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.     

Sexual maturation and moult in juvenile Starlings Sturnus vulgaris in response to different daylengths

Starlings, like most other species, show no gonadal development until spring of the year after they hatch, even though they hatch and attain full body size during long days. This could be because they develop in a physiological state analogous to that of photorefractory adults and so need to experience short days in order to activate the reproductive system. To test this possibility, young were hand-reared under different photoperiodic regimes. Young raised under constant long days showed no gonadal development, nor did birds initially raised under short days and then transferred to long days at 3 weeks of age. However, birds transferred from short to long days at 10 weeks of age did show gonadal development, followed by gonadal regression, while birds raised under constant short days showed slow continual gonadal development. This last group, unlike the other three groups, did not moult into adult plumage. Since 4 weeks of long days are required to terminate photorefractoriness in adult Starlings, these results demonstrate that the reproductive system of young birds is in a similar state to that of photorefractory adults, and hence that puberty is analogous to the termination of photofractoriness.     

The pineal clock affects behavioral circadian rhythms but not photoperiodic induction in the Indian weaver bird (Ploceus philippinus)

We investigated whether pineal is part of the circadian clock system which regulates circadian rhythms of activity and photosensitivity in the Indian weaver bird (Ploceus philippinus). Two experiments were performed. The first experiment examined the induction of testicular growth, and androgen-dependent beak pigmentation and luteinizing hormone (LH)-specific plumage coloration in pinealectomised (pinx) and sham-operated (sham) birds exposed to short day (8 h light: 16 h darkness, 8L:16D) and long day (16L:8D) for 9 months in the late breeding and early regressive phase (October), or the late regressive and preparatory phase (January) of the annual testicular cycle. As expected, short days were non-stimulatory, and long days stimulated testicular growth, beak pigmentation and plumage coloration. There was no difference in the response between pinx and sham birds subjected to short or long days in October, but the response was enhanced in pinx birds that were subjected to long day in January. In the second experiment circadian behavioral rhythms were studied (activity pattern in singly housed birds) in weaver birds first exposed at two different phases of the annual testicular cycle to a stimulatory photoperiod (12L:12D in preparatory phase or 13L:11D in early breeding phase) and then released into dim continuous light (LL_dim). All birds showed synchronization to the light period before and after the pinealectomy; there was no difference in the response between pinx and sham birds. When released into LL_dim, sham birds exhibited circadian rhythmicity continuously, whereas pinx birds lost circadian rhythmicity after some cycles. Considered together, these results suggest that circadian clock residing within the pineal gland regulates the circadian rhythm in activity, but not the circadian rhythm involved in photoperiodic induction of the Indian weaver bird.     

Effect of season and photoperiod on the follicle-stimulating hormone receptors in a subtropical bird

Annual changes in and photoperiodic influence oh the weight of gonads, a parameter of gonadal activity, are much smaller in female birds than in males. Effect of season and photoperiod on the follicle-stimulating hormone receptors in the testis or ovary was studied using a subtropical weaver finch. The number of follicle-stimulating hormone binding sites per unit testicular weight showed a peak in the non-breeding phase; while the total number of binding sites per two testes was maximal in the breeding phase and minimal in the regressive phase. In contrast, seasonal changes in follicle-stimulating hormone binding sites in the ovary were less marked. Exposure to short-day during the breeding phase induced marked decreases in the numbers of binding sites per unit testicular weight and per two testes. These numbers markedly increased after transfer to long-day during the non-breeding phase. However, there was no significant effect of short-day or long-day exposure on follicle-stimulating hormone binding sites in the ovary. These results suggest that photoperiod is an effective environmental factor in the regulation of follicle-stimulating hormone receptors in the testis and the effect is manifested by pronounced changes in the testicular weight during annual breeding cycle.     

Changes in the daily rhythm of plasma corticosterone concentration related to seasonal conditions in the white-throated sparrow, Zonotrichia albicollis.

