Ovarian steroid withdrawal results in GABAA receptor upregulation in the photoperiodic neuroendocrine pathways of the turkey hen.

The mechanism(s) underlying photorefractoriness in temperate zone seasonally breeding birds remains undetermined. Our recent findings reveal a link between the upregulation of GABAA receptors (GABAARs) in the premammillary nucleus (PMM) and the state of photorefractoriness. Gonadal steroid levels fluctuate during the breeding season; increasing after gonadal recrudescence and declining sharply once gonadal regression begins. Here, we examined the effect of gonadal steroid withdrawal on the expression of GABAARs in the turkey PMM. Exogenous ovarian steroids were administered and then withdrawn from turkey hens to mimic the decline of ovarian steroids levels at the end of a breeding season. The upregulation of GABAAR α3, α4, δ, π, and γ2-subunits was observed in the PMM of the steroid withdrawal group when compared to the non-steroid treatment group. The level of tyrosine hydroxylase, photopigment melanopsin, and circadian clock genes in the PMM of the steroid withdrawal group resembled the levels observed in the natural photorefractory hens and were significantly lower than those of the short-day light stimulated group. A reduction in gonadotropin-releasing hormone-I mRNA expressed within the nucleus commissurae pallii was also observed in hens undergoing steroid withdrawal. These results suggest that the natural decline in circulating ovarian steroid levels may modulate the GABAergic system in the PMM through the upregulation of GABAA receptors. This, in turn, could diminish the reproductive neuroendocrine responses to light and favor a condition resembling the state of photorefractoriness.

Annual gonadal cycles in birds: modeling the effects of photoperiod on seasonal changes in GnRH-1 secretion.

This paper reviews current knowledge of photoperiod control of GnRH-1 secretion and proposes a model in which two processes act together to regulate GnRH1 secretion. Photo-induction controls GnRH1 secretion and is directly related to prevailing photoperiod. Photo-inhibition, a longer term process, acts through GnRH1 synthesis. It progresses each day during daylight hours, but reverses during darkness. Thus, photo-inhibition gradually increases when photoperiods exceed 12h, and reverses under shorter photoperiods. GnRH1 secretion on any particular day is the net result of these two processes acting in tandem. The only difference between species is their sensitivity to photo-inhibition. This can potentially explain differences in timing and duration of breeding seasons between species, why some species become absolutely photorefractory and others relatively photorefractory, why breeding seasons end at the same time at different latitudes within species, and why experimental protocols sometimes produce results that appear counter to what happens naturally.

Enhanced GABAergic inhibition in the premammillary nucleus of photorefractory turkey hens via GABAA receptor upregulation.

The premammillary nucleus (PMM) of the turkey mediobasal hypothalamus, where dopamine-melatonin (DA-Mel) neurons are localized, is a site for photoreception and photoperiodic time measurement, which is essential for the initiation of avian reproductive seasonality. In addition, this area could also be responsible for the onset and maintenance of photorefractoriness at the end of the breeding season due to the enhanced inhibitory effect of γ-aminobutyric acid (GABA). GABA is an inhibitory neurotransmitter in the central nervous system which interferes with the photosexual response in the turkey, a seasonally breeding bird. Here, we further characterized the GABAA receptor subunits in the PMM DA-Mel neurons related to reproductive seasonality and the onset of photorefractoriness. GABAA receptor subunits and GABA synthesis enzymes in the PMM of photosensitive and photorefractory turkey hens were identified using real-time qRT-PCR. The upregulation of GABAA receptor α1-3, ß2-3, γ1-3, ρ1-3, δ, and θ mRNA expression were observed in the PMM of photorefractory birds when compared to those of photosensitive ones while there is no change observed in the GABA synthesis enzymes, glutamate decarboxylase 1 and 2. Those upregulated GABAA receptor subunits were further examined using immunohistochemical staining and they appeared to be co-localized within the PMM DA-Mel neurons. The upregulation of GABAA receptor subunits observed in the PMM of photorefractory birds coincides with a lack of responsiveness to a light stimulus provided during the photosensitive phase. This is supported by the absence of c-fos induction and TH upregulation in the PMM and a subsequence inhibition of c-fos and GnRH-I expression in the nucleus commissurae pallii. The augmented GABAA receptor subunits expression may mediate an enhancement of inhibitory GABAergic neurotransmission and the subsequent interference with the photosexual response. This could contribute to the state of photorefractoriness and the termination of breeding activities in the turkey, a temperate zone bird.

The effect of latitude on photoperiodic control of gonadal maturation, regression and molt in birds.

Photoperiod is the major cue used by birds to time breeding seasons and molt. However, the annual cycle in photoperiod changes with latitude. Within species, for temperate and high latitude species, gonadal maturation and breeding start earlier at lower latitudes but regression and molt both occur at similar times at different latitudes. Earlier gonadal maturation can be explained simply by the fact that considerable maturation occurs before the equinox when photoperiod is longer at lower latitudes - genetic differences between populations are not necessary to explain earlier breeding at lower latitudes. Gonadal regression is caused either by absolute photorefractoriness or, in some species with long breeding seasons, relative photorefractoriness. In either case, the timing of regression and molt cannot be explained by absolute prevailing photoperiod or rate of change in photoperiod - birds appear to be using more subtle cues from the pattern of change in photoperiod. However, there may be no difference between absolute and relative photorefractory species in how they utilise the annual cycle in photoperiod to time regression.

