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The development of within-song type variation in song sparrows.

We investigated the development of within-song type variation in song sparrows, Melospiza melodia, with two experiments designed to determine how exposure to within-type variation influences the song-learning process and whether within-type variation itself is a learned trait. In the first experiment, we compared learning between two groups of males, one group tutored exclusively with song models presented with no variation, and the other group tutored exclusively with song models presented with a range of within-type variation that is normally produced by birds in the field. The two groups in this experiment did not differ significantly in any measure of how well they learned, suggesting that exposure to within-type variation has no measurable influence on the learning process overall. Nor did the groups differ in the expression of within-song type variation in their own adult songs, demonstrating that within-type variation is not a learned feature of song sparrow song. In the second experiment, we tutored a single group of birds with both invariant and variable models, allowing us to ask how within-type variability affects learning preferences. Young birds preferentially copied song type models presented with variation significantly more than invariant models. Taken together, these experiments provide insight into the evolution of within-song type variation in song sparrows, although the functional significance of this level of variation and learning preferences based on variation remain enigmatic. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

A species-specific acoustic cue for selective song learning in the white-crowned sparrow.

Song learning in birds is paradoxical. Without tutoring, songbirds do not develop normal songs. Yet despite this inability, birds possess extensive foreknowledge, in a mechanistic sense, about the normal song of their species. When given a choice of tape recordings, young, näive songbirds select sounds of their own species for imitation. We tape-tutored white-crowned sparrows, Zonotrichia leucophrys oriantha, with a set of manipulated songs to investigate whether the introductory whistle universally present in white-crowned sparrow song guides selective song learning in this species. Our results confirm that this whistle serves as a cue for song learning, enabling acquisition of normally rejected sounds of other species, including hermit thrush, Catharus guttatus, notes, which have a sound quality distinct from that of natural white-crowned sparrow phrases. Our results support the conclusion that sensory mechanisms rather than motor constraints are primarily responsible for the selectivity seen in song learning. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Changes in brain GnRH associated with photorefractoriness in house sparrows (Passer domesticus).

Temperate zone birds terminate reproduction when they become photorefractory. In many species, refractoriness is "absolute" in that gonadal regression occurs before day length declines, and individuals are reproductively unresponsive even to continuous light. Based on studies of a few species, this form of refractoriness appears to be associated with a reduction (compared with breeding and/or photosensitive birds) in numbers of hypothalamic cells and fibers that are immunoreactive for gonadotropin releasing hormone (GnRH). Some species display "relative" refractoriness, in that day length must decline before gonadal regression occurs, and individuals never lose the capacity to respond to very long days. A few reports suggest that GnRH levels in the brain do not change in relatively refractory birds. House sparrows (Passer domesticus) regress their gonads sooner when day length declines during summer than they do if photoperiod is not permitted to decline after the summer solstice (i.e., they appear to be "relatively" refractory). However, if held on long days they eventually regress the gonads despite no decline in photoperiod (they appear to be "absolutely" refractory). In this experiment, we tested whether gonadal regression was associated with changes in hypothalamic GnRH in adult male house sparrows. We used immunocytochemistry (primary antibody sensitive to both forms of avian GnRH) to compare the distribution and number of GnRH-immunoreactive cells and fibers among reproductively active individuals on long days, "absolutely" refractory individuals on long days, and presumably "relatively" refractory individuals induced to regress by shortened days. Similar reductions in ir-GnRH cell number and cell size occurred in both groups of refractory birds compared with birds still in breeding condition. Gonadal regression also was correlated with reduced staining intensity of cell bodies and reduced fiber staining, but these reductions were somewhat more pronounced in the "absolutely" than the "relatively" refractory birds. We discuss these results in light of other studies of avian GnRH changes with reproductive stage. Our results are consistent with the idea that house sparrows become absolutely refractory, regardless of whether exposed to a decline in photoperiod. However, the results also suggest that relative refractoriness may induce gonadal regression through a cessation of GnRH secretion, whereas absolute refractoriness involves down-regulation of peptide production as well.

Animals↗

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.

Animals↗

Noninvasive corticosterone treatment rapidly increases activity in Gambel's white-crowned sparrows (Zonotrichia leucophrys gambelii).

