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.     

Photoperiodic control of the termination of breeding and the induction ofmoult in House Sparrows Passer domesticus

Two photoperiodic mechanisms controlling gonadal regression in birds have been identified: absolute photorefractoriness, typical of species with short breeding seasons, where gonadal regression occurs spontaneously during long days, and relative photorefractoriness, where a decrease in daylength is required to induce regression. An experiment was designed to test whether these simply represent extremes of one underlying mechanism. Three groups of male House Sparrows Passer domesticus were transferred from a short photoperiod, 8 h of light: 16 h of darkness per day (8L:16D) to long photoperiods of either 18L:6D, 16L:8D or 13L:11D. Gonadal maturation rates were similar in all three groups; gonadal regression and moult began latest in the 13L:11D group. Four additional groups of sparrows were transferred from 8L:16D to 18L:6D and then transferred to either 13L: 11D or 16L:8D prior to, or shortly after, the onset of gonadal regression. The decrease in daylength prior to regression had no effect on the timing of regression but did advance the onset of moult. Decrease in daylength after the onset of regression increased the rate of regression and the rate of moult. Because a decrease in daylength did not affect the timing of regression, the data do not support the hypothesis that absolute and relative photorefractoriness represent extremes of a single underlying photoperiodic control mechanism. The adaptive significance of the effects of decreasing daylength on the rate of regression and moult is discussed.     

Photosensitivity in body mass and testicular activity of brahminy myna, Sturnus pagodarum

This study analyzed photoperiodic sensitivity of brahminy myna (Sturnus pagodarum), which is a seasonally breeding bird species. During regression phase of the reproductive cycle (in early September), groups of myna were exposed to artificial photoperiods that were either close to or longer than those brahminy myna experiences at this time in wild at 29 degrees N. Following a 14-week exposure to such photoperiods (hours of light: hours of darkness; 13L:11D, 12L:12D, 11.5L:12.5D and 11L:13D), the birds were subjected to a longer day length (16L:8D) for another 9 weeks to test whether pre-treatment with varying photoperiods had an effect on subsequent long day photostimulation. There was a progressive increase in body mass under different pre-treatment photoperiods, with a faster increase in 11L:13D and 11.5L:12.5D than in 12L:12D and 13L:11D. When subjected to 16L: 8D, however, all groups showed decline in body mass. By contrast, the testes were not stimulated under 11L:13D and 11.5L:12.5D, and only one individual of the groups under 12L:12D and 13L: 11D showed a small testis recrudescence. All birds except one individual of the 13L:11D group, however, showed testis recrudescence when subjected to 16L:8D. These results suggest that body mass and testes have a different profile of photoperiodic response and appear to have two different threshold photoperiods for dissipating the post-reproductive refractoriness.     

Photoperiodic history and a changing melatonin pattern can determine the neuroendocrine response of the ewe to daylength

The reproductive neuroendocrine response of Suffolk ewes to the direction of daylength change was determined in animals which were ovariectomized and treated with constant release capsules of oestradiol. Two groups of animals were initially exposed to 16 or 10 h light/day for 74 days. On day zero of the study, when one group of ewes was reproductively stimulated (elevated LH concentrations) and the other reproductively inhibited (undetectable LH concentrations), half the animals from each group were transferred to an intermediate daylength of 13 h light/day. The remaining ewes were maintained on their respective solstice photoperiods to control for photorefractoriness. LH concentrations rose in animals experiencing a 3 h decrease in daylength from 16L:8D to 13L:11D while LH concentrations fell to undetectable values in those that experienced a 3 h increase in daylength from 10L:14D to 13L:11D. The photoperiodic response of the Suffolk ewe, therefore, depends on her daylength history. Such a result could be explained if the 24-h secretory pattern of melatonin secretion, known to transduce photoperiodic information to the reproductive axis, was influenced by the direction of change of daylength. Hourly samples for melatonin were collected for 24 h 17 days before and three times after transfer to 13L:11D. The melatonin secretory profile always conformed to daylength. Therefore, the mechanism by which the same photoperiod can produce opposite neuroendocrine responses must lie downstream from the pineal gland in the processing of the melatonin signal.     