Circadian variations in concentrations of plasma corticosterone were investigated in the white-throated sparrow maintained on short (10-hr) or long (16-hr) daily photoperiods. In addition, the plasma concentrations of corticosterone were determined throughout a day in birds that were in the reproductively photosensitive spring migratory condition, the reproductively photorefractory post nuptial molt condition, and the fall migratory condition. Distinct unimodal rhythms were found in photosensitive birds. The daily rise occurred 12 hr after the offset of light in birds kept on both the short and the long photoperiodic regimens. There was no discernible daily variation in photorefractory birds kept on a 16 hr daily photoperiod and there was a bimodal rhythm in the birds that were in the fall migratory condition. The results are consistent with an hypothesis that assigns an important role to the circadian rhythm of corticosteroid concentration in the photoperiodic mechanism controlling seasonal reproductive and migratory conditions in the white-throated sparrow.     

Importance of photoperiods in determining temporal pattern of annual testicular events in rose-ringed parakeet (Psittacula krameri).

Male rose-ringed parakeets (Psittacula krameri) were transferred to a long photoperiod (LP; LD 16:8) or a short photoperiod (SP; LD 8:16) for 45 or 90 days on four dates corresponding to the beginnings of different reproductive phases in an annual testicular cycle, and testicular responsiveness was evaluated by comparison with the testicular volume, weight, seminiferous tubular diameter, and germ cell profiles of birds in a natural photoperiod (NP). Exposure of birds to LP during the progressive phase (November) led to precocious maturation of testes after 45 days, but induced regression at 90 days. After showing retarded gametogenic functions at 45 days, parallel (November) SP birds exhibited an accelerated rate of germ cell formation at day 90. During the prebreeding phase (January), there were no remarkable differences in any features of testes among NP. LP, and SP birds at 45 days, but gonadal involution in LP parakeets and active spermatogenesis in SP birds occurred after 90 days. The testes did not show any response to LP or SP for 45 and 90 days when the birds were transferred to altered photoperiods during the breeding (March) and preparatory (June) phases, indicating that the parakeets were photorefractory for at least 6 months (March through September). The results also suggest that initiation and termination of seasonal gametogenic activity in parakeets are possibly functions of endogenous rhythmicity or extraphotoperiodic environmental factors. Duration of light may have certain influences on the attainment of annual peak in spermatogenesis, but in all probability the species has a low photoperiod threshold for induction of testicular growth.     

The seasonal development of photoperiodic responsiveness in an equatorial migrant, the garden warblerSylvia borin

Summary Garden warblers (Sylvia borin) are migrating European passerines that spend the winter in tropical Africa. To investigate how photoperiod controls the annual cycles of reproduction, molt and migration, garden warblers were exposed to photoperiodic changes simulating those experienced by conspecifics with an equatorial wintering area. At three different times groups of birds were moved from a constant equatorial photoperiod of 12.8 h to a 15-h photoperiod: group 1 on November 19, group 2 on February 24, and group 3 on April 12. In all birds the changes in body weight, nocturnal migratory restlessness, testicular width or diameter of the largest oocyte, and plasma LH concentrations were determined. In addition, the birds were examined for molt. In all birds of groups 2 and 3 the transfer to long photoperiods induced a gonadal cycle associated with spring migratory restlessness and, in the group 3 birds, an increase in LH. In group 1 only few birds showed marginal gonadal responses and the gonadal cycles of the group 2 birds were less pronounced than those of group 3. The results suggest that the hypothalamo-pituitary-gonadal axis was refractory to stimulation by long photoperiods in autumn but, later in the season, became responsive to the same long photoperiods. Unlike the situation in other temperate zone birds, the transition from the photorefractory to the photosensitive state seems to be a gradual process which develops spontaneously and with the appropriate temporal characteristics in the absence of photoperiodic change.Abbreviation LH luteinizing hormone     

Elimination of short-day melatonin signaling accelerates gonadal recrudescence but does not break refractoriness in male Turkish hamsters

Long days stimulate and short days (SDs) inhibit the reproductive axis of photoperiodic rodents. In long-day Turkish hamsters, unlike most other rodents, elimination of pineal melatonin secretion by constant light or pinealectomy initiates a cycle of gonadal involution and recrudescence outwardly similar to that induced by short days. The present study assessed whether short days and constant light induce the seasonal reproductive cycle via common or different interval timing mechanisms. Male hamsters that had undergone gonadal involution in SDs for 8 or 14 weeks were treated with LL for 14 and 8 weeks, respectively. If SDs and LL act via independent mechanisms, then gonadal quiescence of SD-regressed males, which normally lasts 10 weeks, might be extended by LL treatment; alternatively, if SDs and LL act on the same timer, or the timer cannot be retriggered, then LL will not extend the duration of reproductive quiescence. Neither of these outcomes materialized. Instead, male hamsters exposed to LL while reproductively quiescent exhibited accelerated gonadal recrudescence. Extended LL treatment did not restore responsiveness to SDs in photorefractory hamsters. In Turkish hamsters, photoperiodic history determines whether constant light inhibits or stimulates the hypothalamic-pituitary-testicular axis.     

