Wavelength dependency of light-induced effects on photoperiodic clock in the migratory blackheaded bunting (Emberiza melanocephala)

The effects of light wavelength on photoperiodic clock were determined in the migratory male blackheaded bunting (Emberiza melanocephala). We constructed an action spectrum for photoperiodic induction (body fattening, gain in body mass, and gonadal recrudescence) by exposing birds for 4.5 weeks to 13 h light per day (L:D = 13:11 h) of white (control), blue (450 nm), or red (640 nm) color at irradiances ranging from 0.028 to 1.4Wm(-2). The threshold light irradiance for photoinduction was about 10-fold higher for blue, compared to red and white light. Phase-dependent effects of light wavelength on the photoperiodic clock were further examined in the next two sets of skeleton photoperiods (SKPs). In the first set of SKPs, birds were exposed for four weeks to asymmetrical light periods (L:D:L:D= 6:6:1:11 h) at 0.25+/-0.01 W m(-2); two light periods applied were of the same (450nm: blue:blue, B:B; 640nm, red:red, R:R) or different (blue:red, B:R or red:blue, R:B) wavelengths, or of white:white (W:W, controls). Photoperiodic induction occurred under R:R and B:R, but not under B:B and R:B light conditions; the W:W condition induced an intermediate response. The second set of SKPs used symmetrical light periods (L:D:L:D = 1:11:1:11 h), and measured effects also on the activity rhythm. Birds were first exposed to one of the four SKPs (R:R, B:B, R:B, or B:R) for three weeks, subsequently were released into dim constant light (LLdim; approximately 0.01 Wm(-2), the night light used in an L:D cycle) for two weeks, and then were returned to respective SKPs for another three weeks. Activity was greater in the R:R compared to B:B, and in B:R compared to R:B light condition. Zugunruhe (intense nighttime activity, indicating migratory restlessness in a caged situation) developed under the R:R and B:R, but not the B:B and R:B, light condition. Under LLdim, all birds free-ran with a period >24h, the Zugunruhe had a circadian period longer than the daytime activity, and the re-entrainment to SKPs was influenced by the position of light periods relative to circadian phase of the activity rhythm. Photoperiodic induction at the end of 8 weeks was found in the R:R and B:R, but not in B:B, light conditions; in the R:B condition only one bird had initiated testes. Taken together, these results suggest that in the blackheaded bunting, the circadian photoperiodic clock is differentially responsive to light wavelengths; this responsiveness is phase-dependent, and the development of Zugunruhe reflects a true circadian function. Wavelength-dependent response of the photoperiodic clock could be part of an adaptive strategy in evolution of the seasonality in reproduction and migration among photoperiodic species under wild conditions.     

The photoperiodic clock in blackheaded buntings (Emberiza melanocephala) is mediated by a self-sustaining circadian system

Three experimental protocols were employed to clarify whether the circadian system is involved in photoperiodic time-measurement in the blackheaded bunting, Emberiza melanocephala. In a single-pulse paradigm, one 8-h light pulse was delivered at different times to groups of birds across three days of constant darkness (DD). Photoperiodic induction, as measured by a rise in plasma luteinizing hormone (LH), showed clear circadian rhythmicity. The second experiment examined the LH responses in birds exposed to lighting cycles using a Nanda-Hamner type of protocol and confirmed full photostimulation under 6L:30D. The third experiment measured the time of the first photo-induced rise in LH in birds subjected to 30 h of continuous light following entrainment under short days (6L:18D). This experiment aimed to identify the position of the photoinducible phase ( _i). LH first rose at hour 18 following dawn indicating that _i lies in the middle of the day. Plasma concentrations of melatonin were also measured under 6L:18D and 6L:30D light cycles as another physiological marker of the circadian system in buntings. The pattern of melatonin secretion under these LD cycles showed properties consistent with the driving oscillator being circadian in nature. It is concluded that the circadian pacemaker driving the photoinducible rhythm in blackheaded bunting is strongly self-sustaining and free-runs under constant conditions.     