Timing of photorefractoriness in the European starling: significance of photoperiod early and late in the reproductive cycle

In European starlings, as in many other birds inhabiting higher latitudes, gonads develop in response to the increasing daylengths in early spring. Later in the year, however, the hypothalamo-pituitary-gonadal axis becomes refractory to the previously stimulatory long photoperiods and the gonads regress in summer. The present study addresses the question of when during the gonadal growth phase photorefractoriness is determined. A 13-h photoperiod induces testicular development and subsequent testicular regression associated with refractoriness in male starlings. An 11-h photoperiod, in contrast, induces only testicular development, and photorefractoriness never develops. When starlings were transferred to an 11-h photoperiod, either 12 or 25 days following exposure to a 13-h photoperiod, their testes developed to full size, but remained large to the end of the experiment, i.e. refractoriness did not develop. The same was even true of most birds in a third group that were transferred to an 11-h photoperiod after 46 days of the 13-h photoperiod, when gonads had developed to near maximal size. These data show that, in contrast to some other species of passerine birds, the onset of photorefractoriness does not become fixed before the testes have undergone considerable development, and that the photoperiodic conditions experienced at the end of the testicular growth phase are still effective in determining the precise time of onset of photorefractoriness. It is suggested that this peculiarity of the starling is related to the fact that its gonadal development begins rather early in spring and, hence, under much shorter photoperiods than the other species studied.     

Influence of different photoperiods on development of gonad, cloacal gland and circulating thyroid hormones in male Japanese quail Coturnix coturnix japonica.

One day old chicks of Japanese quail were exposed to different photoperiods (LD, 8:16, 13.5:10.5, 16:8 and LL) and observations (testes weight, cloacal gland size, body weight and circulating thyroxine and triiodothyronine) were taken at the age of 3, 5, 7, 9 and 16 weeks. Results indicate that immediate reproductive development occurred in birds exposed to long photoperiods (greater than 12 hr). Growth under LD 8:16, was not apparent till 7th week and by 16 weeks, degree of gonadal development was similar in all the birds, irrespective of photoperiodic treatment. Whereas body weight of the intermediate and long day (LD 13.5:10.5, 16:8 and LL) treated birds increased upto 5th week and remained constant thereafter. But the chicks maintained under short day length (LD 8:16), showed spontaneous increase till the end of the study and birds were much heavier compared to all other groups. Plasma T4 concentration increased with increasing age till 9th week and remained unaltered thereafter. On the other hand T3 level did not change till 7th week followed by a decline. It is suggested that the initiation and degree of gonadal growth in quail depends on the availability of daily photoperiod, until the achievement of full breeding condition. Peak level of T4 observed in 9 week old birds may be involved in the development of photorefractoriness at that age.     

Regulation of seasonality in the migratory male blackheaded bunting (Emberiza melanocephala)

The present study was carried out on a Palearctic-Indian migratory species, the blackheaded bunting (Emberiza melanocephala), to understand the importance of photoperiodism and circannual rhythms in determining seasonality in changes in body mass and testis size in birds. An initial experiment determined the effects of duration and intensity of light on photoperiodic induction. The birds were exposed to different photoperiods (hours of light:hours of darkness; 11.5L:12.5D, 12L:12D, 12.5L:11.5D and 13L:11D) at the same (approximately 450 lux) light intensity, and to 13L:11D at different light intensities (50-, 100-, 400-, 800- and 1000-lux). The induction and subsequent regression of photoperiodic responses were dependent upon duration and intensity of the light period until these reached threshold. A second experiment investigated if an endogenous seasonal rhythm underlies photoperiodism in buntings. Birds maintained since February on a 8L: 16D photoperiod (a non-inductive short day length invariably used to ensure photosensitivity in photoperiodic species) were subjected periodically to 16L:8D (a long day length), one group every month from mid-March to mid-August. The magnitude of long day response in body mass and testes decreased as the duration of the short days progressed, but testicular response was restored in birds that were exposed to long days in July and August. The birds exposed simultaneously to short, long, and natural day lengths for 32 weeks underwent an induction-regression cycle under long days and natural day lengths, but not under short days in which a decrease in body mass occurred after about 20 weeks. The last experiment examined the importance of latitudinal migration on photoperiodism, by comparing the response to long days of three groups which included birds from populations those were held in the outdoor aviary for 1 or 2 years at 27 degrees N and those immediately arrived from their breeding grounds (approximately 40 degrees N). There was no difference in the photoperiodic induction among the three groups, indicating that neither experience to changing photoperiods during a migratory journey, nor to long photoperiods at breeding grounds, were critical for a subsequent response (initiation-termination-reinitiation) cycle. Taken together, these findings suggest that (1) the blackheaded bunting has its own endogenous timing program, which is regulated by the photoperiod, and (2) the photoperiodic programs of bunting are flexible enough to accommodate variations in the amplitude of environmental cycles. Thus, it appears that photoperiodism has evolved independently of the evolution of migration in this species.     

