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A. R. Goldsmith
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Concentrations of prolactin were measured in the plasma of male and female canaries serially sampled during repeated breeding cycles. Concentrations in female canaries were low during nest building and increased significantly (P < 0·05) during the first few days of incubation. Levels increased further (P < 0·01) in the middle of the incubation period to reach concentrations which were ten times higher than before breeding. Levels remained high for several days after the eggs hatched, as the young were fed in the nest. Prolactin concentrations declined gradually as the young were reared, reaching basal levels by the time the young were fledged, but always increased again in the females as they began incubating eggs in subsequent breeding cycles. Male canaries, which do not incubate but do assist in feeding the young, showed only slight increases in prolactin during the parental period. Male and female canaries which did not breed had low levels of prolactin throughout the experiment. The results show that prolactin secretion is high not only during incubation but also during the parental feeding phase in an altricial passerine bird. These findings are discussed in relation to the different patterns of prolactin secretion which occur during breeding in other birds.

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A. Dawson
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A. R. Goldsmith
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ABSTRACT

Is the extended prepubertal period which occurs in most birds due to a juvenile photorefractoriness analogous to photorefractoriness in adults? Starlings were taken from nest boxes 4 days after hatching and hand-reared on long days or short days. In females on long days, hypothalamic gonadotrophin-releasing hormone (GnRH) content and plasma LH concentration remained low. In intact males on long days, plasma LH remained low and the testes remained small, and in castrated birds, plasma LH also remained low. In females reared on short days, hypothalamic GnRH content began to increase dramatically from 4 weeks of age and plasma LH increased from 6 weeks of age. In intact males on short days, there was a slight increase in plasma LH and testicular size, and in castrated males, plasma LH increased markedly from six weeks of age. All birds on long days moulted into adult plumage, whereas those on short days retained juvenile plumage. The changes in birds reared on short days were similar to those found when photorefractory adult birds are transferred to short days. This adds to the evidence that juvenile photorefractoriness is analogous to photorefractoriness in adults, and therefore that the seasonal termination of photorefractoriness is an annual puberty. However, the responses to short days occur later in birds raised on short days than in older birds transferred to short days, which suggests either that short day-induced changes occur more slowly in young birds, or that birds only respond to short days after a certain age.

Journal of Endocrinology (1989) 123, 189–196

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A. Dawson
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A. R. Goldsmith
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ABSTRACT

Intact and gonadectomized male and female starlings (Sturnus vulgaris) were held in outdoor aviaries for 1 year. Blood samples were taken frequently and assayed for LH and prolactin, and intact males were laparotomized to determine testis size.

Changes in testis size in intact males were similar to those in wild starlings with maximum size during April. Hormone levels in intact males and females also followed a similar pattern to those in wild birds although absolute levels were somewhat lower. Levels of LH increased during spring, were highest in early May and then declined rapidly as birds became photorefractory. Prolactin levels did not begin to increase until April, reached a peak in late May and then declined. In gonadectomized males and females, high levels of LH were maintained throughout the early part of the year when birds were photosensitive, but declined rapidly as they became photorefractory. This occurred at the same time as in intact birds. During photorefractoriness, LH levels in gonadectomized birds were similar to those in intact birds. Prolactin showed the same seasonal pattern in gonadectomized birds as that in intact birds. These results demonstrate that gonadal steroid feedback is unimportant in initiating and maintaining photorefractoriness and confirm that increased prolactin secretion coincides with the onset of photorefractoriness.

J. Endocr. (1984) 100, 213–218

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A. Dawson
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A. R. Goldsmith
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Previous studies have shown that an increase in prolactin secretion coincides with the beginning of the photorefractory phase in starlings (Sturnus vulgaris). To determine how closely this increase is associated with the onset of refractoriness, we kept starlings on photoperiods which would induce refractoriness at different rates. Four groups of males and females were transferred from 8 h light: 16 h darkness (8L : 16D) to either 18L : 6D, 13L : 11D, 11L : 13D or 8L : 16D. Weekly blood samples were assayed for LH, FSH and prolactin and the males were frequently laparotomized to determine testis volume.