The role of the pineal gland in the photoperiodic control of bird song frequency and repertoire in the house sparrow, Passer domesticus

Temperate zone birds are highly seasonal in many aspects of their physiology. In mammals, but not in birds, the pineal gland is an important component regulating seasonal patterns of primary gonadal functions. Pineal melatonin in birds instead affects seasonal changes in brain song control structures, suggesting the pineal gland regulates seasonal song behavior. The present study tests the hypothesis that the pineal gland transduces photoperiodic information to the control of seasonal song behavior to synchronize this important behavior to the appropriate phenology. House sparrows, Passer domesticus, expressed a rich array of vocalizations ranging from calls to multisyllabic songs and motifs of songs that varied under a regimen of different photoperiodic conditions that were simulated at different times of year. Control (SHAM) birds exhibited increases in song behavior when they were experimentally transferred from short days, simulating winter, to equinoctial and long days, simulating summer, and decreased vocalization when they were transferred back to short days. When maintained in long days for longer periods, the birds became reproductively photorefractory as measured by the yellowing of the birds' bills; however, song behavior persisted in the SHAM birds, suggesting a dissociation of reproduction from the song functions. Pinealectomized (PINX) birds expressed larger, more rapid increases in daily vocal rate and song repertoire size than did the SHAM birds during the long summer days. These increases gradually declined upon the extension of the long days and did not respond to the transfer to short days as was observed in the SHAM birds, suggesting that the pineal gland conveys photoperiodic information to the vocal control system, which in turn regulates song behavior.     

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.     

Diapause and seasonal life cycle strategy in the house spider, Achaearanea tepidariorum (Araneae, Theridiidae)

Abstract In order to elucidate the mechanism regulating its seasonal life cycle, the photoperiodic response of Achaearanea tepidariorum has been analysed. Nymphal development was faster in long-day and slower in short-day photoperiods. The combined action of low temperature, poor food supply and short daylength induced diapause at an earlier developmental stage than short days alone. Thus, photoperiod is a primary factor inducing nymphal diapause, but the diapausing instar is influenced by both temperature and food supply. Hibernating nymphs became unresponsive to photoperiod in late December. After hibernation, however, sensitivity was restored and the nymphs remained sensitive to photoperiod throughout their life. This spider could also enter an imaginal or reproductive diapause. Photoperiod was again a primary inducing factor and temperature modified the photoperiodic response to some extent. The induction of the reproductive diapause was almost temperature-compensated whereas development was not. So the involvement of a photoperiodic counter system was suggested. Irrespective of whether the nymph had experienced diapause or not, the imaginal diapause was induced in response to a short-day photoperiod after adult moult. Based on these observations, the seasonal life cycle and the adaptive significance of nymphal and imaginal diapause are discussed.     

Responsiveness to photoperiodic changes in the circannual rhythm of the varied carpet beetle, <Emphasis Type="Italic">Anthrenus verbasci</Emphasis>

The circannual pupation rhythm of Anthrenus verbasci is entrained to an environmental cycle by changes in photoperiod. Exposure of larvae reared under short-day conditions to 4 weeks of long days can induce phase-dependent phase shifts. In the present study, we examined the range of photoperiodic changes effective for phase shifts at 20°C. For larvae under light/dark (LD) 12:12 conditions, 4-week exposure to LD 14:10 caused a clear phase delay, as great an extent as that brought about by exposure to LD 15:9 and LD 16:8. In contrast, the delay brought about by exposure to LD 13:9 was slight. For larvae under LD 10:14, exposure to LD 14:10 and LD 16:8 for 4 weeks induced a phase delay, but exposure to LD 12:12 did not. These results indicate that a clear phase delay is induced when the photoperiodic change exceeds a critical value in the photophase between 13 and 14 h, regardless of the amplitude of the change. Although phase advances were smaller than phase delays, they depended on the amplitude of photoperiodic changes rather than the absolute photophase duration in contrast to the case of the phase delay.     