Support for a rare pattern of temperature-dependent sex determination in archaic reptiles: evidence from two species of tuatara (Sphenodon)

Background

In many birds, day length (=photoperiod) regulates reproductive cycle. The photoperiodic environment varies between different seasons and latitudes. As a consequence, species at different latitudes may have evolved separate photoperiodic strategies or modified them as per their adaptive need. We studied this using house sparrow as a model since it is found worldwide and is widely investigated. In particular, we examined whether photoperiodism in house sparrows (Passer domesticus) at 27°N, 81°E shared features with those exhibited by its conspecifics at high latitudes.

Results

Initial experiment described in the wild and captive conditions the gonad development and molt (only in captives) cycles over a 12-month period. Both male and female sparrows had similar seasonal cycles, linked with annual variations in day length; this suggested that seasonal reproduction in house sparrows was under the photoperiodic control. However, a slower testis and attenuated follicular growth among captives indicated that other (supplementary) factors are also involved in controlling the reproductive cycle. Next experiment examined if sparrows underwent seasonal variations in their response to stimulatory effects of long day lengths. When birds were transferred every month over a period of 1 year to 16 hours light:8 hours darkness (16L:8D) for 17–26 weeks, there was indeed a time-of-year effect on the growth-regression cycle of gonads. The final experiment investigated response of house sparrows to a variety of light-dark (LD) cycles. In the first set, sparrows were exposed for 31 weeks to photoperiods that were close to what they receive in between the period from sunrise to sunset at this latitude: 9L:15D (close to shortest day length in December), 12L:12D (equinox, in March and September) 15L:9D (close to longest day length in June). They underwent testicular growth and regression and molt in 12L and 15L photoperiods, but not in 9L photoperiod. In the second set, sparrows were exposed for 17 weeks to photoperiods with light periods extending to different duration of the daily photosensitivity rhythm (e.g. 2L:22D, 6L:18D, 10L:14D, 14L:10D, 18L:6D and 22L:2D). Interestingly, a slow and small testicular response occurred under 2L and 10L photoperiods; 6L:18D was non-inductive. On the other hand, 14L, 18L and 22L photoperiods produced testicular growth and subsequent regression response as is typical of a long day photostimulation.

Conclusion

Subtropical house sparrows exhibit photoperiodic responses similar to that is reported for its population living at high latitudes. This may suggest the conservation of the photoperiodic control mechanisms in birds evolved over a long period of time, as a physiological strategy in a temporally changing environment ensuring reproduction at the best suited time of the year.     

The influence of light wavelength on reproductive photorefractoriness in migratory blackheaded bunting (Emberiza melanocephala).