Recent evidence supports the hypothesis that corticosteroids influence behavioral changes associated with stressful events. Most investigations into this relationship focus on the long-term behavioral effects of corticosterone. Because many behavioral responses to environmental perturbations occur within minutes, we determined what rapid effects corticosterone may have on behavior. With this goal in mind, we devised and evaluated a method of corticosterone delivery which allowed us to examine immediate effects of corticosterone on behavior in a noninvasive manner. White-crowned sparrows (Zonotrichia leucophrys gambelii) were allowed access to mealworms (Tenebrio molitor) injected with corticosterone. Once ingested, the corticosterone moves across the digestive epithelium into the circulation. This method was evaluated using two vehicles: dimethyl sulfoxide and peanut oil. We tested the efficiency and consistency of corticosterone transfer into the circulation for both vehicles. Dimethyl sulfoxide gave a more efficient transfer of corticosterone. Injecting mealworms with corticosterone (carried in dimethyl sulfoxide) and feeding those worms to white-crowned sparrows increased circulating corticosterone to a discrete, repeatable level which peaked within 7 min and was cleared within 60 min. Using this method, we demonstrated that intermediate levels of corticosterone caused an increase in perch hopping in white-crowned sparrows within 15 min of hormone administration. An increase in perch hopping indicated elevated locomotor activity that is consistent with behavioral responses to natural perturbations. High levels of corticosterone did not induce this behavioral change. In light of the rapid effect of corticosterone on behavior, we propose that corticosterone was acting through a nongenomic mechanism.

Animals↗

Melatonin binding in the house sparrow song control system: sexual dimorphism and the effect of photoperiod.

Avian song is a sexually dimorphic behavior which is regulated seasonally. This regulation involves the construction and growth of song control structures: the high vocal center (HVC), nucleus robustus archistrialis (RA), nucleus magnocellularis anterior (MAN), and Area X. Song behavior and its neural correlates are controlled by steroid-dependent and independent processes. The avian circadian system is known to be involved in both daily processes and seasonal reproduction. A major part of this system is the circadian secretion of melatonin by the pineal gland. To determine possible interactions of the circadian and song control systems, the distribution and density of 2-[125I]iodomelatonin (IMEL) binding, an indicator of melatonin sensitivity, were determined in male and female house sparrow brains. Specific binding was found in visual system centers of both genders, but binding in HVC, RA, and Area X was present only in males. Binding in MAN was present in both sexes. Although the effects of short and long photoperiods on male house sparrow IMEL binding in song structures revealed no systematic changes, there were significant differences in binding under different photoperiods in HVC and RA. IMEL binding in the tectofugal nucleus rotundus, however, was consistently highest under short day conditions. IMEL binding in song control nuclei was independent of testicular influence, since castration did not affect it significantly. The data point to a role for the circadian system of house sparrows in song control, but a specific role for melatonin in the daily or seasonal regulation of the song control system in birds, could not be determined.

Animals↗

Territorial aggression and dawn song are modulated by septal vasotocin and vasoactive intestinal polypeptide in male field sparrows (Spizella pusilla).

Previous experiments demonstrate that lesions of the septum produce opposite effects on intraspecific male aggression in the territorial field sparrow (Spizella pusilla) and the colonial zebra finch (Taeniopygia guttata; facilitate vs inhibit, respectively) and intraseptal infusions of arginine vasotocin (AVT) and vasoactive intestinal polypeptide (VIP) modulate aggression in the male zebra finch (facilitate and inhibit, respectively). The present experiments were conducted to test the hypotheses that (1) septal AVT and VIP modulate both overt territorial aggression and the production of territorial song during the dawn chorus in male field sparrows and (2) these neuropeptides will exert effects opposite of those observed in the zebra finch, consistent with the prediction that social organization is associated with septal neuropeptide function. Wild-caught male field sparrows were fitted with chronic guide cannulae directed at the septum and were tested in outdoor aviaries placed in their natural habitat. Intrusion tests (introduction of a stimulus male) and dawn song observations were conducted following infusion of AVT, VIP, or saline control. Consistent with predictions, infusion of AVT significantly inhibited chases and significantly increased chase latency. No significant effects of VIP on chasing or chase latency were observed, although most subjects were more aggressive following infusion of VIP. Both AVT and VIP produced significant, selective effects on the complex (agonistic) song type (facilitation and inhibition, respectively) and produced no effect on the simple (multipurpose) song type. Thus, song and overt aggression appear be modulated independently by septal neuropeptides, and septal AVT and VIP function may differ between species which differ in the expression of territorial or colonial social organizations.

Aggression↗

The Old World sparrows (genus Passer) phylogeography and their relative abundance of nuclear mtDNA pseudogenes.