A model of pre-pubertal broiler breeder estradiol-17ß levels predicts advanced sexual maturation for birds with high body weight or short juvenile day-length exposure.

As broiler breeders face increased reproductive challenges specifically related to overfeeding, a clear understanding of the physiological effects of BW and rearing photoperiod on reproductive development is needed. The objective was to use mathematical models to compare plasma estradiol-17ß (E2) concentration to characterize the effect of BW and rearing photoperiod on E2 levels. A 2 × 3 factorial arrangement of treatments was used. Hens (n = 180) were fed with a precision feeding system to allocate feed individually to achieve the breeder-recommended BW curve (Standard) or to a BW curve reaching the 21 wk target at 18 wk (High). Hens were on 8L:16D, 10L:14D, or 12L:12D photoschedules during rearing and were photostimulated at 21 wk. Age at first egg (AFE) was recorded. Plasma E2 levels were determined weekly between week 20 and 28. Two modified Gompertz models described E2 level as a function of (a) chronological or (b) physiological (relative to AFE) age. Timing of E2-inflection point was compared between models and treatments. Differences were reported as significant at P ≤ 0.05. The chronological age model inferred that High BW reduced the duration between the E2-inflection point and AFE, whereas the physiological age model inferred that High BW only reduced the duration between photostimulation and the E2-inflection point. Hens on the Standard BW treatment had a longer period between photostimulation and the E2-inflection point compared to hens on the High-BW treatment (11.03 vs. 1.50 wk, respectively, based on physiological age). Hens on the 12L:12D photoschedule had a longer period between photostimulation and the E2-inflection point compared to hens on the 8L:16D or 10L:14D photoschedule, both in the Standard and High BW (28.91 vs. 1.78 and 2.40 wk, 2.65 vs. 0.93 and 0.94 wk, respectively, based on physiological age). The described methodology and results provide quantitative insight into E2 dynamics and serves as a model for future endocrinological studies in poultry reproduction.

Dorsomedial hypothalamic lesions counteract decreases in locomotor activity in male Syrian hamsters transferred from long to short day lengths.

The dorsomedial nucleus (DMN) of the hypothalamus has been implicated in seasonal control of reproduction. Syrian hamsters with DMN lesions, unlike control hamsters, do not undergo testicular regression after transfer from a long day length (14 h of light per day; LD) to a short day length (8 h of light per day; SD). SDs also markedly reduce hamster locomotor activity (LMA). To assess whether the DMN is a component of the neural circuitry that mediates seasonal variation in LMA, neurologically intact males (controls) and hamsters that had sustained lesions of the DMN (DMNx) were housed in an LD or SD photoperiod for 26 weeks. DMNx that prevented testicular regression counteracted decreases in LMA during 8 to10 weeks of SD treatment; steroid-independent effects of SDs did not override high levels of LMA in DMNx males. As in previous studies, testosterone (T) restoration increased LMA in LD but not SD castrated control males. In the present study, T also failed to increase LMA in SD-DMNx hamsters. The DMN is not necessary to maintain decreased responsiveness of locomotor activity systems to T in SDs, which presumably is mediated by other central nervous system androgen target tissues. Finally, DMNx did not interfere with the spontaneous increase in LMA exhibited by photorefractory hamsters after 26 weeks of SD treatment. We propose that DMN is an essential part of the substrate that mediates seasonal decreases in LMA as day length decreases but is not required to sustain decreased SD responsiveness to T or for development of refractoriness to SDs.

Photoperiodic regulation of seasonal reproduction, molt and body weight in the migratory male yellow-breasted bunting (Emberiza aureola).

Photoperiod has been shown to be a major source of temporal information regulating reproduction and associated functions in a number of avian species. We studied seasonal cycles of testicular volume, molt and body weight in natural and temperature-controlled conditions and under different artificial photoperiods in the yellow-breasted buntings. Buntings posses seasonal cycles of testicular volume, molt, body weight and fattening with no major difference between natural and temperature-controlled conditions. These cycles follow an annual solar cycle suggesting the possibility of their photoperiodic control. To confirm this, photosensitive birds were studied under 9L/15D (close to shortest day length), 12L/12D (equinox day length) and 14L/10D (close to longest day length) for 18 months. Buntings showed testicular growth followed by regression and development of photorefractoriness; molt and body weight change only under 12L/12D and 14L/10D but not under 9L/15D. Reinitiation of above responses did not occur following initial cycles under stimulatory photoperiods precluding the possibility of circannual rhythm involvement. Birds exhibited an incomplete prenuptial molt of body feathers during gonadal stimulation under long days followed by complete postnuptial molt of body and primary feathers that progressed with gonadal regression. Exposure of photosensitive birds to light-dark cycles constituting 9-16h of light/day suggested that daily photoperiod of about 12h or more is essential in inducing testicular growth and function. These results clearly indicate that buntings are capable of fine discrimination of photoperiodic information and use annual changes in day length as an environmental factor to time their seasonal responses.