Torpor Bouts Research Articles

Ambient Temperature Cycles Affect Daily Torpor and Hibernation Patterns in Malagasy Tenrecs.

Hibernation and daily torpor (heterothermy) allow endotherms to cope with demanding environmental conditions. The depth and duration of torpor bouts vary considerably between tropical and temperate climates, and tropical hibernators manage to cope with a wider spectrum of ambient temperature (Ta) regimes during heterothermy. As cycles in Ta can have profound effects on activity and torpor patterns as well as energy expenditure, we examined how these characteristics are affected by daily fluctuating versus constant Ta in a tropical hibernator, the lesser hedgehog tenrec (Echinops telfairi). Throughout the study, regardless of season, the tenrecs became torpid every day. In summer, E. telfairi used daily fluctuations in Ta to passively rewarm from daily torpor, which led to synchrony in the activity phases and torpor bouts between individuals and generally decreased energy expenditure. In contrast, animals housed at constant Ta showed considerable variation in timing and they had to invest more energy through endogenous heat production. During the hibernation season (winter) E. telfairi hibernated for several months in constant, as well as in fluctuating Ta and, as in summer, under fluctuating Ta arousals were much more uniform and showed less variation in timing compared to constant temperature regimes. The timing of torpor is not only important for its effective use, but synchronization of activity patterns could also be essential for social interactions, and successful foraging bouts. Our results highlight that Ta cycles can be an effective zeitgeber for activity and thermoregulatory rhythms throughout the year and that consideration should be given to the choice of temperature regime when studying heterothermy under laboratory conditions.

The physiological signature of daily torpor is not orexin dependent.

Under conditions of scarce food availability and cool ambient temperature, the mouse (Mus Musculus) enters into torpor, a state of transient metabolic suppression mediated in part by the autonomic nervous system. Hypothalamic orexins are involved in the coordination of behaviors and autonomic function. We tested whether orexins are necessary for the coordinated changes in physiological variables, which underlie torpor and represent its physiological signature. We performed simultaneous measurements of brain temperature, electroencephalographic, and electromyographic activity allowing objective assessment of wake-sleep behavior, and cardiovascular, respiratory, and metabolic variables in orexin knockout mice (ORX-KO) and wild-type mice (WT) during torpor bouts elicited by caloric restriction and mild cold stress. We found that torpor bouts in WT are characterized by an exquisitely coordinated physiological signature. The characteristics of torpor bouts in terms of duration and rate of change of brain temperature and electromyographic activity at torpor entrance and exit did not differ significantly between ORX-KO and WT, and neither did the cardiovascular, respiratory, and metabolic characteristics of torpor. ORX-KO and WT also had similar wake-sleep state changes associated with torpor bouts, with the exception of a significantly higher rapid-eye movement sleep time in ORX-KO at torpor entrance. Our results demonstrate that orexins are not necessary either for the normal physiological adaptations occurring during torpor in mice or for their coordination, suggesting that mechanisms different from orexin peptide signaling may be involved in the regulation and the coordination of these physiological responses.

Hummingbird torpor in context: duration, more than temperature, is the key to nighttime energy savings

Torpor is an important energy saving strategy in some small birds, but it has rarely been studied in natural field conditions. We compared torpor use across 43 wild‐caught individuals of eight hummingbird species across sites with different natural temperature regimes. Most laboratory studies focus on the relationship between metabolic rate and temperature, but our aim was to evaluate what environmental factors most influence hummingbird nighttime energy management under natural conditions. We found that the probability of an individual entering torpor was weakly correlated with mass but unrelated to nighttime temperature and that hummingbirds at both warm, tropical and cooler, temperate sites used torpor. Energy savings in torpor were maximized as ambient temperatures approached a species’ minimum body temperature, consistent with laboratory studies; energy savings ranged between 65 and 92% of energy per hour in torpor compared to normothermy. However, regardless of the degree of energy savings in torpor, variation in total nighttime energy expenditure was most significantly influenced by torpor bout duration. Lab studies largely assess the effect of temperature on torpor use, but our findings indicate that other environmental conditions are more important in determining hummingbirds’ total nighttime energy expenditure under natural temperature cycles. Our results show that a small endotherm's nighttime energy management in its natural habitat is more affected by torpor bout duration, which is linked to photoperiod, than by temperature. This result suggests that in their natural environments hummingbirds are able to save energy in torpor across a range of nighttime temperatures, indicating that they may have sufficient physiological flexibility to tolerate climatic variation.

Cold-inducible RNA-binding protein Cirp, but not Rbm3, may regulate transcript processing and protection in tissues of the hibernating ground squirrel.

