), mean corpuscular haemoglobin (MCH), blood volume, plasma volume (BV), width of erythrocytes, and length, width and volume of erythrocyte nuclei of squabs almost reached adult values by the 4th week of age. During courtship and mating, while the level of plasma glucose increased, those of albumin, potassium, cholesterol, calcium and uric acid decreased. At nest-building, plasma albumin and plasma calcium increased significantly. The initial phase of incubation showed an elevation in plasma calcium and a decline in cholesterol and sodium, whereas mid-phase of incubation indicated a marked rise in cholesterol and uric acid. Terminal phase of incubation had significantly low plasma protein level. During feeding and brooding period, a significant rise in sodium, protein and glucose levels and a fall in calcium were observed. Following egg-laying, there was a significant rise in calcium and a drop in protein, haemoglobin, cholesterol, sodium and MCH values. Concomitant with the phenomenal rate of growth of squabs, their haematological indices neared adult values by the 4th week of age and during breeding activity significant changes in blood values occurred.     

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.     

Testosterone and prolactin in two songbirds that differ in paternal care: the Blue-headed Vireo and the Red-eyed Vireo

We investigated the hypothesis that species differences in paternal care in birds may result from differences in concentrations of circulating testosterone (T) and prolactin (PRL). Concentrations of plasma T and PRL were compared in breeding Blue-headed Vireos (Vireo solitarius) and Red-eyed Vireos (Vireo olivaceus), passerine congeners with biparental and maternal incubation, respectively. In male Blue-headed Vireos, plasma T remained low from prenesting to fledgling stages; whereas in male Red-eyed Vireos, plasma T was highest during prenesting and progressively decreased during incubation, nestling, and fledgling stages. In male Blue-headed Vireos, plasma PRL was similar to that in female Blue-headed Vireos and was higher than in male Red-eyed Vireos at all breeding stages. Plasma PRL increased in male Red-eyed Vireos at the incubation stage and remained moderately elevated through the nestling and fledgling stages. In male Blue-headed Vireos, the combination of high PRL and low T during the prenesting stage may promote brood patch formation and nest building, and, at later stages, incubation and feeding of young. In male Red-eyed Vireos, high PRL and low T during incubation and nestling stages may facilitate the feeding of young, as seen in males of other species. Our observations support the hypothesis that differences in paternal care reflect differences in circulating T and PRL.     

Alteration in haematocrit values and plasma protein fractions during the breeding cycle of female pigeons, Columba livia

Haematocrit measurements and plasma protein electrophoresis were carried out in female pigeons (Columba livia) during the breeding cycle. Non-laying birds showed higher (p < 0.001) haematocrit values than those involved in courtship and mating, nest-building, incubation and feeding and brooding. Significant differences in relative percentage of various plasma protein fractions were observed as a function of breeding. Haematocrit could be a useful index to the non-laying status of pigeons like that in chickens, Gallus domesticus.     

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.     

Vasoactive intestinal polypeptide-like immunoreactivity during reproduction in doves: influence of experience and number of offspring

To assess the possibility that vasoactive intestinal polypeptide (VIP) plays a role in naturally occurring changes in prolactin secretion in ring doves, we used immunohistochemical techniques to measure VIP-like immunoreactivity in the brain as a function of stage of the reproductive cycle. Differences between parental and nonparental birds in VIP profiles were detected in the ventral portion of the infundibular region. More specifically, there is an increase in cell size and staining intensity in the ventral infundibulum of breeding birds compared to simultaneously processed tissue taken from control animals. In both sexes, an increase in size of VIP-like immunoreactive cells is detectable during courtship and early incubation, anticipating the increase in plasma prolactin levels. VIP cell size remains elevated from about Incubation Day 14 to Brooding Day 14, and a steady decrease is observed during the remaining posthatching period, as squab begin to feed independently. Compared to parents rearing one squab, those with two young have a prolonged interval of increased infundibular VIP immunoreactivity. Furthermore, doves with no previous experience of a breeding cycle exhibit prolonged VIP-like immunoreactivity compared to experienced parents, paralleling previously described differences between these groups in parental behavior.     

