Starling circannual systems: are they arrested in long photoperiods?

Summary In the European starling,Sturnus vulgaris, circannual rhythms in gonadal size, molt and other related functions persist only in photoperiods close to 12 h, but are absent in longer or shorter daylengths. To find out whether the arrhythmia seen in long photoperiods results from an arrest of the underlying clock system, three groups of male starlings were held for 10, 14, or 20 months in a 13 h photoperiod and then transferred to a 12 h photoperiod. A control group was held in the 13 h photoperiod throughout the experiment for 28 months. During the initial exposure to the 13 h photoperiod, all birds went through a gonadal cycle, followed by a complete molt. Subsequently, the control birds retained small testes to the end of the experiment and there was no further molt. In contrast, most of the experimental birds re-initiated a testicular cycle, following transfer to the 12 h photoperiod and molted after its completion. The latency between the transfer to the 12 h photoperiod and the onset of testicular growth was not significantly different among the three groups, indicating that the underlying circannual clock had been arrested in the 13 h photoperiod and restarted in the 12 h photoperiod. The pattern of the second testicular cycle did, however, differ among groups. Particularly its amplitude decreased from group 1 to group 3, suggesting that the capacity of the birds to respond to a 12 h photoperiod decreased with increasing duration of exposure to the 13 h photoperiod.Dedicated to Prof. Dr. C.S. Pittendrigh on the occasion of his seventieth birthday.     

Avian circannual clocks: adaptive significance and possible involvement of energy turnover in their proximate control

Endogenous circannual clocks are found in many long-lived organisms, but are best studied in mammal and bird species. Circannual clocks are synchronized with the environment by changes in photoperiod, light intensity and possibly temperature and seasonal rainfall patterns. Annual timing mechanisms are presumed to have important ultimate functions in seasonally regulating reproduction, moult, hibernation, migration, body weight and fat deposition/stores. Birds that live in habitats where environmental cues such as photoperiod are poor predictors of seasons (e.g. equatorial residents, migrants to equatorial/tropical latitudes) rely more on their endogenous clocks than birds living in environments that show a tight correlation between photoperiod and seasonal events. Such population-specific/interspecific variation in reliance on endogenous clocks may indicate that annual timing mechanisms are adaptive. However, despite the apparent adaptive importance of circannual clocks, (i) what specific adaptive value they have in the wild and (ii) how they function are still largely untested. Whereas circadian clocks are hypothesized to be generated by molecular feedback loops, it has been suggested that circannual clocks are either based upon (i) a de-multiplication ('counting') of circadian days, (ii) a sequence of interdependent physiological states, or (iii) one or more endogenous oscillators, similar to circadian rhythms. We tested the de-multiplication of days (i) versus endogenous regulation hypotheses (ii) and (iii) in captive male and female house sparrows (Passer domesticus). We assessed the period of reproductive (testicular and follicular) cycles in four groups of birds kept either under photoperiods of LD 12L:12D (period length: 24h), 13.5L:13.5D (27 h), 10.5L:10.5D (23 h) or 12D:8L:3D:1L (24-h skeleton photoperiod), respectively, for 15 months. Contrary to predictions from the de-multiplication hypothesis, individuals experiencing 27-h days did not differ (i.e. did not have longer) annual reproductive rhythms than individuals from the 21- or 24-h day groups. However, in line with predictions from endogenous regulation, birds in the skeleton group had significantly longer circannual period lengths than all other groups. Birds exposed to skeleton photoperiods experienced fewer light hours per year than all other groups (3285 versus 4380) and had a lower daily energy expenditure, as tested during one point of the annual cycle using respirometry. Although our results are tantalizing, they are still preliminary as birds were only studied over a period of 15 months. Nevertheless, the present data fail to support a 'counting of circadian days' and instead support hypotheses proposing whole-organism processes as the mechanistic basis for circannual rhythms. We propose a novel energy turnover hypothesis which predicts a dependence of the speed of the circannual clock on the overall energy expenditure of an organism.     

