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F E Wilson

Publications and source records attributed to F E Wilson.

At least 19 recordsLinked to original sources

Thyroid hormone-dependent seasonality in American tree sparrows (Spizella arborea): effects of GC-1, a thyroid receptor beta-selective agonist, and of iopanoic acid, a deiodinase inhibitor.

To explore the role of TH in the control of seasonality [i.e., photoperiodic testicular growth, photorefractoriness, and postnuptial (prebasic) molt] in American tree sparrows (Spizella arborea), we performed experiments in which THX males were simultaneously photostimulated and given TH replacement therapy. In the first experiment, equimolar concentrations (1X = 1.3 nmol) of T4, T3, or GC-1, an iodine-free TRbeta agonist, were administered s.c. daily during the first 21 days of photostimulation. Two additional THX groups received GC-1 at 0.1X or 10X, and THX and THI control groups received vehicle. In the second experiment, T4 or T3, alone or in combination with the deiodinase inhibitor IOP, was injected i.m. twice daily during the first 14 days of photostimulation. In both experiments, end points were testis length and molt score. In the first experiment, THI birds given vehicle and THX birds given T4 replacement therapy exhibited all three components of seasonality. THX birds given T3 or GC-1 (1X or 10X) showed a subdued photoperiodic testicular response, but they did not become photorefractory or initiate molt. THX birds that received 0.1X GC-1 or vehicle exhibited none of the components of seasonality. These data are consistent with the hypothesis that photoperiodic testicular growth, a vernal component of seasonality, is a TRbeta-mediated response and suggest that T4 may activate TRbeta more efficiently than does T3 or GC-1. By contrast, the failure both of T3 and of GC-1, but not of T4, to program photostimulated THX males for photorefractoriness and postnuptial molt suggests that autumnal components of seasonality may be TRalpha-mediated responses solely to T4. In the second experiment, IOP administered alone had no significant impact on seasonality. THX birds that received T4 with or without IOP showed all components of seasonality, whereas birds that received T3 with or without IOP showed only photoperiodic testicular growth. These results challenge the widely held view that T4 is merely a prohormone for T3 and support the emerging view that T4 has intrinsic hormonal activity. Because IOP augmented the photoperiodic testicular response in T3-treated THX birds, T3 may act either independently or co-dependently with T4 in programming vernal seasonal events.

Acclimatization↗

A test of the hypothesis that T3 is the "seasonality" thyroid hormone in American tree sparrows (Spizella arborea): intracerebroventricular infusion of iopanoic acid, an inhibitor of T3 synthesis and degradation.

This study tested the hypothesis that L-3,5,3'-triiodothyronine (T3) is the bioactive "seasonality" thyroid hormone in American tree sparrows (Spizella (arborea). The experimental approach coupled thyroid hormone replacement therapy after radiothyroidectomy with photostimulation and intracerebroventricular infusion of iopanoic acid, an inhibitor of L-3,5,3'-triiodothyronine synthesis and degradation. Endpoints were testis length, molt score, and hypothalamic content of chicken gonadotropin-releasing hormone 1. The hypothesis predicts that thyroidectomized male tree sparrows moved to long days and given thyroxine in combination with iopanoic acid will lack L-3,5,3'-triiodothyronine and so will not express thyroid hormone-dependent photoperiodic testicular growth (a vernal component of seasonality) and photorefractoriness or postnuptial molt (autumnal components of seasonality). It further predicts that iopanoic acid will enhance the efficacy of L-3,5,3'-triiodothyronine and so will facilitate the expression of seasonality in thyroidectomized males given L-3,5,3'-triiodothyronine replacement therapy. Iopanoic acid had no significant effect on any component of seasonality in thyroid-intact males given vehicle, or in thyroidectomized males given thyroxine or L-3,5,3'-triiodothyronine. Thyroid-intact males, as well as thyroidectomized males infused with thyroxine alone, commonly expressed all components of seasonality. Thyroidectomized males given L-3,5,3'-triiodothyronine alone exhibited photoperiodic testicular growth, but did not become photorefractory or initiate molt. While these results confirm that thyroid hormone acts centrally to program American tree sparrows for seasonality, they do not support the hypothesis that L-3,5,3'-triiodothyronine is the bioactive "seasonality" thyroid hormone, and they challenge the view that thyroxine is merely a prohormone.