Regulation of seasonality in the migratory male blackheaded bunting (Emberiza melanocephala)

The present study was carried out on a Palearctic-Indian migratory species, the blackheaded bunting (Emberiza melanocephala), to understand the importance of photoperiodism and circannual rhythms in determining seasonality in changes in body mass and testis size in birds. An initial experiment determined the effects of duration and intensity of light on photoperiodic induction. The birds were exposed to different photoperiods (hours of light:hours of darkness; 11.5L:12.5D, 12L:12D, 12.5L:11.5D and 13L:11D) at the same (approximately 450 lux) light intensity, and to 13L:11D at different light intensities (50-, 100-, 400-, 800- and 1000-lux). The induction and subsequent regression of photoperiodic responses were dependent upon duration and intensity of the light period until these reached threshold. A second experiment investigated if an endogenous seasonal rhythm underlies photoperiodism in buntings. Birds maintained since February on a 8L: 16D photoperiod (a non-inductive short day length invariably used to ensure photosensitivity in photoperiodic species) were subjected periodically to 16L:8D (a long day length), one group every month from mid-March to mid-August. The magnitude of long day response in body mass and testes decreased as the duration of the short days progressed, but testicular response was restored in birds that were exposed to long days in July and August. The birds exposed simultaneously to short, long, and natural day lengths for 32 weeks underwent an induction-regression cycle under long days and natural day lengths, but not under short days in which a decrease in body mass occurred after about 20 weeks. The last experiment examined the importance of latitudinal migration on photoperiodism, by comparing the response to long days of three groups which included birds from populations those were held in the outdoor aviary for 1 or 2 years at 27 degrees N and those immediately arrived from their breeding grounds (approximately 40 degrees N). There was no difference in the photoperiodic induction among the three groups, indicating that neither experience to changing photoperiods during a migratory journey, nor to long photoperiods at breeding grounds, were critical for a subsequent response (initiation-termination-reinitiation) cycle. Taken together, these findings suggest that (1) the blackheaded bunting has its own endogenous timing program, which is regulated by the photoperiod, and (2) the photoperiodic programs of bunting are flexible enough to accommodate variations in the amplitude of environmental cycles. Thus, it appears that photoperiodism has evolved independently of the evolution of migration in this species.     

Food availability affects circadian clock-controlled activity and Zugunruhe in the night migratory male blackheaded bunting (Emberiza melanocephala)

This study investigated the functional linkage between food availability and activity behavior in the Palaearctic Indian night migratory blackheaded bunting (Emberiza melanocephala) subjected to artificial light-dark (LD) cycles. Two experiments were performed on photosensitive birds. In the first one, birds were exposed to short days (LD 10/14; Experiment 1A), long days (LD 13/11; Experiment 1B), or increasing daylengths (8 to 13?h light/d; Experiment 1C) and presented with food either for the whole or a restricted duration of the light period. In Experiments 1A and 1B, illumination of the light and dark periods or of the dark period, alone, was changed to assess the influence of the light environment on direct and circadian responses to food cycles. In the second experiment, birds were exposed to LD 12/12 or LD 8/16 with food availability overlapping with the light (light and food presence in phase) or dark period (light and food presence in antiphase). Also, birds were subjected to constant dim light (LL(dim)) to examine the phase of the activity rhythms under synchronizing influence of the food cycles. Similarly, the presentation of food ad libitum (free food; FF) during an experiment examined the effects of the food-restriction regimes on activity rhythms. A continuous measurement of the activity-rest pattern was done to examine both the circadian and direct effects of the food and LD cycles. Measurement of activity at night enabled assessment of the migratory phenotype, premigratory restlessness, or Zugunruhe. The results show that (i) light masked the food effects if they were present together; (ii) birds had a higher anticipatory activity and food intake during restricted feeding conditions; and (iii) food at night alone reduced both the duration and amount of Zugunruhe as compared to food during the day alone. This suggests that food affects both the daily activity and seasonal Zugunruhe, and food cycles act as a synchronizer of circadian rhythms in the absence of dominant natural environmental synchronizers, such as the light-dark cycle.     