RNA-binding proteins (RBPs) have important roles in transcription, pre-mRNA processing/transport, mRNA degradation, translation, and non-coding RNA processing, among others. RBPs that are expressed in response to cold stress, such as Cirp and Rbm3, could regulate RNA stability and translation in hibernating mammals that reduce their body temperatures from 37°C to as low as 0-5°C during torpor bouts. RBPs including Cirp, Rbm3, and stress-inducible HuR translocate from the nucleus to stabilize mRNAs in the cytoplasm, and thereby could regulate which mRNA transcripts are protected from degradation and are translated, versus stored, for future protein synthesis or degraded by nucleases during cell stress associated with metabolic rate depression. This is the first study to explore the transcriptional/translational regulation, and subcellular localization of cold-inducible RBPs in a model hibernator, the 13-lined ground squirrel (Ictidomys tridecemlineatus). Cirp protein levels were upregulated in liver, skeletal muscle, and brown adipose tissue throughout the torpor-arousal cycle whereas Rbm3 protein levels stayed constant or decreased, suggesting an important role for Cirp, but likely not Rbm3, in the hibernator stress response. Increased cytoplasmic localization of Cirp in liver and muscle and HuR in liver during torpor, but no changes in the relative levels of Rbm3 in the cytoplasm, emphasizes a role for Cirp and possibly HuR in regulating mRNA processing during torpor. This study informs our understanding of the natural adaptations that extreme animals use in the face of stress, and highlight natural stress response mediators that could be used to bolster cryoprotection of human organs donated for transplant.

Do small precocial birds enter torpor to conserve energy during development?

Precocial birds hatch feathered and mobile, but when they become fully endothermic soon after hatching, their heat loss is high and they may become energy depleted. These chicks could benefit from using energy-conserving torpor, which is characterised by controlled reductions of metabolism and body temperature (Tb). We investigated at what age the precocial king quail Coturnix chinensis can defend a high Tb under a mild thermal challenge and whether they can express torpor soon after achieving endothermy to overcome energetic and thermal challenges. Measurements of surface temperature (Ts) using an infrared thermometer showed that king quail chicks are partially endothermic at 2-10 days, but can defend high Tb at a body mass of ∼13 g. Two chicks expressed shallow nocturnal torpor at 14 and 17 days for 4-5 h with a reduction of metabolism by >40% and another approached the torpor threshold. Although chicks were able to rewarm endogenously from the first torpor bout, metabolism and Ts decreased again by the end of the night, but they rewarmed passively when removed from the chamber. The total metabolic rate increased with body mass. All chicks measured showed a greater reduction of nocturnal metabolism than previously reported in quails. Our data show that shallow torpor can be expressed during the early postnatal phase of quails, when thermoregulatory efficiency is still developing, but heat loss is high. We suggest that torpor may be a common strategy for overcoming challenging conditions during development in small precocial and not only altricial birds.

Disease recovery in bats affected by white-nose syndrome.

Processes associated with recovery of survivors are understudied components of wildlife infectious diseases. White-nose syndrome (WNS) in bats provides an opportunity to study recovery of disease survivors, understand implications of recovery for individual energetics, and assess the role of survivors in pathogen transmission. We documented temporal patterns of recovery from WNS in little brown bats (Myotis lucifugus) following hibernation to test the hypotheses that: (1) recovery of wing structure from WNS matches a rapid time scale (i.e. approximately 30 days) suggested by data from free-ranging bats; (2) torpor expression plays a role in recovery; (3) wing physiological function returns to normal alongside structural recovery; and (4) pathogen loads decline quickly during recovery. We collected naturally infected bats at the end of hibernation, brought them into captivity, and quantified recovery over 40 days by monitoring body mass, wing damage, thermoregulation, histopathology of wing biopsies, skin surface lipids and fungal load. Most metrics returned to normal within 30 days, although wing damage was still detectable at the end of the study. Torpor expression declined overall throughout the study, but bats expressed relatively shallow torpor bouts - with a plateau in minimum skin temperature - during intensive healing between approximately days 8 and 15. Pathogen loads were nearly undetectable after the first week of the study, but some bats were still detectably infected at day 40. Our results suggest that healing bats face a severe energetic imbalance during early recovery from direct costs of healing and reduced foraging efficiency. Management of WNS should not rely solely on actions during winter, but should also aim to support energy balance of recovering bats during spring and summer.

Optimal hibernation theory

AbstractHibernation has received considerable attention from physiologists and natural historians, but theoretical and ecological treatments of hibernation are rarer. There is ample recent evidence that costs associated with hibernation affect the degree to which hibernation is expressed in nature, but we currently lack a quantitative framework under which to make predictions about how the costs and benefits of hibernation interact under various environmental conditions. Here, we attempt the first steps towards building an optimal hibernation theory for making specific predictions about the expression of hibernation (i.e. the depth and duration of torpor bouts), metabolic functioning, and the total period of hibernation in mammals and birds. Our current understanding of the costs associated with hibernation do not allow for parameterisation of optimal hibernation theory, but we hope this work provides a roadmap for physiologists and ecologists to collect the necessary data in the future.

