Different Phase Responses of the Two Circadian Oscillators in Gonyaulax

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Abstract
Two circadian oscillators have been previously shown to exist in the unicellular alga Gonyau lax polyedra by virtue of different periods for two different rhythms, which occur under certain experimental conditions. Here we show that phase shifts in response to light pulses also differ for these two rhythms. The phase response curves (PRCs) for white or blue light pulses are similar in shape and amplitude, but are somewhat displaced in time and have a slightly larger delay portion for the aggregation rhythm, corresponding to the A oscillator. In contrast, the aggregation rhythm PRC for dark pulses in a red light background has a much larger amplitude than that for the bioluminescence rhythm (B oscillator). These results suggest that the light input mechanism of the B oscillator is mainly blue-sensitive, whereas that of the A oscillator is sensitive to both red and blue light. This is supported by entrainment experiments: Dark pulses given as a skeleton 24-hr scotoperiod in a blue light background act as a strong zeitgeber for both oscillators. But if red light is used as background, there is bistability in the entrainment of the B oscillator, whereas the A oscillator is entrained to a definitive phase angle, regardless of the cells' prior phase of entrainment. Finally, the two oscillators can be differentially entrained with 22-hr T cycles using 3-hr dark pulses interrupting red light (19 hr), whereas both oscillators are entrained when the dark pulses interrupt blue light.