A tropical bird can use the equatorial change in sunrise and sunset times to synchronize its circannual clock

At higher latitudes, most organisms use the periodic changes in day length to time their annual life cycle. At the equator, changes in day length are minimal, and it is unknown which cues organisms use to synchronize their underlying circannual rhythms to environmental conditions. Here, we demonstrate that the African stonechat (Saxicola torquatus axillaris)-an equatorial songbird-can use subtle solar cues for the annual timing of postnuptial moult, a reliable marker of the circannual cycle. We compared four groups that were kept over more than 3 years: (i) a control group maintained under constant equatorial day length, (ii) a 12-month solar time group maintained under equatorial day length, but including a simulation of the annual periodic change in sunrise and sunset times (solar time), (iii) a 14-month solar time group similar to the previous group but with an extended solar time cycle and (iv) a group maintained under a European temperate photoperiod. Within all 3 years, 12-month solar time birds were significantly more synchronized than controls and 14-month solar time birds. Furthermore, the moult of 12-month solar time birds occurred during the same time of the year as that of free-living Kenyan conspecifics. Thus, our data indicate that stonechats may use the subtle periodic pattern of sunrise and sunset at the equator to synchronize their circannual clock.     

Circannual variations in plasma luteinizing hormone levels in castrated male European starlings (Sturnus vulgaris)

Intact and castrated male European starlings were held for about 2 years in a constant 12-hr photoperiod and constant temperature conditions. At 1- to 2-month intervals, testicular width was measured by laparotomy, and blood samples were taken for analysis of plasma luteinizing hormone (LH). Most of the control birds went through at least one circannual cycle of testicular width and plasma LH concentration. In the castrates, a similar proportion of birds went through circannual LH cycles with periods indistinguishable from those of the controls. It is concluded that the testes and their hormones are not essential components of the mechanism that generates circannual gonadal cycles in male European starlings.     

Temperature alters the photoperiodically controlled phenologies linked with migration and reproduction in a night-migratory songbird

We investigated the effects of temperature on photoperiodic induction of the phenologies linked with migration (body fattening and premigratory night-time restlessness, Zugunruhe) and reproduction (testicular maturation) in the migratory blackheaded bunting. Birds were exposed for four weeks to near-threshold photoperiods required to induce testicular growth (11.5 L:12.5 D and 12 L:12 D) or for 18 weeks to a long photoperiod (13 L:11 D) at 22°C or 27°C (low) and 35°C or 40°C (high) temperatures. A significant body fattening and half-maximal testicular growth occurred in birds under the 12 L, but not under the 11.5 L photoperiod. Further, one of six birds in both temperature groups on 11.5 L, and four and two of six birds, respectively, in low- and high-temperature groups on 12 L showed the Zugunruhe. Buntings on 13 L in both temperature groups showed complete growth-regression cycles in body fattening, Zugunruhe and testis maturation. In birds on 13 L, high temperature attenuated activity levels, delayed onset of Zugunruhe by about 12 days, reduced body fattening and slowed testicular maturation. The effect of temperature seems to be on the rate of photoperiodic induction rather than on the critical day length. It is suggested that a change in temperature could alter the timing of the development of phenologies linked with seasonal migration and reproduction in migratory songbirds.     

Stable daily light regimens as inductive factors of endogenous testicular cycles in the European starling,Sturnus vulgaris

European starlings (Sturnus vulgaris ) maintained under chronic 12L:12D exhibit testicular cycles with a periodicity of 9–10 months. These circannual testicular cycles incorporate all of the physiologically distinct phases observed during gonadal cycles in starlings under temperate-zone photoperiods. Starlings maintained under chronic 6L-18D also undergo testicular cycles but these cycles: (a) have a relatively short periodicity (about 6 months); (b) include periods of testicular involution, though not to the minimal quiescent level for this species; and (c) do not include the physiologically distinct photorefractory phase separating testicular cycles in starlings under chronic 12L:12D and under temperate-zone photoperiods. While it is possible that testicular cycles in starlings under certain daily light regimens of fixed duration are a function of an endogenous circannual reproductive rhythm, we believe that the testicular cycles generated under both 12L:12D and 6L:18D are the product of gonadotropin secretion rates controlled by circadian (not circannual) oscillations periodically entrained by these chronic photoperiods.     

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.     

Photoperiodic regulation of seasonal responses in Indian weaver bird (Ploceus philippinus)

