Circadian organization in reindeer.

The light/dark cycle of day and night synchronizes an internal 'biological clock' that governs daily rhythms in behaviour, but this form of regulation is denied to polar animals for most of the year. Here we demonstrate that the continuous lighting conditions of summer and of winter at high latitudes cause a loss in daily rhythmic activity in reindeer living far above the Arctic Circle. This seasonal absence of circadian rhythmicity may be a ubiquitous trait among resident polar vertebrates.

A Mini-Review of Ixodes Ticks Climate Sensitive Infection Dispersion Risk in the Nordic Region.

Climate change in the Nordic countries is projected to lead to both wetter and warmer seasons. This, in combination with associated vegetation changes and increased animal migration, increases the potential incidence of tick-borne diseases (TBD) where already occurring, and emergence in new places. At the same time, vegetation and animal management influence tick habitat and transmission risks. In this paper, we review the literature on Ixodes ricinus, the primary vector for TBD. Current and projected distribution changes and associated disease transmission risks are related to climate constraints and climate change, and this risk is discussed in the specific context of reindeer management. Our results indicate that climatic limitations for vectors and hosts, and environmental and societal/institutional conditions will have a significant role in determining the spreading of climate-sensitive infections (CSIs) under a changing climate. Management emerges as an important regulatory "tool" for tick and/or risk for disease transfer. In particular, shrub encroachment, and pasture and animal management, are important. The results underscore the need to take a seasonal view of TBD risks, such as (1) grazing and migratory (host) animal presence, (2) tick (vector) activity, (3) climate and vegetation, and (4) land and animal management, which all have seasonal cycles that may or may not coincide with different consequences of climate change on CSI migration. We conclude that risk management must be coordinated across the regions, and with other land-use management plans related to climate mitigation or food production to understand and address the changes in CSI risks.

Behavioral Timing without Clockwork: Photoperiod-Dependent Trade-Off between Predation Hazard and Energy Balance in an Arctic Ungulate.

Occurrence of 24-h rhythms in species apparently lacking functional molecular clockwork indicates that strong circadian mechanisms are not essential prerequisites of robust timing, and that rhythmical patterns may arise instead as passive responses to periodically changing environmental stimuli. Thus, in a new synthesis of grazing in a ruminant (MINDY), crepuscular peaks of activity emerge from interactions between internal and external stimuli that influence motivation to feed, and the influence of the light/dark cycle is mediated through the effect of low nocturnal levels of food intake on gastric function. Drawing on risk allocation theory, we hypothesized that the timing of behavior in ruminants is influenced by the independent effects of light on motivation to feed and perceived risk of predation. We predicted that the antithetical relationship between these 2 drivers would vary with photoperiod, resulting in a systematic shift in the phase of activity relative to the solar cycle across the year. This prediction was formalized in a model in which phase of activity emerges from a photoperiod-dependent trade-off between food and safety. We tested this model using data on the temporal pattern of activity in reindeer/caribou Rangifer tarandus free-living at natural mountain pasture in sub-Arctic Norway. The resulting nonlinear relationship between the phasing of crepuscular activity and photoperiod, consistent with the model, suggests a mechanism for behavioral timing that is independent of the core circadian system. We anticipate that such timing depends on integration of metabolic feedback from the digestive system and the activity of the glucocorticoid axis which modulates the behavioral responses of the animal to environmental hazard. The hypothalamus is the obvious neural substrate to achieve this integration.

Where clocks are redundant: weak circadian mechanisms in reindeer living under polar photic conditions.

Biological rhythms are a result of interplay between endogenous clocks and the ambient light-dark (LD) cycle. Biological timing in resident polar organisms presents a conundrum because these experience distinct daily LD cycles for only a few weeks each year. We measured locomotor activity in reindeer, Rangifer tarandus platyrhynchus (SR, n = 5 and 6) and R. tarandus tarandus (NR, n = 6), ranging freely at 78 and 70 degrees N, respectively, continuously throughout 1 year using data loggers. NR, but not SR, are gregarious which enabled us to examine the integrated effects of differences in social organisation and the photic environment at two different latitudes on the organisation of activity. In both sub-species, ultradian bouts of activity and inactivity alternated across the 24-h day throughout the year. This pattern was modified by the LD cycle in NR but barely at all in SR. Periodogram analysis revealed significant ultradian rhythmicity in both sub-species; the frequency of daily cycles of activity increased from three per day in winter to nearly five in summer. We conclude that this increase, and a concomitant increase in the level of daily activity, reflected the seasonal increase in the animals' appetite and the quality of their forage. Secondly, the combination, most evident in SR, of a weak photic response, weak circadian mechanisms and a weak social synchronization reduces the constraints of biological timing in an environment which is effectively non-rhythmic most of the year and permits expression of the basic ultradian pattern of ruminant activity. Third, the weaker 24-h rhythmicity in SR compared to NR indicates a latitudinal decrease in circadian organization and photic responsiveness in Rangifer.

Adaptations for life in the Arctic: evidence that melatonin rhythms in reindeer are not driven by a circadian oscillator but remain acutely sensitive to environmental photoperiod.

In reindeer Rangifer tarandus, a high latitude species, the rhythmic production of melatonin periodically dissipates under natural photoperiods when, in mid-winter, there is near permanent darkness and again, in summer, when there is permanent light. In spring and autumn, as expected, melatonin production reflects the ambient light:dark (LD) cycle. We investigated the expression of circadian mechanisms on blood levels of melatonin in reindeer. Two experiments were conducted in which animals were transferred from natural photic conditions into continuous darkness for 3 days: (i) in February, when they had been exposed to an LD cycle (11L:13D) and (ii) in July, when they had been exposed to permanent light. In July, plasma levels of melatonin rose abruptly on exposure to darkness but then declined over 24 hr before displaying a second rise and decline over the following 36 hr. In contrast, in February, levels of melatonin rose abruptly but then remained elevated for more than 60 hr in darkness. Melatonin secretion upon exposure to darkness did not conform to a circadian pattern and did not, therefore, support the hypothesis that pineal activity in reindeer is tightly regulated by circadian mechanisms. Instead the secretion of melatonin appeared to be acutely and directly sensitive to ambient lighting. The results are consistent with a model in which Arctic resident animals have adapted to extreme photic conditions by disconnecting the generation of the pineal melatonin signal from their circadian machinery and relying, instead, on its being driven by the LD cycle for just a few weeks annually in spring and autumn.