There are two effects of long day length on reproductive responses in birds, one is the photoinduction of gonadal growth and maturation and the other is the induction of gonadal regression and photorefractoriness. Although it is likely that the same photoreceptors are involved in the photoinduction of gonadal growth and the onset and maintenance of photorefractoriness. and so the influence of wavelength should be similar, this has not been investigated. Therefore, we investigated the influence of light wavelength on reproductive photorefractoriness in the migratory male blackheaded bunting held under long photoperiods. In mid May, when photoperiod was approximately 14L:10D (14 hours light:10 hours darkness), eight groups of sexually mature birds were moved indoors on an artificial photoperiod of 14L:10D (L - 450 lux. D - 0 lux). Then after 3 weeks, for six groups, a 4-h light period in the morning (zt 0-4; zt 0 [zeitgeber time 0] refers to the beginning of lights-on period) or in the evening (zt 10-14) was substituted with green (428 nm), red (654 nm) or white light at 16 +/- 2 lux intensity. Of the remaining two groups, one was maintained on 14L: 10D and the other transferred to 10L:14D: these served as controls. At the end of 4 weeks, all birds were found to have undergone testicular regression, irrespective of LD cycle they were exposed to. When these gonadally regressed birds were subjected to 16L:8D for another 4 weeks, to test their responsiveness to the stimulatory effects of long day lengths, only those exposed to 10L:14D and 14L:10D with a 4-h green light period showed testicular regrowth. On the other hand, birds exposed to 14L:10D with a 4-h white or red light period remained fully regressed, similar to 14L:10D controls. Except for some individual difference, there was no difference in response between the groups that received a 4-h light period in the morning and that received it in the evening. These results suggest that the wavelengths of light influence induction of buntings from the photosensitive state into the photorefractory state. Whereas the short light wavelengths facilitated recovery from the photorefractoriness, the long light wavelengths were more effective in maintaining the photorefractoriness.     

The thyroid and photoperiodic control of seasonal reproduction in American tree sparrows (Spizella arborea)

To explore the role of the thyroid gland in the control of seasonal reproduction in obligately photoperiodic American tree sparrows (Spizella arborea), the effects of (1) thyroxine administered in drinking water to thyroid-intact photosensitive or photorefractory birds, and (2) radiothyroidectomy before and after photostimulation and during photorefractoriness were examined. Chronic administration of pharmacological doses of thyroxine induced testicular growth and usually regression in initially photosensitive birds held on short or intermediate daylengths. Some thyroxine-treated birds with regressed testes were absolutely photorefractory, but most remained photosensitive. Exogenous thyroxine never induced testicular growth in photorefractory birds moved to short days, though it often impeded, and sometimes even blocked, the recovery of photosensitivity. Although circumstantial, these effects of exogenous thyroxine are consistent with an hypothesis that assigns to thyroid hormones two roles — one stimulatory and the other inhibitory — in the control of seasonal reproduction. Radiothyroidectomy before photostimulation inhibited (but did not prevent) photoinduced testicular growth, blocked spontaneous testicular regression, suppressed molt, and prevented photorefractoriness. Moreover, as demonstrated by testicular growth after thyroxine replacemnt therapy, radiothyroidectomy during photorefractoriness later restored photosensitivity despite continued photostimulation. Thus, euthyroidism is an essential condition for maximizing (but not for initiating) photoinduced testicular growth and for triggering and maintaining photorefractoriness in photostimulated tree sparrows. However, when performed early during photostimulation, radiothyroidectomy neither immediately induced nor later blocked spontaneous testicular regression. Thus, endogenous thyroid hormones and long days may interact during a critical period to program a sequence of physiological events that plays out as photorefractoriness in chronically photostimulated birds. Such an organizational event cannot be permanent, for seasonal reproduction is episodic and its control mechanism necessarily cyclic. Because thyroidectomy simulated the well-known restorative effect of short days (and exogenous thyroxine impeded it), short days may dissipate photorefractoriness by creating a milieu wherein thyroid hormones are deficient or inactive.Abbreviations ANOVA analysis of variance - bTSH bovine thyroid stimulating hormone - GnRH gonadotropin-releasing hormone - LH luteinizing hormone - nL: nD daily light: dark regime (n is duration in hours) - SEM standard error of the mean - SNK Student-Newman-Keuls test - T₄ thyroxine - TH thyroid hormone - TR thyroid hormone receptor     

Response to experimental photoperiods by a migratory bunting, Emberiza melanocephala