Photoperiod (=day length) is the vital factor for the regulation of behavioral and physiological activities in many avian species. This study investigated the seasonal cycles of testicular growth and secondary sexual characteristics of Indian weaver bird under natural day length (NDL) and the effects of duration and intensity of light on photoperiodic induction. In the first experiment, groups of birds (n = 7 each) were exposed to under NDL in April 2008 and May 2009 for 8 and 12 months, respectively. In second and third experiment, birds (n = 6 each group) were exposed to different photoperiods (11.5L:12.5D, 12L:12D, 13L:11D, and 15L:9D) at the same (500 lux) light intensity, and to 13L:11D at different light intensities (10-, 50-, 500-, and 800-lux). Observations on testis size, molt, and plumage score were recorded 2-week (molt and plumage) or at 4-week intervals (testes). Both the NDL groups showed similar seasonal cycles of testicular growth-regression and secondary sexual characteristics. Second and third experiments suggest that the photoperiodic induction was depending upon duration and intensity of the light. Birds showed testicular growth-regression cycle followed by molt and plumage color change only under 13L:11D and 15L:9D and only 500- and 800-lux under 13L:11D photoperiod but not under 11.5L:12.5D and 12L:12D and 10- and 50-lux light intensities. Pre- and post-nuptial molting on body feathers were progressed with gonadal stimulation–maturation and regression cycle under 13L:11D and 15L:9D. Results under different light–dark cycles suggest that day length of about 12 h or more and above the threshold level of light intensity are essential for the induction of photoperiodic responses.     

Latitude of origin influences photoperiodic control of reproduction of deer mice (Peromyscus maniculatus)

Three subspecies of Peromyscus maniculatus originating from different latitudes were maintained from birth in light dark cycles that provided between 10 and 18 h of light per day. At 50 days of age, Chihuahua, Mexico mice (latitude of origin 27 degrees N) and South Dakota, U.S.A. mice (44 degrees N) kept in the 10L:14D photoperiod had reduced gonadal and seminal vesicle weights and a lower spermatogenic index than corresponding mice kept in a 14L:10D photoperiod. Some Chihuahua and South Dakota mice, apparently constituting nonphotoperiodic subpopulations, developed their gonads while kept in the short-day photoperiod. The critical day length for stimulation of sexual maturation was greater for mice from Manitoba, Canada (55 degrees N) than for mice from the lower latitudes. At 70 days of age, testes and seminal vesicle weights, and the spermatogenic index of Manitoba mice in the 14L:10D photoperiod, were lower than those of animals maintained in 16L:8D and 18L:6D photoperiods. Responsiveness to short day lengths was greater among adult South Dakota than adult Chihuahau mice and melatonin treatment significantly reduced testes weights of South Dakota but not of Chihuahua adult mice. Photoperiodic regulation of the reproductive system varies with latitude of origin. Differences in the critical day length necessary for stimulating development of functional reproductive activity and variations in the percent of photoperiodic animals within each subspecies, appear to contribute to latitudinal gradients in reproduction.     

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.     

Persistence of a circannual rhythm of plasma prolactin concentrations in ewes exposed to a constant equatorial photoperiod

Circulating concentrations of prolactin were monitored for 3 yr in intact ewes kept either outdoors or indoors in a fixed equatorial photoperiod (12L:12D) and restricted range of environmental temperatures. Prolactin data were analyzed by spectral analysis. In all ewes kept outdoors, concentrations of prolactin showed robust circannual rhythms with a single predominant period of 359 days. In ewes kept indoors, the range of significant periods varied from 35 to 532 days. Although all ewes kept indoors showed a significant rhythm with a period of 354 days, this clearly was not the predominant period in all. The amplitude of the rhythm in ewes kept indoors was significantly lower (p less than 0.01) than that of ewes kept outdoors. Although the annual rhythm of circulating prolactin typical of ewes kept outdoors was significantly compromised in animals kept under a constant 12L:12D photoperiod and restricted environmental temperature range, there was evidence of an endogenous circannual rhythm.     

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.     

The effects of daylength and testosterone on the initiation and progress of moult in Starlings Sturnus vulgaris

The effects of daylength and of testosterone implants, before and after the beginning of moult, on the timing and rate of primary moult have been quantified. Female Starlings Sturnus vulgaris were moved from natural daylength in February to 13 h or 18 h of light per day (13L: 11D or 18L: 6D). Some of the birds on 18L: 6D were left on 18L: 6D throughout the experiment and others were transferred to 13L: 11D after 6 weeks, before moult had begun, or after 12 weeks, after moult had begun. Birds kept on 18L: 6D began to moult before birds kept on 13L: 11D, but the subsequent rate of moult was the same in both groups. A decrease in daylength before moult started slightly advanced the onset of moult. A decrease after moult had begun increased the speed of moult. Castrated male Starlings on 18L: 6D were given testosterone implants for different periods before or after the beginning of moult. Testosterone treatment which ended before moult would normally have started had little effect. Treatment extending beyond the normal start of moult considerably delayed or even prevented the onset of moult. Moult was arrested in birds which received testosterone after moult had begun. On removal of testosterone implants, moult began again from the point where it had stopped, but in some birds, all of the feathers which had been regrown recently were dropped again and regrown. These results are discussed in relation to the different patterns of moult seen amongst different species.     