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Thyroid hormone acts centrally to programme photostimulated male american tree sparrows (Spizella arborea) for vernal and autumnal components of seasonality.

Thyroid hormone and long days interact to programme American tree sparrows (Spizella arborea) for seasonality (i.e. thyroid hormone-dependent photoperiodic gonadal growth, photorefractoriness, and postnuptial moult). This study explored in radiothyroidectomized (THX) males given thyroid hormone replacement therapy whether thyroid hormone acts within the brain and, additionally, the identity of the putative tissue-active thyroid hormone. The minimum dose (30 ng) of L-thyroxine (T4) that restored all components of seasonality when given i.c.v. daily during the first 21 days of photostimulation restored no component of seasonality when given s.c. The same dose of L-triiodothyronine (T3) also was ineffective when administered s.c., but restored photoperiodic testicular growth (though neither photorefractoriness nor postnuptial moult) when admiministered i.c.v. Three of seven birds given a 10-fold lower dose of T4 (3 ng) exhibited thyroid hormone-dependent photoperiodic testicular growth, albeit damped. The other four birds given 3 ng T4 and all birds given 3 ng T3 responded like THX controls, exhibiting only slight thyroid hormone-independent photoperiodic testicular growth. The highest dose (300 ng) of T3 restored all components of seasonality only when administered i.c.v. daily during the first 49 days of photostimulation. This demonstration in American tree sparrows is the first in any species that the thyroid-dependent transition from the breeding season to the non-breeding season can be effected by T3. The same dose of reverse T3 administered daily over the same 49 days restored photoperiodic testicular growth in only half of 10 subjects and photorefractoriness and moult in none. Collectively, the data support the hypothesis that thyroid hormone acts centrally to programme photostimulated male American tree sparrows for all components of seasonality. The most parsimonious interpretation of the data, including the threshold-like effect of 3 ng T4, favours T4 as the tissue-active thyroid hormone for vernal as well as autumnal events, but does not entirely exclude T3.

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Long days and thyroxine program american tree sparrows for seasonality: evidence for temporal flexibility of the breeding season of euthyroid females.

To explore the role of the thyroid in the control of seasonality, photosensitive female American tree sparrows (Spizella arborea) were thyroidectomized (THX), moved to long days, and given daily injections of thyroxine (T4) for 3 weeks; THX and thyroid-intact (THI) controls received daily injections of alkaline vehicle. Birds were retained on long days 4 additional weeks and then moved to constant light and given T4 in drinking water for 5 weeks in order to test for photorefractoriness. Endpoints were ovarian mass, molt score, and hypothalamic cGnRH-I (chicken gonadotropin-releasing hormone I) content; data were collected as independent measures at intervals of 1 to 5 weeks. THX females given T4 replacement therapy (THXT4 females) exhibited all components of seasonality (i.e., photoperiodic ovarian growth, photorefractoriness, and postnuptial molt), as did THI females. THX females not given replacement T4 were aseasonal: They showed only minor thyroid-independent photoperiodic ovarian growth; they remained photosensitive, despite chronic photostimulation; and they did not initiate postnuptial molt. Collectively, these observations support, and extend to female tree sparrows, our model of seasonality in male tree sparrows, wherein control circuits are programmed for gonadal growth, photorefractoriness, and postnuptial molt by interactive effects of long days and thyroid hormone during the first 3 weeks of photostimulation. The unexpected finding that constant light and/or exogenous T4 extended the simulated breeding seasons of some THI females prompted us to investigate the nature and expression of photorefractoriness. Our approach was to evaluate the same endpoints as before in chronically photostimulated THI females either retained on long days, with or without T4 in drinking water, or moved to constant light, with or without T4. The results showed that exposure to constant light (not T4) near the end of a simulated breeding season can-though usually it does not-temporarily extend the breeding season and prevent the onset of postnuptial molt. It remains unclear whether these perturbations caused by constant light reflect a transitional relative photorefractoriness or merely a delay in the onset of absolute photorefractoriness. In either case, the thyroid- and daylength-dependent programs controlling seasonality in female American tree sparrows have a heretofore undemonstrated potential for limited temporal flexibility.