鸟类的父本行为及其内分泌调控机制研究进展

飞翔能力帮助鸟类极大地扩展了生存空间,使其能够迅速移动来应对各种局部环境的变化。然而,个体发育早期,不具备这样的能力,使其极不适应环境,需要大量的亲本照顾。不同于哺乳类动物主要依赖母本行为,雄性鸟类表现出了大量的父本行为,诸如筑巢行为、孵卵行为和育雏行为等。针对这一系列行为及其调控机制,本文进行了归纳:1)筑巢行为常常伴随着雄性的求偶行为,睾酮对两种行为均有促进作用,加压素也促进筑巢行为。2)孵卵行为可以传递热量,保护和帮助子代发育;同时会刺激泌乳素释放,而泌乳素又会进一步促进孵卵行为;此时的睾酮水平下降。3)育雏行为与雏鸟的发育速度有关,晚成鸟需要更多的父本照顾;与孵卵行为的调控相似,泌乳素升高,睾酮降低;孕酮也参与调控父本行为。4)环境压力通过影响皮质酮和泌乳素水平的变化,调节鸟类的父本行为。综上,在鸟类父本行为形成和表现过程中,睾酮和泌乳素先后发挥着主要的调控作用,并且受到环境压力的影响。然而,在神经内分泌水平上,研究非常缺乏,需要加强。     

DEVELOPMENT OF THE MAGNUM REGION OF THE OVIDUCT IN RELATION TO THE REPRODUCTIVE CYCLE OF THE BARBARY DOVE STREPTOPELIA ROSEOGRISEA VAR.

The magnum region of oviducts of female Barbary Doves was examined at different phases of development before egg laying, and during incubation and brooding of the young. The tubular glands in the mucosal folds of the magnum region increased in size with increase in precursor albumen granule content during the pre-egg-laying phase, and decreased during incubation and brooding. Enlargement of tubular glands was correlated with an increase in plasma progesterone level. The temporal relationships between ovarian development, ovarian hormones, oviduct development and nesting behaviour are discussed.     

BREEDING BIOLOGY OF THE MANX SHEARWATER PUFFINUS PUFFINUS

Studies on the breeding biology of Puffinus puffinus were carried out in 1963 and 1964 at the large colony on Skokholm, Wales. During the six weeks before laying the birds spent up to a quarter of the days in the burrows, but the ten days immediately prior to laying were normally spent at sea. There is a prolonged laying period, with a marked peak in the first half of May. Details are given of a second egg being laid when the first was deserted immediately after being laid. The male took the first incubation spell. The incubation spells ranged from one to 26 days and averaged six. The incubation period was about 51 days. The frequency of visits to land by breeding birds, unlike those by non-breeders, was not affected by the moon. On hatching, the chicks grew rapidly and reached maximum weights of between 505 and 755 gm. sometime between 39 and 61 days. There was a variable desertion period, usually eight or nine days, before the chicks left the island about 70 days after hatching. During the feeding period the chick received about two feeds every three days. There is evidence that adults visited the chicks more frequently than this. There was no correlation between growth of the chicks, their feeding rates or fledging weights and the time of laying. There was a high survival (about 95 %) of chicks during the fledging period but some eggs were lost in disputes for burrows. Nine pairs in 1964 were unable to raise two young simultaneously. Parents altered their feeding rhythms to try to feed two young but did not themselves lose weight. It is suggested that the critical factor in the production of young is the availability of food for the young immediately after they leave the colonies.     

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.     

Behavioural insensitivity to supplementary testosterone during the parental phase in the chestnut-collared longspur,Calcarius ornatus

In seasonally breeding male birds that show paternal care, plasma testosterone (T) peaks in the early spring coincident with territory establishment and pair bonding. Once nests are initiated, T declines precipitously and remains low as nestlings hatch and males become parental. A growing body of evidence suggests there is a trade-off between a male's level of T and his expression of paternal care: high T enhances sexual behaviour but reduces paternal care. Male chestnut-collared longspurs show high levels of paternal care. They feed and brood young, act as nest sentries and aggressively defend the nest against predators. We experimentally altered males' plasma levels of T such that the early spring T peak was extended through the parental phase. T enhanced song behaviour typical of the sexual phase and decreased nest sentry behaviour during incubation, but did not reduce brooding or feeding of nestlings. Thus, males responded behaviourally to T during incubation, but, unlike most other temperate-breeding birds studied, did not respond to T after eggs hatched. Based on these data and similar results in an Arctic passerine, we suggest that the pattern of males' reduced responsiveness to T during the nestling phase may reflect the relative importance of male care to nest success in this species, or a phylogenetic constraint common to the genusCalcarius . Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.