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Keeping Time Under the Midnight Sun: Behavioral and Plasma Melatonin Profiles of Free-Living Lapland Longspurs (Calcarius lapponicus) During the Arctic Summer


Noah T. Ashley, Department of Biology, Western Kentucky University, Bowling Green, KY 42101. Email:


Polar environments are characterized by discrete periods of continuous light or darkness during the summer and winter months, respectively. Because the light/dark cycle serves as the primary Zeitgeber to synchronize rhythms of most organisms, its seasonal absence in polar regions poses challenges to the circadian organization of organisms that reside in these environments. Although some species become arrhythmic, others, such as migratory songbirds, are able to maintain an intact diurnal rhythm during polar summer. This suggests that birds may switch to alternative environmental cues, such as daily changes in light intensity and ambient temperature, which may have the potential to reset the biological clock. However, identifying the low-amplitude Zeitgeber that synchronizes rhythms in free-living polar-dwelling animals has been difficult to demonstrate. In this study, we measured behavioral and melatonin profiles of free-living Lapland longspurs (Calcarius lapponicus) near Barrow, Alaska (71°N) during the continuous daylight of summer in the Arctic. Diel cycles in activity and male singing were apparent throughout the polar day with a quiescence period of 4–5 hr starting around 24:00 Alaska Daylight Time. This inactivity corresponded with elevated melatonin profiles. In contrast, territorial aggression of males in response to a conspecific intruder was not dependent upon time-of-day. Diel changes in light intensity and ambient temperature were negatively associated with daily melatonin profiles after taking into account time-of-day effects. These results suggest that photic and thermal cues may act either as alternative Zeitgeber cues, or possibly masking agents. Distinguishing between these two possibilities will require further study. J. Exp. Zool. 319A:10–22, 2013. © 2012 Wiley Periodicals, Inc.