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Time of thyroidectomy variably affects seasonality in female American tree sparrows (Spizella arborea).

Female American tree sparrows (Spizella arborea) were injected with Na131I before, at, or after the onset of photostimulation in order to study the effect of time of thyroidectomy on three components of seasonality: thyroid-dependent photoperiodic ovarian growth, photorefractoriness, and postnuptial (prebasic) molt. Thyroidectomy before or at the onset of photostimulation abolished all components of seasonality; birds exhibited only minor thyroid-independent photoperiodic ovarian growth. Thyroidectomy on day 7 of photostimulation blocked the transition from photosensitivity to photorefractoriness; although birds showed thyroid-dependent photoperiodic ovarian growth, they neither exhibited ovarian regression nor initiated postnuptial molt. Thyroidectomy on day 14, 21, or 28 of photostimulation had no remarkable effect on any component of seasonality. We conclude that separate mechanisms control photoperiodic ovarian growth and photorefractoriness/molt and that, early during photostimulation, the thyroid has a codependent role in programming female tree sparrows for vernal as well as autumnal seasonal events.

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Effect of withdrawing long days from male American tree sparrows (Spizella arborea): implications for understanding thyroid-dependent programming of seasonal reproduction and postnuptial molt.

In previous studies, we withdrew thyroid hormones by thyroidectomy before, at, or after the onset of photostimulation and showed that male American tree sparrows (Spizella arborea) are programmed for seasonal reproduction and postnuptial molt by or before Week 3 on long days. In this corollary study, we withdrew long days before or after the control circuits had been programmed. After 1 day to 4 wk on long days, groups of thyroid-intact males were returned to short days until Week 7, when they were moved to constant light and evaluated for photosensitivity or photorefractoriness and postnuptial molt. Long-day controls held 7 wk on long days showed robust testicular growth through Week 6 and then spontaneous testicular regression. Testes of short-day controls and of males photostimulated for 1 day remained small. In all other groups, photostimulation induced testicular growth, which gave way to regression during exposure to short days. Long-day controls tested photorefractory at Week 7 and initiated molt by Week 10. All other groups tested photosensitive and did not molt. Our demonstration that long days are required for expression of seasonal reproduction and postnuptial molt in thyroid-intact male tree sparrows previously programmed for these events suggests that long days create a milieu that is permissive for expression.

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Effects of thyroxine (T4) or triiodothyronine (T3) replacement therapy on the programming of seasonal reproduction and postnuptial molt in thyroidectomized male American tree sparrows (Spizella arborea) exposed to long days.