Persistent free-running circannual reproductive cycles during prolonged exposure to a constant 12L:12D photoperiod in laboratory woodchucks (Marmota monax).

Serum levels of gonadal steroid were assayed at approximately 3-month intervals in groups of 5 to 8 male or female woodchucks which were exposed to a natural photoperiod for 1 year as yearlings or 3 years as adults (Study 1), or a constant photoperiod of 12L:12D from birth for 4.5 years (Study 2). After 4.5 years of 12L:12D, food intake was measured in November and compared with that in natural photoperiod animals (Study 3). Other groups of 11 males and 3 females were housed in 12L:12D for 2.5 years after capture at 2 months of age, and gonadal structure and serum steroid levels in November were compared with those of animals at selected times in the normal annual cycle (Study 4). All animals were provided food and water ad libitum and were not induced to hibernate. In Study 1, normal circannual breeding season elevations in testosterone in males and in progesterone in females were detected in most animals maintained in natural photoperiod. In Study 2, similar cycles persisted for 4.5 years in animals exposed to 12L:12D. However, based on quarterly blood samples, obvious asynchrony relative to natural light animals appeared to develop after 2, 3, or 4 years, with apparent free-running intervals of about 10 to 11 months. In Study 3, mean daily food consumption in late autumn for woodchucks in the 12L:12D group was 72% greater than animals in the natural photoperiod. In Study 4, some woodchucks exposed to 12L:12D for only 2.5 years had prematurely increased spermatogenic activity, Leydig tissue development, and elevated serum testosterone levels in November. They were similar in November to those in natural photoperiod animals in March, and significantly greater than those in natural photoperiod animals in November when normal regression and repair of the testis was complete. Likewise, females in the 12L:12D group had luteinized follicles and elevated progesterone in November which were not noted in natural photoperiod animals and which were similar to those observed during the spring in unbred females under normal conditions. The results suggest that circannual cycles of metabolic and reproductive activity in woodchucks persist in the absence of normal changes in photoperiod, are entrained to seasonal changes in the natural photoperiod, and can recede to a periodicity of less than 12 months within 2.5 to 4 years of laboratory maintenance in 12L:12D.     

Seasonal changes in moult,body mass and reproductive condition in siskins Carduelis spinus exposed to daylength regimes simulating different latitudes

Birds use change in daylength during the year to time events during their annual cycles. Individual Eurasian siskins Carduelis spinus can breed and winter in widely separated areas in different years. Birds at different latitudes will experience different changes in photoperiod. So how does latitude affect photoperiodic control? Our aim in this study was to find whether Siskins caught from the wild in Britain and exposed to different photoperiodic regimes, typical of widely separated latitudes, would differ in the subsequent timing and duration of their moults and associated processes. Siskins were caught in late February and early March, and initially kept outside on natural photoperiods. From the spring equinox (21 March), they were divided into three groups kept under photoperiodic regimes that simulated latitudes 40°, 55° and 70°N respectively. All three groups showed highly significant subsequent changes in body mass, fat scores and cloacal protuberance size. Moult of the primary feathers started during June – August (mean 9 July), and lasted 61–99 days (mean 75 days). Birds that started to moult late in the season had shorter moult durations. All individuals showed lower mass and fat levels during moult than before or after moult. Crucially, there were no significant differences in the timing of these events between the three photoperiodic groups. Apparently these birds did not use prevailing absolute photoperiod or the prevailing rate of change in photoperiod to time moult‐related seasonal events, but used instead some other feature of the annual photoperiod cycle or some form of interval timer linked to photoperiod.     

Photoperiodic control of testicular regression and moult in male House Sparrows Passer domesticus

Testis size, bill colour and moult were monitored in male House Sparrows Passer domesticus kept under a natural daylength regime between February and November. On three occasions (at the summer solstice, 25 days later and 39 days later), groups of birds were transferred to a daylength of 18 h of light and 6 h of darkness per day (18L: 6D), the natural daylength at the solstice. In birds under natural daylengths, the testes had regressed significantly by 2 5 days after the solstice. In those transferred to 18L:6D at the solstice, the onset of regression was delayed by about 4 weeks. Transfer to 18L: 6D after the solstice did not cause recrudescence; the testes continued to regress. In birds transferred to 18L: 6D at the solstice, moult was delayed by 4 weeks and progressed more slowly. These results suggest that photoperiodically induced gonadal regression in this species contains elements characteristic of both absolute and relative photorefractoriness.     