The phylogenetic relationships of genus Passer (Old World sparrows) have been studied with species covering their complete world living range. Mitochondrial (mt) cyt b genes and pseudogenes have been analyzed, the latter being strikingly abundant in genus Passer compared with other studied songbirds. The significance of these Passer pseudogenes is presently unclear. The mechanisms by which mt cyt b genes become pseudogenes after nuclear translocation are discussed together with their mode of evolution, i.e., transition/transversion mitochondrial ratio is decreased in the nucleus, as is the constraint for variability at the three codon positions. However, the skewed base composition according to codon position (in 1st position the percentage is very similar for the four bases, in 2nd position there are fewer percentage of A and G and more percentage of T, and in 3rd codon position fewer percentage of G and T and is very rich in A and C) is maintained in the translocated nuclear pseudogenes. Different nuclear internal mechanisms and/or selective pressures must exist for explaining this nuclear/mitochondrial differential DNA base evolutive variability. Also, the phylogenetic usefulness of pseudogenes for defining relationships between closely related lineages is stressed. The analyses suggest that the primitive genus Passer species comes from Africa, the Cape sparrow being the oldest: P. hispaniolensis italiae is more likely conspecific to P. domesticus than to P. hispaniolensis. Also, Passer species are not included within weavers or Estrildinae or Emberizinae, as previously suggested. European and American Emberizinae sparrows are closely related to each other and seem to be the earliest species that radiated among the studied songbirds (all in the Miocene Epoch).

Cell Nucleus↗

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.

Animals↗

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.

Animals↗

Short-term changes in plasma levels of hormones during establishment and defense of a breeding territory in male song sparrows, Melospiza melodia.

When territorial male song sparrows are captured and removed from their territories, previously unmated and nonterritorial males will take over those vacant territories within 12-72 hr. Plasma levels of testosterone are elevated in these replacement males as well as in their neighbors. Since the latter already have territories, it is suggested that the agonistic interactions over territory boundaries, or behavioral stimuli from challenging males, rather than ownership of a territory per se, stimulates secretion of testosterone. To test this hypothesis further, male song sparrows were challenged by experimental simulation of a territorial intrusion. This procedure involved placing a caged male song sparrow in the center of the subject's territory and playing tape recorded conspecific song through an adjacent speaker. Responding males were then captured at intervals after onset of the intrusion. Plasma levels of testosterone were significantly higher in males exposed to experimental territorial intrusion than they were in controls, supporting the hypothesis that behavioral stimuli emanating from an intruding male can act as supplementary information stimulating secretion of testosterone.

Aggression↗

Red cell organic phosphates and Bohr effects in house sparrow blood.

CO2 and fixed acid Bohr effects (d log PO2/d pH) were determined for fresh whole blood of adult house sparrow (Passer domesticus) at 35, 41 and 45 degrees C. At each temperature, the effects of titrating blood with CO2 at constant base excess (CO2 Bohr effect) were similar at P50 (35 degrees, -0.48; 41 degrees, -0.49; 45 degrees, -0.49). The CO2 Bohr slopes were also reasonably saturation independent between 10 and 90% S. Fixed acid Bohr values, determined by titrating sparrow blood with HCl and NaHCO3 at 4% CO2, were significantly less than the corresponding CO2 coefficients at half saturation (35 degrees, -0.43; 41 degrees, -0.39; 45 degrees, -0.41). The difference between CO2 and H+ Bohr effects, assumed here to represent carbamino CO2 binding to hemoglobin, decreased in magnitude with increasing saturation at each temperature. Inositol pentaphosphate (IPP, 3.1 mumol/ml RBC) and ATP (7.7 mumol/ml RBC) were the major organic phosphates present in Passer erythrocytes. CO2 and organic phosphates are known to complete for common binding sites on the Hb molecule. Because of IPP's strong binding affinity and high concentration in most avian red cells, carbamate formation is generally suppressed in bird blood. The presence of a small but significant specific CO2 effect in Passer blood may indicate that one or both sparrow isohemoglobins has reduced affinity for IPP and/or ATP, permitting CO2 to compete more effectively in Hb-carbamate formation.

Adenosine Triphosphate↗

Effect of melatonin on circadian rhythm, locomotor activity and body temperature in the intact house sparrow, Japanese quail and owl.

We compared the effect of melatonin on circadian rhythm, body temperature, and locomotion in the intact house sparrow, Japanese quail and owl. Daily treatment with melatonin at a fixed time did not entrain the free-running rhythm of locomotor activity in the house sparrow and the disrupted rhythm in Japanese quail under constant dim light. However, melatonin clearly inhibited movement for several hours after treatment. The duration of resting after injection of melatonin was dose-dependent. Body temperature was significantly decreased after melatonin treatment, the effect being more potent during the active phase than in the resting phase. Although this effect of melatonin on body temperature was also dose-dependent, the magnitude of the decrease in body temperature after injection of melatonin was greater in the house sparrow than in the Japanese quail. On the other hand, melatonin induced a further large decrease of body temperature in a nocturnal bird, the owl, whose pineal gland is degenerate. The decrease of body temperature was larger in the active phase than in the resting phase, and melatonin did not prevent movement in spite of the decrease in body temperature. These results suggest that the effects of melatonin on circadian rhythm, locomotor activity and body temperature differ among avian species, and that these mechanisms may not be linked to each other.

Animals↗

Intracerebroventricular corticotropin-releasing factor decreases food intake in white-crowned sparrows.