This study tested the hypothesis that T3 (triiodothyronine) is the tissue-active "seasonality" hormone by determining whether T3 could mimic T4 (thyroxine) and program photostimulated thyroidectomized (THX) male American tree sparrows (Spizella arborea) for three components of seasonality (i.e., full-blown testicular growth, photorefractoriness, and postnuptial molt). Photosensitive males were radiothyroidectomized, transferred to long days 4 weeks later, and administered 14 daily injections (s.c.) of alkaline saline (V) containing 0.1, 1, or 10 micrograms T4 or T3. THX and thyroid-intact (THI) controls received only V. After 5 additional weeks on long days, all birds were tested for photosensitivity/photorefractoriness. Periodically during the experiment, primary flight feathers were scored for molt, and testis length was monitored by laparotomy. As an independent measure of reproductive (i.e., photosensitive vs. photorefractory) state, hypothalami collected at the end of the experiment were assayed for cGnRH-I (chicken gonadotropin-releasing hormone I) content. Like THI controls, THX males administered 1 or 10 micrograms T4 exhibited full-blown testicular growth and then regression, initiated molt, and had low hypothalamic cGnRH-I, indicating that photostimulated birds that received mid- or high-dose T4 replacement therapy had been programmed for all three components of seasonality. On the other hand, both THX controls and THX males administered low-dose (0.1 microgram) T3 replacement therapy exhibited only modest testicular growth, signifying that neither group had been programmed for any component of seasonality. By contrast, photostimulated THX males that received 0.1 microgram T4, or 1 or 10 micrograms T3, were programmed for testicular growth, but not for photorefractoriness or molt. Collectively, these results show that subcutaneously administered T3 mimicked T4 imperfectly and suggest either that T3 does not program photostimulated male tree sparrows for photorefractoriness and postnuptial molt, or that T3 does not cross the blood-brain barrier as efficiently as does T4.

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Testis-dependent and -independent effects of photoperiod on volumes of song control nuclei in American tree sparrows (Spizella arborea).

Songbirds exhibit seasonal changes in the volumes of song control nuclei. Birds on long, spring-like days have larger nuclei than do birds on short, winter-like days. The mechanisms mediating volumetric changes have not been determined unequivocally, but testosterone (T) is probably involved. This study examined whether testicular factors are uniquely responsible for seasonal changes in the song system, or whether photoperiod has testis-independent effects. Male American tree sparrows were exposed to one of three photoperiodic conditions: (1) Photosensitive birds were retained on short days (8L:16D). Plasma T is rarely detected in such birds. (2) Photosensitive birds were moved from short days to long days (20L:4D) and photostimulated for three weeks. Photostimulation elevates circulating T in photosensitive birds. (3) Photorefractory birds were held at least four months on 20L:4D. Such birds seldom have detectable levels of T, even though they are on long days. In each condition, there were both intact and castrated birds. Castration typically removes circulating T in tree sparrows. The volumes of the high vocal center (HVC), nucleus robustus archistriatalis (RA), and area X were measured. Song nuclei were largest in intact photostimulated birds. Other long-day birds (i.e. castrated photostimulated, and intact and castrated photorefractory groups) had larger song nuclei than did short-day intact or castrated photosensitive birds and did not differ from each other. These data indicate that photoperiod has both testis-dependent and -independent effects on the volumes of song control nuclei.

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The thyroid and the hypothalamus-pituitary-ovarian axis in American tree sparrows (Spizella arborea).

Prompted by evidence of a linkage between the thyroid gland and seasonal reproduction in obligately photoperiodic American tree sparrows (Spizella arborea), we measured circulating T4 (L-thyroxine), hypothalamic cGnRH-I (chicken gonadotropin-releasing hormone, variant I), pituitary and plasma LH (luteinizing hormone), and ovarian mass in euthyroid females moved from short to long days. Our purpose was to correlate temporal changes in T4 and cGnRH-I with each other and with pituitary and plasma LH and ovarian mass. T4 increased early during photostimulation and peaked at Week 1 or 2 on 20L:4D (20 hr light:4 hr dark per day). Hypothalamic cGnRH-I also increased during photostimulation and peaked between Weeks 2 and 4. Pituitary and plasma LH peaked at Weeks 4 and 1, respectively, and ovarian mass increased significantly by Week 6. In a separate study, we charted the progression of postnuptial molt. Molt of the primary remiges began at Week 9, proceeded rapidly through Week 15, and was complete by Week 18. In view of a demonstrated role for the thyroid gland in the control of seasonal reproduction in female tree sparrows and the time-dependent effects of thyroidectomy after photostimulation in male tree sparrows, the observation that circulating T4 increases early during photostimulation suggests the possibility of a causal relationship between the thyroid gland and the hypothalamus-pituitary-ovarian axis, hence a thyroid-hypothalamus-pituitary-ovarian axis.