In males, both 18L and 13L induced marked increases in plasma gonadotrophin levels and rapid gonadal maturation. A photoperiod of 18L induced an earlier onset of refractoriness (2–4 weeks) than 13L (6 weeks). In both cases prolactin levels began to increase immediately before the onset of refractoriness. Males on 11L showed a slight increase in gonadotrophin levels and slow gonadal maturation. There was no increase in prolactin levels and these birds did not become refractory. Hormonal data from the females were similar to those from males. These results confirm that increased prolactin secretion is associated, in some way, with the onset of photorefractoriness in this species, although whether it is a cause or consequence of photorefractoriness, or simply of gonadal regression, remains unclear.

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A. R. Goldsmith
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T. J. Nicholls
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ABSTRACT

Male starlings which had fully developed testes under 11-h daily photoperiods were subjected for 10 weeks to daylengths either increasing or decreasing by 30 min per week. In both cases testicular regression occurred; in the former case this was complete due to photorefractoriness, and was accompanied by a postnuptial moult and a marked (P <0·01) rise in plasma prolactin concentration. In the latter case the rate of gonadal regression was slow and variable and no moulting was observed. Plasma prolactin remained very low. Eventually the birds which had experienced decreasing photoperiods were transferred to long (16 h) days; they were shown to be photosensitive since their testes redeveloped rapidly. There was also a rise in plasma prolactin, the birds eventually showed the full testicular regression characteristic of photorefractoriness, and moulting occurred. These results show that raised prolactin secretion is not associated with short-day-induced testicular regression but only with long-day-induced photorefractoriness.

J. Endocr. (1984) 102, 353–356

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A. R. GOLDSMITH
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D. M. WILLIAMS
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The concentrations of prolactin and LH were measured in the plasma of male and female mallards (Anasplatyrhynchos) during the breeding season. The concentration of prolactin in the female birds was low before and during egg-laying, increased threefold (P < 0·001 ) during incubation and returned to basal levels immediately after the young were hatched. The drakes, which do not participate in incubation in this species, had fairly low prolactin levels throughout the breeding period; levels being slightly higher during the incubation phase than at other times but still much lower (P < 0·001) than in the incubating females. Concentrations of LH in the females were high during egg-laying but declined at the onset of incubation. In the drakes levels of LH remained high for much longer and did not appear to decline at a fixed time relative to the reproductive cycle of the female birds. These observations support the view that prolactin is associated with incubation in birds, though the data do not permit a conclusion as to whether an increase in prolactin secretion causes incubation behaviour or is stimulated by it.

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A. R. GOLDSMITH
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C. EDWARDS
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M. KOPRUCU
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R. SILVER
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The concentrations of prolactin and LH were measured in the plasma of male and female ring doves (Streptopelia risoria) during the breeding cycle. Prolactin concentrations were low during courtship and early incubation but increased markedly (P<0·01 in both sexes) between days 7 and 15 of incubation, during the period of rapid growth of the crop sac. Levels remained raised for 4 days after the young had hatched but then declined as the crop sacs gradually regressed. These observations showed that concentrations of prolactin were high at the time when the crop sacs were proliferating and producing 'milk' for feeding the young, but that there was no increase in prolactin secretion at the onset of incubation as occurs in ducks, hens and other birds. In samples taken from doves at various times of the day and night there was no increase in plasma prolactin during the first few days of incubation. Concentrations of LH in plasma were generally higher in male doves than in females, but in both sexes were significantly higher during courtship than during incubation and brooding the young. Concentrations increased again when the young were independent and the parents began a second round of courtship.