Investigations of photoperiodically induced fattening in migratory blackheaded bunting (Emberiza melanocephala)(Ayes)

The body fattening and weight gain preceding vernal migration in birds is timed by a set of environmental factors of which daylength is predominant. However, the mechanism(s) by which these events is determined is poorly understood. Previous investigations on a photoperiodic migratory species, the blackheaded bunting ( Emberiza melanocephala ), indicate the involvement of a light-sensitive circadian rhythm during initiation of fat deposition and body weight gain. This communication presents data from another set of experiments aimed to characterize further the mechanism(s) of fat deposition in the same species.
Groups of photosensitive, unstimulated and stimulated birds were subjected to transfer and superimposition experiments for 30 days. While the former set included shifting of long-day (LD) birds to DD, SD (short days), DD/LD and SD/LD, in the latter a 90-minute bright light was superimposed at two different times of the day during the dim-green lighted phase 15L:9D of varying intensity. Birds were weighed at the beginning and at the end of experiments. Those in transfer cycles were also weighed at 10-day intervals. The results suggest that the premigratory body fattening and weight gain in blackheaded buntings is light dependent and timed by environmental daylength in accordance with the photosensitive endogenous circadian rhythm (ECR). They also show that the photoperiodic responses in birds in general are mediated by circadian rhythm(s).     

The influence of light wavelength on reproductive photorefractoriness in migratory blackheaded bunting (Emberiza melanocephala).

There are two effects of long day length on reproductive responses in birds, one is the photoinduction of gonadal growth and maturation and the other is the induction of gonadal regression and photorefractoriness. Although it is likely that the same photoreceptors are involved in the photoinduction of gonadal growth and the onset and maintenance of photorefractoriness. and so the influence of wavelength should be similar, this has not been investigated. Therefore, we investigated the influence of light wavelength on reproductive photorefractoriness in the migratory male blackheaded bunting held under long photoperiods. In mid May, when photoperiod was approximately 14L:10D (14 hours light:10 hours darkness), eight groups of sexually mature birds were moved indoors on an artificial photoperiod of 14L:10D (L - 450 lux. D - 0 lux). Then after 3 weeks, for six groups, a 4-h light period in the morning (zt 0-4; zt 0 [zeitgeber time 0] refers to the beginning of lights-on period) or in the evening (zt 10-14) was substituted with green (428 nm), red (654 nm) or white light at 16 +/- 2 lux intensity. Of the remaining two groups, one was maintained on 14L: 10D and the other transferred to 10L:14D: these served as controls. At the end of 4 weeks, all birds were found to have undergone testicular regression, irrespective of LD cycle they were exposed to. When these gonadally regressed birds were subjected to 16L:8D for another 4 weeks, to test their responsiveness to the stimulatory effects of long day lengths, only those exposed to 10L:14D and 14L:10D with a 4-h green light period showed testicular regrowth. On the other hand, birds exposed to 14L:10D with a 4-h white or red light period remained fully regressed, similar to 14L:10D controls. Except for some individual difference, there was no difference in response between the groups that received a 4-h light period in the morning and that received it in the evening. These results suggest that the wavelengths of light influence induction of buntings from the photosensitive state into the photorefractory state. Whereas the short light wavelengths facilitated recovery from the photorefractoriness, the long light wavelengths were more effective in maintaining the photorefractoriness.     

Differential responses of the photoperiodic clock in two passerine birds possessing a strongly self-sustained circadian system

To investigate whether the photoperiodic clocks of species possessing strongly self-sustaining circadian clocks share identical features, we compared the full response cycle (initiation and termination of the response) in body mass and testes of the non-migratory house sparrow (Passer domesticus) with that of the migratory redheaded bunting (Emberiza bruniceps) under Nanda-Hamner experiments. Birds were exposed to a 36 h day (L:D=6:30 h), controls exposed to a 24 h day (L:D=6:18 h), for a period of 31 weeks. By week 18 of L:D=6:18 h, there was a small increase in body mass among sparrows, but not among buntings, and the testes of bunting did not grow, while those of sparrow grew slightly. The response to L:D=6:30 h is of particular interest. There was a rapid gain and subsequent loss in the body mass of bunting, but not of sparrows. Further, both species underwent a testicular cycle as if they were exposed to long days, but the response of sparrows was slower and hence delayed the attainment of peak testicular size. Such a differential response to exotic light cycles between these two photosensitive species, despite their similar circadian oscillatory properties (strong self-sustainment), could suggest a species-specific adaptation of the endogenous clock involved in photoperiodic regulation of avian seasonality.     

