Does Unusual Entrainment of the Circadian System Under T36h Photocycles Reduce the Critical Daylength for Photoperiodic Induction in Japanese Quail?

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Abstract
In photoperiodic species, short daylength resonance cycles of modulo t + ½t (t = 24 h) behave like long days because they entrain the circadian system so that alternate light pulses coincide with the photoinducible phase (ø1) in castrated quail. However, while a long-day response after exposure to a single long daylength is readily detected by a rise in plasma LH (photoinduction), long-term exposure to LD 6:30 is ineffective in this respect. To discover whether this occurs because of unusual entrainment, circadian rhythms in quail and starlings were investigated. Whereas starlings entrained in the expected way with alternate pulses falling at different circadian phases, activity bouts in quail appeared to follow 24 h after successive light pulses. Because of this, activity was examined in free-running conditions to confirm that the pacemaker in quail was indeed being reset to a constant phase (reset to circadian time [CT] 0) by successive pulses. Examination of the circadian rhythms of plasma melatonin secretion under LD 6:30 also showed a resetting to CT 0. The positioning of all light pulses at the same circadian phase in the early subjective day explains the lack of photoinduction in quail since ø1 in the early subjective night phase remains unilluminated. A second feature in quail is that when the length of the photophase is gradually increased within T36h cycles, there is a progressive increase in the degree of photoinduction although the photophase length remains well below the critical daylength for induction in normal T24h cycles. We therefore tested whether ø1 is reset to a constant phase by successive pulses in LD 6:30, and that this phase is also advanced relative to light onset so that photophases shorter than the critical daylength can interact with ø1 to cause induction. Such a reduction in critical daylength relative to successive LD 6:30 pulses was confirmed by transferring quail to various types of long day and measuring the change in LH secretion. When the long-day test was replaced with continuous light, stimulation of LH secretion occurred 5-7 h earlier in quail pretreated with LD 6:30 and LD 6:54 compared to quail pretreated with LD 6:18 or LD 6:42, implying that ø 1 had been markedly phase advanced under resonance cycles.