Little is known about the effects of photoperiod on avian migrants that visit southeast Asia. In this paper, we report experiments performed on an emberizid finch, the Black-headed Bunting Emberiza melanocephala , to investigate its photoperiodic responses under artificial photoperiods, and continuous light and darkness.
Two series of experiments were performed with the photosensitive male birds. In the first series, different groups were exposed to seven different artificial photoperiods: 3L/21D, 6L/18D, 8L./16D, 11L/13D, 12L/12D, 15L/9D and 20L/4D, for 30 days. They were weighed and laparotomized at the beginning and end of the experiments. The birds responded to 12L/12D, 15L/9D and 20L/4D, but not to 3L/21D, 6L/18D, 8L/16D and 11L/13D. In the second series, photosensitive birds were placed under continuous light (LL) and dark (DD) conditions for 130 and 90 days. Periodic observations indicated that testicular growth and fattening followed by involution and fat-depletion had resulted in birds under LL, indicating the onset of photorefractoriness, while DD had no effect either on gonads or fattening in the buntings.
Our results demonstrate that light stimulation is a prerequisite to reproductive and metabolic activities (pre-migratory and migratory changes, fattening and weight gain) in the Black-headed Bunting, which has a photoperiodic threshold to these events at between 11 and 12 h daily photoperiods.     

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.     

Reception of photoperiodic information by fetal Siberian hamsters: role of the mother's pineal gland

The rate of reproductive development in juvenile male Siberian hamsters is strongly influenced by daylength (photoperiod). Recent studies indicate that reception of photoperiodic cues begins during fetal life. The present experiments yielded a further demonstration that developing male Siberian hamsters receive information about the photoperiod to which their mother is exposed during pregnancy. The possibility that photoperiodic information is transmitted from mother to young after birth was investigated by cross-fostering young gestated on 12L and 16L to mothers from the other photoperiod. Litters were cross-fostered on the day of birth and then were transferred, along with their foster mothers, to 14L. We found no influence of the mother after birth, indicating that transmission of photoperiodic information from mother to young must occur during gestation. To determine if the pineal gland of the mother is required for this response, adult females were pinealectomized or sham-operated and paired with intact males in 12L, 14L, or 16L. After parturition parents and offspring were exposed to 14L. The influence of prenatal photoperiod on postnatal testicular development in 14L was blocked by pinealectomy of the mother. Postnatal testicular development was retarded in offspring that experienced a photoperiod transfer from either 15L to 14L or 8L to 12L at birth. In contrast, the inhibitory effect of a transfer from 16L to 14L at birth was abolished when juvenile males were exposed to a single long photoperiod (16.3 h light) at age 17-21 days and then were returned to 14L.     

Effects of pinealectomy and melatonin or 5-methoxytryptamine on testicular LH and PRL receptors in Syrian hamsters (Mesocricetus auratus)

The pineal has been previously shown to be an important factor in the regulation of testicular function in photoperiodic mammals. The effects of lack or increase in pineal hormones on testicular hormonal receptors has, therefore, been examined. Pinealectomy decreased the concentration of testicular LH receptors in hamsters exposed to either a long or short photoperiod but had no effect on the concentration of testicular PRL receptors. In animals exposed to a short photoperiod, pinealectomy prevented testicular regression and the concomitant decreases in total LH and PRL receptor contents. Treatment for 12 weeks with either melatonin or 5-methoxytryptamine caused a decrease in testicular PRL receptor levels, whereas the only changes in LH receptor levels were due to melatonin-induced testicular regression. The present results indicate that some of the effects of pineal hormones on the testes are independent of the pineal-induced changes in testes mass and are the consequence of long-term action. Furthermore, testicular function appears to be affected by both the lack or the increase in pineal hormones.     

Extraocular control of photorefractoriness in American tree sparrows (Spizella arborea)

Photorefractoriness, a reversible state of unresponsiveness to daylengths of gonadostimulatory duration, terminates seasonal breeding in many photoperiodic species of birds. Whether the eyes are components of the mechanism that triggers photorefractoriness is an important, but heretofore unresolved, question. Although a role for extraocular photoreception in the mechanism of photoinduced gonadal growth is well documented, the eyes may be important in the mechanism of photorefractoriness if, as some evidence suggests, they are gonadoinhibitory. With American tree sparrows (Spizella arborea), I here confirm that the absence of eyes does not impede photoinduced testicular growth and establish that an extraocular mechanism mediates the transition from photosensitivity to photorefractoriness: Tree sparrows blinded by bilateral ocular enucleation, when photosensitivity to long days or by miniature self-powered lights implanted atop the skull, showed marked testicular growth and then, as evidenced by spontaneous testicular regression, became photorefractory, as did sighted controls.     