Daily light regulates seasonal responses in the migratory male redheaded bunting (Emberiza bruniceps)

This study analyzed the role of day length in regulation of seasonal body fattening and testicular growth in a latitudinal Palaearctic-Indian migrant, the redheaded bunting (Emberiza bruniceps). When exposed to increasing photoperiods (hours of light: hours of darkness; 11.5L:12.5D, 12L:12D, 12.5L:11.5D, 13L:11D, 14L:10D, and 18L:6D) for 9-12 weeks, buntings responded in a photoperiod-dependent manner and underwent growth and regression cycle under photoperiods of > or =12 hr per day. Also, the response to a long photoperiod of birds that were held under natural photoperiods at 27 degrees N for 2 years was similar to those who arrived the same year from their breeding grounds ( approximately 40 degrees N), suggesting that the experience of higher amplitude day-night (light-dark, LD) cycles during migratory and breeding seasons were not critical for the subsequent response (initiation-termination-reinitiation) cycle. Another experiment examined entrainment of the circadian photoperiodic rhythm in buntings by subjecting them to T=24+/-2 hr LD-cycles with 8 hr photophase and to T=22 and 24 hr with 11 hr photophase. The results showed a reduction in critical day length under T=22 hr LD-cycle. In the last experiment, we constructed an action spectrum for photoperiodic induction by exposing birds for 4.5 weeks to 13L:11D of white (control), blue (450 nm), or red (640 nm) light at irradiances ranging from 0.028 to 1.4 W m(-2). The threshold light irradiance for photoinduction was about 10-fold higher for blue light, than for red and white lights. These results conclude that the daily light of the environment regulates the endogenous program that times seasonal responses in body fattening and testicular cycles of the redheaded bunting.     

A short red light pulse during dark phase of LD-cycle perturbs the hamster's circadian clock

In this study we investigated the influence of red light, which naturally occurs during dawn and dusk, on locomotor activity and body temperature rhythms of Djungarian hamsters (Phodopus sungarus). A single weak red light pulse given 2 h before regular lights on had acute as well as long-term effects persisting for several days following exposure. The hamsters immediately stopped their locomotor activity, accompanied by a drop in body temperature. In the following undisturbed nights (LD 168) the nocturnal activity stopped earlier than usual. This lasting effect of the light pulse was more pronounced than the acute effect. The activity phase compressed gradually during 3 to 5 days after the light pulse was administered while time of activity onset was almost unaffected. It took 6 to 11 days for complete recovery of the original activity phase. The maximal activity compression and the recovery period depended on the duration of the single red light pulse and its intensity. Red light pulses of 15 min duration were about twice effective as 1 min pulses; and the effect of a red light pulse of 130 mW/m² was about 1.5 times stronger than a 30 mW/m² red light pulse. The maximal value of activity phase compression reached in this experiment was 2.5+0.2 h with a recovery period of 11.1±0.3 days following a given red light pulse of 90 mW/m² and 15 min. The morning oscillator seems to be persistently affected. This indicates a very high photosensitivity of the Djungarian hamster's circadian system to red light.Abbreviations T _b body temperature - DD constant darkness - LD light:dark cycle - LL constant light - duration of activity phase - CT circadian time - PRC phase response curve - SCN suprachiasmatic nuclei     

Biochronometry of photoperiodically induced fat deposition in a migratory finch, the Blackheaded bunting (Emberiza melanocephala) (Aves)

This investigation attempts to identify the mechanism(s) involved in the fat deposition in a photoperiodic migratory species, the Blackheaded bunting ( Emberiza melanocephala ). Groups of photosensitive male buntings were exposed to resonance, interrupted night, and ultra-short-day light cycles for 35, 42 and 75 days, respectively. Birds were weighed at the beginning and at the end of the experiments. Those exposed to ultra-short-day light cycles were also weighed at critical intervals during the treatment period. Our results indicate that: (a) a light-sensitive rhythm with a period of about 24 hr is involved in the photoperiodic induction of premigratory fattening and weight gain in Blackheaded buntings: (b) buntings possess a bimodal pattern 'of sensitivity to photoperiods that induce fattening, and (c) this endogenous circadian rhythm can be entrained by an ultrashort photophase of 3 h if the latter is coupled with scotophases of specific duration.     

