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Many animals adjust the timing of seasonal events, such as reproduction, molt, migration, and hibernation, in response to interannual variation and directional climate-driven changes in temperature. However, the mechanisms by which temperature influences seasonal timing are relatively under-explored. Seasonal timing involves retrograde signaling in which thyrotropin (TSH) in the pars tuberalis (PT) alters expression of thyroid hormone (TH) deiodinases (Dio2 / Dio3) in tanycyte cells lining the third ventricle of the hypothalamus. This, in turn, affects the availability of triiodothyronine (T3) within the mediobasal hypothalamus—increased hypothalamic T3 restores a summer phenotype and activates the reproductive axis in long-day breeders. Recently, we showed that retrograde TH signaling is activated during late hibernation in arctic ground squirrels (Urocitellus parryii) held in constant darkness and constant ambient temperature. Sensitivity of seasonal pathways to nonphotic cues, such as temperature, is likely particularly important to hibernating species that are sequestered in hibernacula during spring. To address this issue, we exposed captive arctic ground squirrels of both sexes to an ecologically relevant increase in ambient temperature (from −6 to −1°C) late in hibernation and examined the effects of warming on the seasonal retrograde TSH/Dio/T3 signaling pathway, as well as downstream elements of the reproductive axis. We found that warmed males tended to have higher PT TSHβ expression and significantly heavier testis mass whereas the TSH/Dio/T3 signaling pathway was unaffected by warming in females, although warmed females exhibited a slight decrease in ovarian mass. Our findings suggest that temperature could have different effects on gonadal growth in male and female arctic ground squirrels, which could lead to mismatched timing in response to rapid climate change. [ABSTRACT FROM AUTHOR]

Hibernation involves prolonged intervals of profound metabolic suppression periodically interrupted by brief arousals to euthermy, the function of which is unknown. Annual cycles in mammals are timed by a photoperiodically-regulated thyroid-hormone-dependent mechanism in hypothalamic tanycytes, driven by thyrotropin (TSH) in the pars tuberalis (PT), which regulates local TH-converting deiodinases and triggers remodeling of neuroendocrine pathways. We demonstrate that over the course of hibernation in continuous darkness, arctic ground squirrels (Urocitellus parryii) up-regulate the retrograde TSH/Deiodinase/TH pathway, remodel hypothalamic tanycytes, and activate the reproductive axis. Forcing the premature termination of hibernation by warming animals induced hypothalamic deiodinase expression and the accumulation of secretory granules in PT thyrotrophs and pituitary gonadotrophs, but did not further activate the reproductive axis. We suggest that periodic arousals may allow for the transient activation of hypothalamic thyroid hormone signaling, cellular remodeling, and re-programming of brain circuits in preparation for the short Arctic summer. Arctic ground squirrels hibernating in darkness activate the pars tuberalis - hypothalamus thyroid hormone signaling pathway, remodel hypothalamic tanycytes, and activate the reproductive axis. [ABSTRACT FROM AUTHOR]

Copyright of Canadian Journal of Zoology is the property of Canadian Science Publishing and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Physical inactivity generates muscle atrophy in most mammalian species. In contrast, hibernating mammals demonstrate limited muscle loss over the prolonged intervals of immobility during winter, which suggests that they have adaptive mechanisms to reduce disuse muscle atrophy. To identify transcriptional programs that underlie molecular mechanisms attenuating muscle loss, we conducted a large-scale gene expression profiling in quadriceps muscle of arctic ground squirrels, comparing hibernating (late in a torpor and during torpor re-entry after arousal) and summer active animals using next generation sequencing of the transcriptome. Gene set enrichment analysis showed a coordinated up-regulation of genes involved in all stages of protein biosynthesis and ribosome biogenesis during both stages of hibernation that suggests induction of translation during interbout arousals. Elevated proportion of down-regulated genes involved in apoptosis, NFKB signaling as well as significant under expression of atrogenes, upstream regulators (FOXO1, FOXO3, NFKB1A), key components of the ubiquitin proteasome pathway (FBXO32, TRIM63, CBLB), and overexpression of PPARGC1B inhibiting proteolysis imply suppression of protein degradation in muscle during arousals. The induction of protein biosynthesis and decrease in protein catabolism likely contribute to the attenuation of disuse muscle atrophy through prolonged periods of immobility of hibernation. [ABSTRACT FROM AUTHOR]

Hibernation is used by a variety of mammals to survive seasonal periods of resource scarcity.Reactive oxygen species (ROS) released during periodic rewarming throughout hibernation, however, may induce oxidative damage in some tissues. Telomeres, which are the terminal sequences of linear chromosomes, may shorten in the presence ofROS, and thus the telomere length of an individual reflects the degree of accrued oxidative damage. This study quantified telomere length dynamics throughout hibernation in arctic ground squirrels (Urocitellus parryii). We hypothesized that telomere dynamics are tissue specific and predicted that telomere shortening would be most pronounced in brown adipose tissue (BAT), the organ that directly supports non-shivering thermogenesis during arousals. We used qPCR to determine relative telomere length (RTL) in DNA extracted fromliver, heart, skeletal muscle (SM) andBATof 45 juvenile and adult animals sampled either at mid- or late hibernation. Age did not have a significant effect on RTL in any tissue. At mid-hibernation, RTL of juvenile females was longer in BAT and SM than in liver and heart. In juvenile females, RTL in BAT and SM, but not in liver and heart, was shorter at late hibernation than at mid-hibernation. At late hibernation, juvenile males had longer RTL in BAT than did juvenile females, perhaps due to the naturally shorter hibernation duration of male arctic ground squirrels. Finally, BAT RTL at late hibernation negatively correlated with arousal frequency. Overall, our results suggest that, in a hibernating mammal, telomere shortening is tissue specific and that metabolically active tissues might incur higher levels of molecular damage. [ABSTRACT FROM AUTHOR]