Mutation in TERMINAL FLOWER1 reverses the photoperiodic requirement for flowering in the wild strawberry Fragaria vesca

Photoperiodic flowering has been extensively studied in the annual short-day and long-day plants rice (Oryza sativa) and Arabidopsis (Arabidopsis thaliana), whereas less is known about the control of flowering in perennials. In the perennial wild strawberry, Fragaria vesca (Rosaceae), short-day and perpetual flowering long-day accessions occur. Genetic analyses showed that differences in their flowering responses are caused by a single gene, SEASONAL FLOWERING LOCUS, which may encode the F. vesca homolog of TERMINAL FLOWER1 (FvTFL1). We show through high-resolution mapping and transgenic approaches that FvTFL1 is the basis of this change in flowering behavior and demonstrate that FvTFL1 acts as a photoperiodically regulated repressor. In short-day F. vesca, long photoperiods activate FvTFL1 mRNA expression and short days suppress it, promoting flower induction. These seasonal cycles in FvTFL1 mRNA level confer seasonal cycling of vegetative and reproductive development. Mutations in FvTFL1 prevent long-day suppression of flowering, and the early flowering that then occurs under long days is dependent on the F. vesca homolog of FLOWERING LOCUS T. This photoperiodic response mechanism differs from those described in model annual plants. We suggest that this mechanism controls flowering within the perennial growth cycle in F. vesca and demonstrate that a change in a single gene reverses the photoperiodic requirements for flowering.     

Timing of breeding in variable environments: tropical birds as model systems

Animals need to adjust reproductive decisions to environmental seasonality. In contrast to species from the well-studied temperate zones, little is known for tropical birds about the environmental cues that stimulate reproductive activity and the physiological mechanisms that regulate reproduction. I am investigating the environmental and endocrine mechanisms that underlie the timing of reproduction in spotted antbirds from the near-equatorial Panamanian rainforest and in small ground finches from the equatorial arid Galápagos islands. Spotted antbirds live in a fairly predictable seasonal environment and show regular changes in gonad sizes and some reproductive hormones. Despite the small annual variation in photoperiod close to the equator, these birds can measure slight photoperiodic increases and use it to initiate reproductive activity. Spotted antbirds also respond to seasonal changes in food availability, which allows them to flexibly adjust gonad growth to environmental conditions. Testosterone is involved in regulating song and aggressive behavior in these year-round territorial birds, although it can remain at low plasma levels throughout the year. In contrast, small ground finches exposed to a rather unpredictable climate on Galápagos appear to grow their gonads whenever heavy rains fall and have regressed gonads during other times of the year. The lack of a physiological preparation for the breeding season and their response to short-term cues related to rainfall indicate a striking flexibility in the regulation of breeding in small ground finches. I suggest that tropical birds can serve as model systems to study the physiological adaptations to different environments. Unraveling the neuroendocrine mechanisms behind the flexibility in reproductive timing will clarify whether differences found between temperate and tropical birds represent variations of the same basic mechanism or instead reflect a fundamental divergence in physiological control systems.     

Photoreceptors and signals in the photoperiodic control of development

Many plant species are sensitive to changes in the seasons, especially with regard to their reproductive behaviour. Sexual or vegetative reproductive structures are often only formed at times of the year when days are sufficiently long, or short. Plants perceive daylength in the leaves, but reproductive changes occur in shoot apices in response to the movement of signals throughout the plant. There is good evidence that phytochrome A is an essential component of the daylength-sensing mechanism in long-day plants, and preliminary evidence suggests a potential interaction between phytochrome C and daylength sensitivity in short-day plants. The sensitivity of reproductive processes to photoperiodic control is directly altered by photoreceptor action, particularly in the case of phytochrome B, which has a major influence on whether flowering or tuberization occurs under non-inductive conditions in both long- and short-day plants, but is not involved in daylength measurement. The signals which move from leaves to the sites of reproductive development are not known, but there is good evidence that gibberellins may be important and some preliminary indication that brassinosteroids might also be involved in photoperiodic signalling.     