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F. J. McNaughton
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A. Dawson
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A. R. Goldsmith
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ABSTRACT

If adult starlings are kept on long days, they rapidly become refractory to long days (photorefractory). The recovery of the ability to respond to long days (photosensitivity) occurs after transfer to short days. Photosensitivity is associated with an increase in hypothalamic content of gonadotrophin-releasing hormone (GnRH), and, in castrated birds, with an increase in plasma LH. If photorefractory adult or juvenile starlings are transferred to short days, these events begin 3 and 4 weeks respectively after transfer. Starlings apparently hatch in a photorefractory state. However, if nestlings are hand-reared under short days, the increases in GnRH and LH begin at 6 and 7 weeks of age respectively. Is this 3-week delay, which corresponds to the time to reach adult body size, simply due to a slower response, or are nestlings incapable of responding to short days until fully grown? Castrated male nestling starlings were hand-reared under short days, under long days or under long days initially and transferred to short days at 2, 3, 6 or 9·5 weeks of age. The increase in plasma LH in birds transferred to short days at or before 3 weeks began at the same time as in birds kept on short days (7 weeks). In birds transferred at 3 weeks or later, the increase in LH began 4 weeks after transfer, irrespective of age. Intact female starlings showed similar responses. Birds kept on short days did not moult whereas birds transferred from long days to short days at 3 weeks did. These results suggest that nestlings are capable of perceiving daylength, but that the GnRH neurosecretory system does not respond to short days until birds are fully grown.

Journal of Endocrinology (1992) 132, 411–417

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A. R. Goldsmith
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S. Burke
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J. M. Prosser
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ABSTRACT

Prolactin and LH concentrations were measured in the plasma of female canaries sampled during the breeding cycle and after disruption and reinitiation of incubation behaviour. The late incubation period was characterized by low LH and high prolactin concentrations, and canaries separated from their nests at this stage showed an increase in LH and a decline in prolactin within 3 h. In one experiment mean (±s.e.m.) concentrations before and 24 h after nest deprivation were: prolactin 397 ± 86 and 18 ± 5 μg/l; LH 1·04±0·21 and 2·03±0·17 μg/l. Female canaries which abandoned their nests after the eggs had been removed also showed an increase in LH together with a fall in prolactin 24 h after egg removal. When nest-deprived canaries were allowed to resume incubation, plasma prolactin increased again within 5 h and after 2 days had reached levels normal for incubating birds (398± 46 μg/l). Concentrations of LH changed more gradually but had decreased 2 days after the resumption of incubation. Thus prolactin and LH show inverse changes after the disruption and reinitiation of incubation behaviour; it is not clear, however, if the change in one is dependent on the other or if both hormones are responding to the same external stimulus.

J. Endocr. (1984) 103, 251–256

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A. R. Goldsmith
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T. J. Nicholls
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G. Plowman
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ABSTRACT

Sexually immature male starlings were radiothyroidectomized while held under short daylengths. They were then subjected to long (18-h) photoperiods and the testes developed rapidly and apparently normally to full maturity. As expected, thyroidectomy prevented an onset of photorefractoriness and after 140 days the testes were still fully developed (testicular width 6·5 ± 0·8 mm). Half of the birds were then given thyroxine (0·011 mmol/l) in the drinking water for 14 days while the others were maintained as controls. At the end of this time the birds still had fully developed gonads (testicular width 7·8 ± 0·3 mm) but after a further 14 days testicular regression began in the thyroxine-treated birds. The testes of all individuals in this group were fully regressed (width 1·8 ± 0·1 mm) by 56 days after the end of thyroxine administration and moulting of the flight feathers had begun. No recrudescence of the gonads was subsequently noted in the next 2 months and the birds were apparently photorefractory. The short period of thyroxine treatment also caused a rapid and prolonged increase in plasma prolactin levels from 2·0 ± 0·3 to 16·8 ± 2·6 μg/l. No testicular regression or moulting was observed in the control birds and their plasma prolactin levels remained very low (below 4·0 μg/l) throughout the experiment.

In thyroidectomized and castrated starlings held on 18-h daylengths, 14 daily injections of thyroxine 100 μg/bird per day) caused a rapid and permanent decrease in circulating FSH to basal levels (reached about 36 days after thyroxine treatment began, at which time the birds moulted). The treatment also caused a marked increase in plasma prolactin lasting for about 50 days.

J. Endocr. (1985) 104, 99–103

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