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Thyroid dysfunction and thyroxine-dependent programming of photoinduced ovarian growth in American tree sparrows (Spizella arborea).

The time course of thyroid dysfunction after injecting 0.30 mCi Na131I was charted in female American tree sparrows (Spizella arborea) transferred to long days on Day 0 and challenged with 30 micrograms bovine thyroid-stimulating hormone on Postinjection Days 4, 7, 46, and 105. Serum T4 (L-thyroxine) was not detectable in thyroidectomized birds, indicating complete thyroid dysfunction by Day 4 and no restoration of thyroid function by Day 105. By contrast, serum T4 concentrations in similarly challenged thyroid-intact controls greatly exceeded assay sensitivity. To determine whether a single injection of T4 can program thyroidectomized female tree sparrows for seasonal reproduction and postnuptial molt, as it can thyroidectomized male tree sparrows, thyroidectomized females were injected with T4 (100 micrograms) or alkaline vehicle (V) either on the first day of photostimulation or 1 week before photostimulation (during which time injected T4 likely was cleared metabolically). Females injected with T4 on the first day of photostimulation showed robust ovarian growth similar to that reported for photostimulated euthyroid females. Both groups of V-injected females, as well as females injected with T4 1 week before photostimulation, showed slow, but reliable, ovarian growth, indicating that thyroidectomized birds can detect an increase in day length. When birds were moved at Week 7.5 or 9 to constant light and given T4 in drinking water (a qualitative assay for absolute photorefractoriness), only one tested photorefractory and molted. The remaining birds, including those injected with T4 1 week before photostimulation and both groups of V-injected controls, tested photosensitive and did not molt. Taken together, these results indicate that T4, or one of its metabolites, programs thyroidectomized female tree sparrows for ovarian growth early during photostimulation. A single T4 injection (100 micrograms) on the first day of photostimulation usually also programs thyroidectomized male tree sparrows for photorefractoriness and molt. Such an injection is far less effective in programming females for these two late-season events.

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The timing of thyroid-dependent programming in seasonally breeding male American tree sparrows (Spizella arborea).

There is convincing evidence that euthyroid male American tree sparrows are already programmed for seasonal reproduction and postnuptial molt by Week 4 of photostimulation. To explore more precisely when, during early photostimulation, thyroid-dependent programming of seasonal events occurs, photosensitive male tree sparrows were radiothyroidectomized or sham thyroidectomized on the first day of photostimulation (Week 0) or at Weeks 1 or 3 thereafter. Birds were monitored for testicular growth and regression over 8 or 12 weeks and then tested for photosensitivity or photorefractoriness by exposing them to constant light and exogenous L-thyroxine for 4 weeks. Molt of the primary flight feathers was scored periodically, and at the end of the experiment, hypothalami were saved for cGnRH-I (chicken gonadotropin-releasing hormone I) assay. Because (1) the thyroid is already dysfunctional by Day 4 after radiothyroidectomy, (2) thyroid-dependent photoinduced gonadal growth is programmed growth, and as demonstrated here, (3) testis length at Week 6 on long days (i.e., maximum or near-maximum testis size) is independent of the time of thyroidectomy at or after the onset of photostimulation, we conclude that male American tree sparrows were programmed for photoperiodic testicular growth during the first week of photostimulation. Based on measurements of testis length and hypothalamic cGnRH-I content, only 2 of 11 birds thyroidectomized at Week 1 were already programmed for photorefractoriness by Week 1, whereas all birds thyroidectomized at Week 3 were so programmed by Week 3. Marked differences in molt scores between photorefractory (thyroidectomized and euthyroid) and photosensitive (thyroidectomized) birds argue that postnuptial molt also was programmed roughly between Weeks 1 and 3. To explain these results, we ascribe organizational-like actions directly or indirectly to endogenous thyroid hormones and argue that the onset of photostimulation opens a window, during which time euthyroid male American tree sparrows are programmed for seasonality. A clear dissociation of photorefractoriness from photoperiodic testicular growth in birds thyroidectomized at Week 1 suggests that separate control circuits mediate these two annually periodic events.