Copyright of Canadian Journal of Zoology is the property of Canadian Science Publishing and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Although hypoxia tolerance in heterothermic mammals is well established, it is unclear whether the adaptive significance stems from hypoxia or other cellular challenge associated with euthermy, hibernation, or arousal. In the present study, blood gases, hemoglobin O2 saturation (So2), and indexes of cellular and physiological stress were measured during hibernation and euthermy and alter arousal thermogenesis. Results show that arterial O2 tension (Pao2) and So2 are severely diminished during arousal and that hypoxia inducible factor (HIF)-1α accumulates in brain. Despite evidence of hypoxia, neither cellular nor oxidative stress, as indicated by inducible nitric oxide synthase (iNOS) levels and oxidative modification of biomolecules, was observed during late arousal from hibernation. Compared with rats, hibernating Arctic ground squirrels (Spermophilus parryii) are well oxygenated with no evidence of cellular stress, inflammatory response, neuronal pathology, or oxidative modification following the period of high metabolic demand necessary for arousal. In contrast, euthermic Arctic ground squirrels experience mild, chronic hypoxia with low So2 and accumulation of HIF-1α and iNOS and demonstrate the greatest degree of cellular stress in brain. These results suggest that Arctic ground squirrels experience and tolerate endogenous hypoxia during euthermy and arousal. [ABSTRACT FROM AUTHOR]

We investigated influences of the thermal environment on patterns of body temperature (Tb), activity, and use of burrows during the active season in a population of free-living arctic ground squirrels (Spermophilus parryii). Arctic ground squirrels normally exhibited a daily 5°C range in Tb, and had higher Tb when above ground than when in burrows (P < 0.0001). This difference decreased as standard operative temperature (Tes; an index of environmental heat load) increased. Ground squirrels entered burrows more frequently on warmer compared to average or cooler days and when Tb exceeded 39°C. On cool days with heavy precipitation, ground squirrels remained almost exclusively below ground, and peaks in Tb were associated with brief aboveground forays. Time on the surface was maximal (about 80% between 0500 and 2200 h) at Tes = 17-33°C and decreased proportionately with decreasing Tes from 17°C to -2°C. Forty-six percent of variation in timing of presence above ground could be explained by a series of thermal and nonthermal variables related to environmental heat transfer. This suggests that diurnal activity patterns in this arctic environment with 24-h daylight result from a strategy that minimizes thermoregulatory costs. [ABSTRACT FROM AUTHOR]

Presents information on a study on the effects of ambient temperature on metabolic rate, respiratory quotient and torpor in arctic ground squirrels. Details on the energetic costs and substrate use in hibernation under arctic conditions; Methodology of the study; results and discussion; Conclusion.

Presents a study determining the expression pattern of uncoupling protein (Ucp) homologues and whether Ucp2 and Ucp3 mRNA levels are differentially regulated in arctic ground squirrels, maintained at different ambient temperatures and in different states of hibernation. Methodology and materials used to conduct the study; Information on the tissue disruption of Ucp1, -2, and 3- mRNA; Findings of the study.

Climate warming is rapid in the Arctic, yet impacts to biological systems are unclear because few long-term studies linking biophysiological processes with environmental conditions exist for this data-poor region. In our study spanning 25 years in the Alaskan Arctic, we demonstrate that climate change is affecting the timing of freeze-thaw cycles in the active layer of permafrost soils and altering the physiology of arctic ground squirrels (Urocitellus parryii). Soil freeze has been delayed and, in response, arctic ground squirrels have delayed when they up-regulate heat production during torpor to prevent freezing. Further, the termination of hibernation in spring has advanced 4 days per decade in females but not males. Continued warming and phenological shifts will alter hibernation energetics, change the seasonal availability of this important prey species, and potentially disrupt intraspecific interactions. [ABSTRACT FROM AUTHOR]

Social interactions can shape daily activity patterns, and this is an area of growing research interest. The propensity for individuals to be active at certain times of day may structure interactions with competitors and potential mates, influencing fitness outcomes. Aspects of daily activity such as timing of activity onset and offset exhibit within-individual repeatability, which raises the possibility that they may be subject to natural and sexual selection. This study used a biologging approach in free-living Arctic ground squirrels, Urocitellus parryii , to test the hypothesis that interactions between the sexes shape temporal patterns of daily activity. We predicted that males would become active earlier than females during the mating period, consistent with sexual selection on activity timing. We also examined repeatability in activity timing and how repeatability estimates are affected by sampling methodology. We deployed collar-mounted light loggers on ground squirrels and used timing of light transitions to determine daily onset and offset of above-ground activity. We observed 87 animals over 3 years, collecting over 7500 observations of daily activity. We found that daily activity timing had moderate (onset) to low (offset) adjusted repeatability. The difference in daily activity timing between the sexes varied seasonally: males remained active later than females during mating, while females initiated activity earlier and extended activity later during lactation. Adjusted repeatability estimates were sensitive to sampling methodology: they varied seasonally and declined with increased behavioural sampling. These results suggest that offset, not onset, may be shaped by sexual selection and that life history differences shape sex-dependent activity patterns in ground squirrels. We propose that activity onset may be under more strict circadian control than offset. This study suggests that researchers must consider seasonality and sampling design when estimating behavioural repeatability. Biologging could transform studies of individual variation by enabling high-frequency sampling of free-living animals. • Arctic ground squirrels showed evidence of circadian chronotypes. • Activity onset and offset differed by sex, especially during mating and lactation. • Activity onset reflected life-history energetics rather than sexual selection. • Repeatability estimates were affected by sampling frequency and design. [ABSTRACT FROM AUTHOR]