Pathways of ocular entrainment in Marpissa marina (Araneae,Salticidae)

Depending on the animal species, photoreceptors are located in the visual organs, in non-visual organs or in both. Because of unique characteristics of vision containing several different pairs of eyes, I chose the jumping spider (salticid) Marpissa marina (Araneae: Salticidae; Goyen, 1892) for this study. Eyes in spiders are categorized in two groups of principal and secondary. Specifically, my aim was to determine which eyes are dedicated to regulation of the central circadian rhythm and to illuminate the pathway(s) of ocular entrainment in jumping spiders. To achieve this, I used an opaque elastic paste to prevent entry of light to the photoreceptors. My procedure was to measure spider activity levels over eight days as well as spiders responses to a 6 h delay shift in light/dark cycle. This would be made first with uncovered eyes (and sham covers) and then with distinct pairs of eyes covered. The results revealed that, unlike the secondary eyes, light information gathered through AMEs did not lead directly or indirectly to the parts of the circadian system that contain circadian pacemakers.     

The complex regulation of ferredoxin/thioredoxin-related genes by light and the circadian clock

The biochemical properties of the ferredoxin/thioredoxin transduction pathway regulating the activity of key carbon-fixation enzymes through post-translational modifications are well characterized but little is known about the regulation of the different genes. In the present study, we investigated in Chlamydomonas reinhardtii the regulation of the expression of ferredoxin, thioredoxin m, ferredoxin-NADP reductase, phosphoribulokinase, as well as that of cytosolic thioredoxin h, the function of which is still largely unknown. The effects of light, the circadian clock and active cell division were investigated by northern blotting. The five genes were found to be regulated by light and the circadian clock but with different kinetics and amplitudes. This leads for the first time to the proposal that an extra-chloroplastic thioredoxin is possibly implicated in light and/or circadian-related processes. An interplay between several light-transduction pathways in controlling the expression of the genes is suggested by the expression studies and the theoretical analysis of the promoters. Received: 2 December 1998 / Accepted: 19 March 1999     

Entrainment to light of circadian activity rhythms in tench (Tinca tinca)

The present article analyzes locomotor activity rhythms in Tinca tinca. To that end, three different experiments were conducted on 24 animals (20 g body weight) kept in pairs in 60-liter aquaria fitted with infrared sensors connected to a computer to continuously record fish movements. The first experiment was designed to study the endogenous circadian clock under free-running conditions [ultradian 40:40 min LD pulses and constant dark (DD)] and after shifting the LD cycle. Our results demonstrate that tench has a strictly nocturnal activity pattern, an endogenous rhythm being evident in 45.8% of the fish analyzed. The second experiment was conducted to test the influence of different photoperiods (LD 6:18, 12:12, 18:6, and 22:2) on locomotor activity, the results showing that even under an extremely long photoperiod, tench activity is restricted to dark hours. The third experiment examined the effect of light intensity on locomotor activity rhythms. When fish were exposed to decreasing light intensities (from 300:0 lux to 30:0, 3:0, and 0.3:0 lux) while maintaining a constant photoperiod (LD 12:12), the highest percentage of locomotor activity was in all cases associated with the hours of complete darkness (0 lux). In short, our results clearly show that (a) tench is a species with a strictly nocturnal behavior, and (b) daily activity rhythms gradually entrain after shifting the LD cycle and persist under free-running conditions, pointing to their circadian nature. However, light strongly influences activity rhythms, since (c) the length of the active phase is directly controlled by the photophase, and (d) strictly nocturnal behavior persists even under very dim light conditions (0.3 lux). The above findings deepen our knowledge of tench behavior, which may help to optimize the aquacultural management of this species, for example, by adjusting feeding strategies to their nocturnal behavior.     

A circadian system is involved in photoperiodic entrainment of the circannual rhythm of Anthrenus verbasci

In the circannual pupation rhythm of the varied carpet beetle, Anthrenus verbasci, entrainment to annual cycles is achieved by phase resetting of the circannual oscillator in response to photoperiodic changes. In order to examine whether a circadian system is involved in expression of the periodic pattern and phase resetting of the circannual rhythm as photoperiodic responses, we exposed larvae to light-dark cycles with a short photophase followed by a variable scotophase (the Nanda-Hamner protocol). When the cycle length (T) was a multiple of 24 h, i.e., 24, 48, or 72 h, short-day effects were clearer than when T was far from a multiple of 24 h, i.e., 36 or 60 h. Exposure to light-dark cycles of T = 36 h had effects similar to exposure to long-day cycles of T = 24 h. The magnitude of phase shifts depended on the duration and the phase of exposure to the cycles of T = 36 or 60 h. It was therefore concluded that a circadian system is involved in photoperiodic time measurement for phase resetting of the circannual oscillator of A. verbasci.     