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The photoperiodic control circuit in euthyroid American tree sparrows (Spizella arborea) is already programmed for photorefractoriness by week 4 under long days.

The aim of this investigation was to determine whether spontaneous testicular regression in male American tree sparrows (Spizella arborea) that were thyroidectomized at week 4 of photostimulation manifests photorefractoriness, as it does in chronically photostimulated euthyroid controls. On the basis of our demonstration that exogenous thyroxine stimulates the gonads only when recipient birds are photosensitive, male tree sparrows were thyroidectomized at week 4 of photostimulation (20 h light:4 h dark) and given thyroxine periodically to assay for photosensitivity. When initiated at weeks 4, 7, 10, 13 and 16 of photostimulation, thyroxine replacement therapy had no effect on testis size until week 16, when most recipient birds showed robust testicular growth. The inductive effect of exogenous thyroxine at week 16 confirms that chronic thyroidectomy dissipates photorefractoriness and simulates the effect of short days. The failure of replacement thyroxine to halt spontaneous testicular regression between week 7 and week 13 establishes that spontaneous testicular regression after thyroidectomy manifests photorefractoriness. Moreover, the failure of replacement thyroxine to induce testicular growth between week 4 and week 7 indicates that by week 4 of photostimulation, at least 3 weeks before photoinduced testicular growth ends, male tree sparrows are programmed for photorefractoriness. This conclusion is strengthened by the finding that thyroidectomy at week 4 of photostimulation does not uncouple photorefractoriness and postnuptial moult, which in euthyroid tree sparrows are tightly linked. In another experiment, photosensitive thyroid-intact tree sparrows were moved from 8 h light:16 h dark to 20 h light:4 h dark and given exogenous thyroxine or vehicle through week 6 of photostimulation. Exogenous thyroxine augmented testicular growth.

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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 replacement 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.

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Photorefractory Harris' sparrows (Zonotrichia querula) exposed to a winter-like daylength gradually regain photosensitivity after a lag.

The time course of regaining photosensitivity was monitored in intact and in castrated Harris' sparrows (Zonotrichia querula) held on short days. Measured endpoints, assumed to be equivalent indicators of the photosensitive state, were photoinduced testicular growth in intact males and a photoinduced elevation in plasma LH (luteinizing hormone) concentration in castrated males. Intact males were initially photorefractory when they were moved from long days to short days, and they remained so for at least 3 weeks thereafter. Photosensitivity was partially restored between Weeks 3 and 5 on short days and then gradually increased to a maximum by Week 13. No further change was detected through Week 25 on short days, when the experiment ended. Castrated males, also photorefractory when moved from long days to short days, regained photosensitivity as early as Week 7 on short days, but not uniformly until Week 16. As reflected by linear regression analysis, recovery of photosensitivity was a gradual process through Week 22 on short days, when the experiment ended. Discrepancies between intact and castrated males raise the possibility that the measured endpoints are not equivalent indicators of the photosensitive state. In accordance with an earlier report (Wilson, 1990), recovery of photosensitivity in castrated Harris' sparrows held on short days was not signaled by a spontaneous increase in plasma LH concentration. Indeed, plasma LH concentrations of castrated males remained suppressed even after photosensitivity had been fully restored in intact males.

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On the recovery of photosensitivity in two passerine species, American tree sparrows (Spizella arborea) and Harris' sparrows (Zonotrichia querula).