Effect of differential photostimulation on induction of Fos-like immune-reactivity in the MBH region of Indian weaver bird

This study investigated the neuronal activation (c-fos) in the mediobasal hypothalamus (MBH) of the Indian weaver bird, after exposure to a single long-day. Wild-caught photosensitive birds were exposed to the short-days (LD 8L:16D). After four days of short-days photoperiod, the LD cycle was programmed such that the light illumination was continuous after zeitgeber time (ZT) 8. Birds were perfused on the same day at ZT 4 or ZT 20. Brains were processed for the immunohistochemistry of c-fos (Fos) in the MBH. We found a significant higher activation of Fos in neurons within the ventral tuberal division (containing infundibular nucleus) of the MBH, in group that received 20-h light than that received 4-h light on first long-day. But in the dorsal tuberal division, there was no noticeable difference in Fos-lir activation on after 4-h and 24-h light exposure. The results suggest that the ventral tuberal division of mediobasal hypothalamus is principally involved in detecting the photoperiodic information from the external environment, and hence, suggested as the key neural center, involved in the photoperiodic mechanism in Indian weaver bird.     

Nest survival rates of Red‐crested Cardinals increase with nest age in south‐temperate forests of Argentina

ABSTRACT The main cause of nest mortality for most bird species is predation and nest survival rates often vary in relation to time‐specific variables. Few investigators have examined time‐specific patterns of nest survival in Neotropical birds, and most such studies have focused on tropical and subtropical species. To better understand age‐related patterns of nest survival, we studied nest survival of Red‐crested Cardinals (Paroaria coronata, Thraupidae) in a south‐temperate forest in Argentina. We modeled daily nest survival rates (DSR) using program MARK. We examined the relationship between nest age and nest survival rate, controlling for the effects of physical characteristics of nest sites and progression of the breeding season. We monitored 367 nests for a total of 4018 exposure days. We found that DSR increased with nest age and was higher in small isolated patches than in large continuous patches of forests. The increase of DSR with nest age could be a consequence of more vulnerable nests being predated early in the nesting cycle or a result of parents defending nests more vigorously as nestlings age because of their increasing reproductive value. Open areas of grassland that surrounded the small isolated patches of forests in our study may have been a barrier to predator movements, possibly explaining the lower predation rates. Nest survival rates in our study were lower than those reported for tropical or Nearctic temperate birds, but similar to those reported in other studies of Neotropical temperate birds. Reasons for the low nest survival rates of Neotropical temperate birds remain unclear, and additional studies of predator communities are needed to help elucidate this topic.     

Photoperiod affects reproductive responsiveness to 6-methoxy-2-benzoxazolinone in house mice

House mice (Mus musculus) have traditionally been characterized as nonphotoperiodic because reproductive function is unaffected by day length in the laboratory. In the present study, the reproductive responsiveness of CF1 mice to a naturally occurring plant metabolite was assessed in animals that were maintained in either long (16L:8D) or short (8L:16D) photoperiods from birth until the end of the study. Males and females were implanted i.p. with either an empty Silastic capsule or one containing 6-methoxy-2-benzoxazolinone (6-MBOA) at either 21 or 60 days of age for 3 days. Other 31-day-old mice were implanted with capsules for 8 wk. Three-day exposure to 6-MBOA stimulated uterine growth in short-day, adult females, but did not affect adult females housed in long photoperiods. Short-term treatment with 6-MBOA did not significantly affect reproductive parameters in either long- or short-day peripubertal house mice, or in adult males regardless of photoperiod, nor did exposure to 6-MBOA for 8 wk influence reproduction in males in either photoperiodic condition. However, short-day female mice had significantly reduced ovarian and uterine masses after 8 wk chronic 6-MBOA treatment as compared to long-day animals or mice unexposed to 6-MBOA. Short-day females exposed to 6-MBOA for 8 wk developed a denser pelage compared to long-day mice treated with this compound. Photoperiod-mediated differential responsiveness to 6-MBOA indicates that female house mice can discriminate long from short days, and these results suggest that the physiological mechanisms for photoperiodic responsiveness remain extant in this species previously characterized as nonphotperiodic.