Importance of changing photoperiod and melatonin secretory pattern in determining the length of the breeding season in the Suffolk ewe

Three groups of ovariectomized Suffolk ewes bearing s.c. Silastic implants of oestradiol were subjected to a 90-day priming treatment of an inhibitory long photoperiod (16 h light/day; 16L:8D). On Day 0 of the experiment, they were moved to stimulatory photoperiods. One control group was transferred to 12L:12D and a second control group was transferred to 8L:16D; both groups remained in those photoperiods to determine the timing of reproductive induction and refractoriness. The experimental group was transferred to 12L:12D on Day 0 and then to 8L:16D on Day 55 to determine whether the further reduction in daylength could delay the development of refractoriness. Reproductive neuroendocrine condition was monitored by serum concentrations of LH and FSH. Both gonadotrophins remained elevated for a longer period of time in the experimental group receiving the second reduction in daylength than in either control group, indicating that the second photoperiodic drop delayed the onset of photorefractoriness. Measurement of 24-h patterns of circulating melatonin suggests that the prolonged stimulation of reproductive neuroendocrine activity in the experimental group resulted from a lengthening of the nocturnal melatonin rise. These findings indicate that refractoriness to an inductive photoperiod can be temporarily overcome by exposure to a shorter daylength, and that the change in duration of the nocturnal increase in melatonin secretion is important in photoperiodic signalling. Thus, in natural conditions, the decreasing autumnal daylength, and the resulting expansion of the nocturnal elevation in melatonin secretion, may be utilized to produce a breeding season of normal duration.     

Effect of intranasal irrigation with zinc sulphate on male-induced oestrus in the wild mouse, Mus musculus domesticus

Adult males of the subtropical house sparrow (Passer domesticus) were subjected to continuous darkness (DD) or short days (8L:16D) after photostimulation (15L:9D) for 30 days. One group was retained on the long day (15L:9D) schedule. Another two groups of photosensitive birds were placed under DD or 8L:16D. The photostimulated birds under DD or 8L:16D showed testicular regression and the rate of regression was similar in both the treatments. The photosensitive birds, on the other hand, did not show any initiation in testicular growth and development in either of the treatments. Long day birds (15L:9D) maintained enlarged testes without regression (the onset of photorefractoriness) during the treatment period. These results conform with the external coincidence model, but could also be explained by a modified internal coincidence model.     

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.     

Testosterone accelerates the onset of photorefractoriness in baya weaver Ploceus philippinus.

The baya weaver Ploceus philippinus, despite its purely tropical origin and tropical/subtropical distribution, exhibits all features of a typically photoperiodic species. The onset of reproduction in this species is triggered by increasing daylength and breeding is terminated by the development of distinct photorefractory phase. In order to ascertain the involvement of androgens in the development of photorefractoriness the long day response of birds was tested after a prior exposure to varied doses of testosterone. Testosterone treatment accelerated the onset of photorefractoriness as judged from the gonadal status, LH-dependent yellow plumage and testosterone-dependent beak pigmentation. While gonadal development and yellow plumage occurred in the control birds due to the stimulatory long photoperiod, the testosterone administered birds failed to show such developments. These findings indicate that testosterone might impair the higher photoresponding mechanisms resulting in the development of photorefractoriness.     

The LH-RH system of the male European starling: photoperiod induces changes to a possible multifunctional peptide system

In many birds reproduction is triggered by long daylengths but, paradoxically, continued exposure to long days leads to photorefractoriness and a complete shut down of the reproductive system. As these effects are thought to be mediated through the secretion of LH-RH, immunocytochemical techniques were used to investigate changes in the LH-RH system when European starlings were exposed to different photoperiods. Starlings exposed to 11L:13D and with mature testes show strong immunostaining both of LH-RH perikarya and fibers. Photosensitive short-day (8L:16D) starlings with undeveloped testes show an almost identical distribution of strongly immunoreactive perikarya but with less dense fibre staining. However, long-day (18L:6D) photorefractory starlings with fully regressed testes, show a profound reduction in LH-RH immunostaining. Perikarya have the same distribution but show a much reduced intensity of staining and fibers had almost entirely disappeared from all regions of the brain. Preliminary observations on the ultrastructure of immunocytochemically identified LH-RH neurones are also reported.