To test the hypothesis that a spontaneous increase in plasma luteinizing hormone (LH) signals recovery of photosensitivity in castrated passerine birds, LH concentrations were measured weekly in intact and in castrated photorefractory tree sparrows and Harris' sparrows transferred to short days. After 7 weeks on short days (Experiment 1) or after 1, 3, 5, 7, and 10 weeks (Experiment 2) were challenged with long days (1 week) to determine if photosensitivity had been restored. As evidenced by a significant LH response to photostimulation, tree sparrows had regained at least partial photosensitivity after 7 weeks on short days. However, during exposure to short days, plasma LH concentrations in castrated males did not differ from those in intact males, and plasma LH concentrations in intact or castrated males did not vary with time. The first indication that photosensitivity had been partially restored in Harris' sparrows came after 7 weeks on short days, when castrated males responded to photostimulation with a fourfold elevation in plasma LH concentration. However, before week 7 and through week 10, LH concentrations remained suppressed in both intact and castrated males retained on short days. These data show that recovery of photosensitivity in castrated tree sparrows and Harris' sparrows held on short days is not signaled by a spontaneous elevation in plasma LH concentration, and that plasma LH concentrations in acutely photosensitive intact male tree sparrows and Harris' sparrows held on short days are not suppressed because of gonadal negative feedback.

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Extraocular control of seasonal reproduction in female tree sparrows (Spizella arborea).

The work reported here explored extraocular control of seasonal reproduction in a photoperiodic species. Photosensitive female tree sparrows (Spizella arborea) were subjected to bilateral or unilateral ocular enucleation or to sham operation and exposed to a photoregimen designed to stimulate luteinizing hormone (LH) secretion, to induce photorefractoriness, and to restore photosensitivity. As revealed by plasma LH profiles constructed from measurements taken over 181 days, neither bilateral nor unilateral ocular enucleation had a substantive effect on photoinduced LH release, on the development of photorefractoriness, or on the recovery of photosensitivity. In a related experiment, photorefractory sighted females were implanted epicranially with miniature self-powered lights (SPLs) or with non-emitting helium blanks (HBs) and transferred to short days. After 8 weeks, SPLs and HBs were removed, and birds were challenged with long days. As indicated by plasma LH and ovarian responses to the challenge, previously HB-implanted birds had regained photosensitivity, but previously SPL-implanted birds remained photorefractory. When interpreted within the context of the effects of blinding, these findings suggest that an extraocular encephalic mechanism maintains photorefractoriness. Collectively, the data support the hypothesis that photoinduced LH release and the biannual transitions between photosensitivity and photorefractoriness are controlled by an extraocular mechanism(s).

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The drive on luteinizing hormone secretion in castrated tree sparrows (Spizella arborea) exposed to short days is daylength independent.

For many, if not most, photoperiodic species of birds, short days are nongonadostimulatory. The tacit assumption that short days are also nonphotostimulatory was tested by determining whether plasma concentrations of luteinizing hormone (LH) in castrated tree sparrows exposed to short daily photoperiods (8 hr or less) are daylength dependent or independent. Castration of tree sparrows held on an 8-hr daily photoperiod evoked, within 2 weeks, a fivefold elevation in plasma LH concentration. Over the next 24 weeks, plasma LH concentrations of castrated birds were resistant, first to a stepwise reduction in daylength from 8 to 2 hr and later to a 2-, 4-, or 6-hr increase therein. At no time did LH concentrations differ among castrated birds held on the same or different short-day photoregimes. Conversely, at all times, LH concentrations of castrated birds, regardless of photoperiodic history, exceeded those of intact males held on an 8-hr daily photoperiod and sampled at the beginning of the experiment. These data, which argue that the drive on LH secretion in castrated tree sparrows exposed to short days is daylength independent and, therefore, likely intrinsic, verify the tacit assumption that short days are nonphotostimulatory.

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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.

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