Neuropeptides such as corticotropin-releasing factor (CRF) may play a role in regulating the pronounced seasonal changes in food intake shown by white-crowned sparrows (Zonotrichia leucophrys gambelii). White-crowned sparrows held on short day length received injections into the third ventricle (icv) of saline or 5.0, 15.0, and 30 microg/kg. Meal size over the subsequent 180 min was significantly depressed in a dose-dependent fashion. Other non-specific behaviors such as preening, hopping, and immobile behaviors appeared to not be affected by a dose that suppressed food intake. This experiment suggests that white-crowned sparrows, when weight-stable, respond to CRF in a manner comparable with several mammalian species.

Animals↗

Adjustment of house sparrow circadian rhythms to a simultaneously applied light and food zeitgeber.

Periodic food availability has been shown to be an effective circadian zeitgeber in many vertebrates. It is still unclear, however, i) whether light-active species like most birds can synchronize with food cycles in the presence of a strong light-dark (LD) cycle and ii) whether it is common among non-mammalian vertebrates to use a separate circadian oscillator to synchronize with food cycles as most mammals do. We investigated these questions experimentally by exposing house sparrows simultaneously to two zeitgebers: light and food. The LD cycle was set at 1410 h; food was always available for 12 hour per day, but at different phases of the LD cycle. The effects of the two zeitgebers were analyzed by observing two behavioral outputs of the birds' circadian system, the rhythms of locomotion and feeding. The data revealed that light acted as the dominant zeitgeber in most conditions. Food cycles affected the phase of the behavioral rhythms of the birds only when the food was presented no later than 3 h after the onset of light. Apart from their synchronizing actions both light and food cycles also exerted direct (masking) effects on the behavioral rhythms of the birds. The results suggest that the circadian system of house sparrows can indeed adjust to two simultaneous environmental periodicities, i.e. light and food. We propose that light is the stronger zeitgeber and plays a 'permissive' role in determining the phases at which synchronization with food cycles comes into effect. We did not find evidence that the house sparrows' behavioral rhythms are controlled by a food-entrainable circadian oscillator that is distinct from the light-entrainable system as is the case in most mammals. The differences in the patterns of food synchronization and organization of the circadian system that appear to exist between different species can be interpreted in two ways: i) species of different phylogenetic origin (e.g., mammals versus birds) evolved different circadian system or ii) regardless of phylogeny, species with different ecological requirements show a specialization in their circadian organization which is adjusted to the importance of zeitgebers in nature.

Adaptation, Psychological↗

A phylogenetic analysis of the emberizid sparrows based on three mitochondrial genes.

Previous molecular phylogenetic studies have examined the taxonomic relationships among a number of typical emberizid sparrow genera. To help clarify these relationships, we sequenced a 1673 base pair fragment for the complete sequence of three mitochondrial genes: adenosine triphosphatase (Atp8 and Atp6) and cytochrome oxidase subunit III (COIII) for 38 sparrow species, along with Passerina amoena (Cardinalidae) and Piranga ludoviciana (Thraupidae) which were selected as the outgroups. Our analysis confirms the monophyly of traditional genera such as Junco, Melospiza, and Zonotrichia. Although Calcarius and Plectrophenax are often thought to be putative emberizids, all our analyses placed these genera basal to all other sparrows examined. As observed with Calcarius, Spizella did not form a monophyletic group, with S. arborea being the sister-taxon to Passerella iliaca. Our analyses also suggest that Aimophila ruficeps is probably more closely related to the "brown towhees" (Pipilo aberti, P. crissalis, and P. fuscus) than its putative congeners. The genus Ammodramus was also not monophyletic, since it appears that Passerculus sandwichensis is more closely related to A. henslowii and A. leconteii then either one is related to its congener A. savannarum. Finally, our analyses exhibited other unsuspected associations, such as the sister-taxon relationships between Amphispiza bilineata and the Chondestes grammacus/Calamospiza melanocorys clade, and Amphispiza belli and Pooecetes gramineus.

Adenosine Triphosphatases↗

Song frequency as a cue for recognition of species and individuals in the field sparrow (Spizella pusilla).

Estimate the amount of frequency variation that affected individual and species recognition by song in territorial male field sparrows with field playbacks of neighbors' songs lowered in frequency in 6 increments between 324 and 1296 Hz. Males gave strong responses to neighbors' songs that had been lowered by about 400 Hz, which is about 11% of the mean midfrequency in field sparrow song. Territorial responses were eliminated when neighbors' songs were lowered by more than about 900 Hz. Thus, there is a fairly narrow range of frequencies within which field sparrow songs can vary while functioning in communication. Ss' responses appear to be constrained by the normal range of song frequency variation within the species, but the range of variability within songs of individuals did not influence the degree of frequency shift required to render neighbors' songs unfamiliar.

Animal Communication↗

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.

Animals↗