Control of the annual cycle in birds: endocrine constraints and plasticity in response to ecological variability

This paper reviews information from ecological and physiological studies to assess how extrinsic factors can modulate intrinsic physiological processes. The annual cycle of birds is made up of a sequence of life-history stages: breeding, moult and migration. Each stage has evolved to occur at the optimum time and to last for the whole duration of time available. Some species have predictable breeding seasons, others are more flexible and some breed opportunistically in response to unpredictable food availability. Photoperiod is the principal environmental cue used to time each stage, allowing birds to adapt their physiology in advance of predictable environmental changes. Physiological (neuroendocrine and endocrine) plasticity allows non-photoperiodic cues to modulate timing to enable individuals to cope with, and benefit from, short-term environmental variability. Although the timing and duration of the period of full gonadal maturation is principally controlled by photoperiod, non-photoperiodic cues, such as temperature, rainfall or food availability, could potentially modulate the exact time of breeding either by fine-tuning the time of egg-laying within the period of full gonadal maturity or, more fundamentally, by modulating gonadal maturation and/or regression. The timing of gonadal regression affects the time of the start of moult, which in turn may affect the duration of the moult. There are many areas of uncertainty. Future integrated studies are required to assess the scope for flexibility in life-history strategies as this will have a critical bearing on whether birds can adapt sufficiently rapidly to anthropogenic environmental changes, in particular climate change.     

Endogenous control of migration and calendar effects in an intratropical migrant, the yellow-green vireo

Endogenous circannual rhythms control the time course for moult, migratory fattening and autumn migration in juveniles of several bird species that breed in the temperate zone. Exogenous factors, such as daylength, can also exert a measure of control: photoperiodic cues detected by birds that hatch late in the season induce accelerated juvenile development, assuring that late-hatched young migrate at the same time as early-hatched young. Whether these same mechanisms of control also apply to birds that migrate between breeding and nonbreeding areas entirely within the tropics at latitudes characterized by little seasonal variation in photoperiod is unknown. We conducted a common-garden experiment in Panama (9°N) in which we hand-reared wild-caught nestling yellow-green vireos, Vireo flavoviridis, under a constant photoperiod and monitored them for the expression of juvenile moult, migratory fattening and migratory activity. Even in the absence of a seasonal photoperiodic cue, juveniles moulted, accumulated fat reserves and initiated migratory activity, suggesting endogenous control of these processes. Age at the onset of moult, migratory fattening and migratory activity were each significantly negatively correlated with hatch date, however, so that the expression of these three processes was synchronized among juveniles with respect to the time of the season. We suggest that the slight differences in daylength perceived either by the nestlings themselves in the short time before they were collected (at 6-8 days of age) or by the adult females during egg production influenced rate of nestling development, thus allowing later-hatched young to moult, accumulate fat and migrate at a younger age than early-hatched young. Remarkably, photoperiod differed between the earliest and latest hatch dates in this study by only 33 min.     

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.     

Lack of gonadotrophin-releasing hormone (GnRH) neuron response to decreasing photoperiod in thyroidectomized male starlings (Sturnus vulgaris)

Exposure of starlings to long days initially causes reproductive maturation, but eventually leads to photorefractoriness. During photorefractoriness, gonadotrophin-releasing hormone (GnRH) decreases in the GnRH cell bodies and fibers emanating from these to the median eminence, circulating gonadotrophin concentrations decrease to a minimum, and the gonads regress. Thyroidectomy profoundly affects these photoperiodic responses. In chronically thyroidectomized starlings, gonadal responses to changes in day length are attenuated. This investigation was conducted to determine whether, in the absence of gonadal responsiveness, the GnRH system of chronically thyroidectomized starlings responds to changes in day length. Two groups of thyroidectomized male starlings were transferred from short days (8L:16D) to long days (18L:6D) for four weeks, and testicular volume increased. One group was kept on long days (TxLD) and the other was returned to short days (TxSD). Testicular volume did not decrease in the TxSD group. The GnRH neurons of the two thyroidectomized groups were compared to those of two groups of intact starlings, one of them on long days and photorefractory (ILD), the other on short days and photosensitive (ISD). Group ILD had lower numbers of GnRH-stained cells than groups TxLD, TxSD and ISD, which did not differ in this respect. Similar differences were observed for GnRH cell size in the pre-optic area (POA) and for density of staining of GnRH fibers in the median eminence. The results confirm that thyroidectomy attenuates gonadal responses to change in day length and suggest that this results from an effect upon the central nervous system rather than a peripheral effect.