Daily torpor in the Djungarian hamster (Phodopus sungorus): photoperiodic regulation,characteristics and circadian organization

1. The daily torpor was measured by oxygen uptake in Djungarian hamsters during adaptation to a short photoperiod (SP: 10L, 14D) at 20 degrees C. In these constant conditions the torpor presented metabolic characteristics and a daily time course independent of the duration of adaptations to SP. 2. The frequency of torpor bouts increased during SP exposure and its maximum was reached after about 130 days. The frequency of torpor was greater in males than in females. 3. The incidence of torpor was increased by constant dark exposure and this is discussed as a protective mechanism for the individual animal's ability to survive. 4. The temporal organization of daily torpor was demonstrated to be directly synchronized by the day-night cycle and to be controlled by an endogenous circadian function.     

Circadian rhythm in plasma aldosterone concentration and its seasonal modulation in the camel (Camelus dromedarius) living in the Algerian Sahara desert

Five male camels dwelling in the Algerian Sahara were studied for circadian rhythmicity in plasma aldosterone concentration and its seasonal modulation. Blood was sampled at a frequency of 1 h or less for a span of 27 h during each season of the year. The mean plasma aldosterone concentration exhibited a significant circadian rhythmicity in every season of the year. Plasma aldosterone concentration was lowest in the morning, increased in the afternoon, and generally highest in the late evening. The peak of the circadian rhythm exhibited seasonal variation; it occurred at 20:04h in October, 16:41h in December, 20:40h in March, and 24:16h in June. The rhythm's 24h mean also exhibited seasonal variability, being significantly higher in March and June compared to October.     

Wavelength Dependency of Light-Induced Effects on Photoperiodic Clock in the Migratory Blackheaded Bunting (Emberiza melanocephala)

The effects of light wavelength on photoperiodic clock were determined in the migratory male blackheaded bunting (Emberiza melanocephala). We constructed an action spectrum for photoperiodic induction (body fattening, gain in body mass, and gonadal recrudescence) by exposing birds for 4.5 weeks to 13 h light per day (L:D = 13:11 h) of white (control), blue (450 nm), or red (640 nm) color at irradiances ranging from 0.028 to 1.4 W m^?2. The threshold light irradiance for photoinduction was about 10-fold higher for blue, compared to red and white light. Phase-dependent effects of light wavelength on the photoperiodic clock were further examined in the next two sets of skeleton photoperiods (SKPs). In the first set of SKPs, birds were exposed for four weeks to asymmetrical light periods (L:D:L:D = 6:6:1:11 h) at 0.25 ± 0.01 W m^?2; two light periods applied were of the same (450 nm: blue:blue, B:B; 640 nm, red:red, R:R) or different (blue:red, B:R or red:blue, R:B) wavelengths, or of white:white (W:W, controls). Photoperiodic induction occurred under R:R and B:R, but not under B:B and R:B light conditions; the W:W condition induced an intermediate response. The second set of SKPs used symmetrical light periods (L:D:L:D = 1:11:1:11 h), and measured effects also on the activity rhythm. Birds were first exposed to one of the four SKPs (R:R, B:B, R:B, or B:R) for three weeks, subsequently were released into dim constant light (LL_dim; ?0.01 W m^?2, the night light used in an L:D cycle) for two weeks, and then were returned to respective SKPs for another three weeks. Activity was greater in the R:R compared to B:B, and in B:R compared to R:B light condition. Zugunruhe (intense nighttime activity, indicating migratory restlessness in a caged situation) developed under the R:R and B:R, but not the B:B and R:B, light condition. Under LL_dim, all birds free-ran with a period >24 h, the Zugunruhe had a circadian period longer than the daytime activity, and the re-entrainment to SKPs was influenced by the position of light periods relative to circadian phase of the activity rhythm. Photoperiodic induction at the end of 8 weeks was found in the R:R and B:R, but not in B:B, light conditions; in the R:B condition only one bird had initiated testes. Taken together, these results suggest that in the blackheaded bunting, the circadian photoperiodic clock is differentially responsive to light wavelengths; this responsiveness is phase-dependent, and the development of Zugunruhe reflects a true circadian function. Wavelength-dependent response of the photoperiodic clock could be part of an adaptive strategy in evolution of the seasonality in reproduction and migration among photoperiodic species under wild conditions.