The evolution of a bat population with white-nose syndrome (WNS) reveals a shift from an epizootic to an enzootic phase

BackgroundWhite-nose Syndrome (WNS) is a mycosis caused by a cutaneous infection with the fungus Pseudogymnoascus destructans (Pd). It produces hibernation mortality rates of 75–98% in 4 bats: Myotis lucifugus, M. septentrionalis, M. sodalis, and Perimyotis subflavus. These high mortality rates were observed during the first several years after the arrival of P. destructans at a hibernation site. Mortality is caused by a 60% decrease in torpor bout duration, which results in a premature depletion of depot fat prior to spring.ResultsLittle is known about the long-term effects of Pd on torpor and mortality, thus we conducted a 9-year study on M. lucifugus at 5 of the hibernation sites where Pd first appeared in North America during the winter of 2007–08. The M. lucifugus hibernating at one of these sites one year after the arrival of Pd (2008–09) had: a) a mean torpor bout duration of 7.6 d, b) no depot fat reserves by March, and c) an apparent over-winter mortality rate of 88%. The M. lucifugus hibernating at this same site 6–9 years after the arrival of Pd, in contrast, had: a) a mean torpor bout duration of 14.7 d, b) depot fat remaining in March, and c) an apparent mortality rate of 50%. The number of M. lucifugus hibernating at 2 of these sites has consistently increased since 2010 and is now more than 3.0-fold higher than the number remaining after the winter of 2008–09.ConclusionsThese findings indicate that this population of M. lucifugus has evolved mechanisms to hibernate well in the presence of Pd, thus reducing over-winter mortality.

Leptin: Is It Thermogenic?

Animals that lack the hormone leptin become grossly obese, purportedly for 2 reasons: increased food intake and decreased energy expenditure (thermogenesis). This review examines the experimental evidence for the thermogenesis component. Analysis of the data available led us to conclude that the reports indicating hypometabolism in the leptin-deficient ob/ob mice (as well as in the leptin-receptor-deficient db/db mice and fa/fa rats) derive from a misleading calculation artefact resulting from expression of energy expenditure per gram of body weight and not per intact organism. Correspondingly, the body weight-reducing effects of leptin are not augmented by enhanced thermogenesis. Congruent with this, there is no evidence that the ob/ob mouse demonstrates atrophied brown adipose tissue or diminished levels of total UCP1 mRNA or protein when the ob mutation is studied on the inbred C57BL/6 mouse background, but a reduced sympathetic nerve activity is observed. On the outbred “Aston” mouse background, brown adipose tissue atrophy is seen, but whether this is of quantitative significance for the development of obesity has not been demonstrated. We conclude that leptin is not a thermogenic hormone. Rather, leptin has effects on body temperature regulation, by opposing torpor bouts and by shifting thermoregulatory thresholds. The central pathways behind these effects are largely unexplored.

Perineuronal net expression in the brain of a hibernating mammal.

During hibernation, mammals like the 13-lined ground squirrel cycle between physiological extremes. Most of the hibernation season is spent in bouts of torpor, where body temperature, heart rate, and cerebral blood flow are all very low. However, the ground squirrels periodically enter into interbout arousals (IBAs), where physiological parameters return to non-hibernating levels. During torpor, neurons in many brain regions shrink and become electrically quiescent, but reconnect and regain activity during IBA. Previous work showed evidence of extracellular matrix (ECM) changes occurring in the hypothalamus during hibernation that could be associated with this plasticity. Here, we examined expression of a specialized ECM structure, the perineuronal net (PNN), in the forebrain of ground squirrels in torpor, IBA, and summer (non-hibernating). PNNs are known to restrict plasticity, and could be important for retaining essential connections in the brain during hibernation. We found PNNs in three regions of the hypothalamus: ventrolateral hypothalamus, paraventricular nucleus (PVN), and anterior hypothalamic area. We also found PNNs throughout the cerebral cortex, amygdala, and lateral septum. The total area covered by PNNs within the PVN was significantly higher during IBA compared to non-hibernating and torpor (P < 0.01). Additionally, the amount of PNN coverage area per Nissl-stained neuron in the PVN was significantly higher in hibernation compared to non-hibernating (P < 0.05). No other significant differences were found across seasons. The PVN is involved in food intake and homeostasis, and PNNs found here could be essential for retaining vital life functions during hibernation.

Incorporating evaporative water loss into bioenergetic models of hibernation to test for relative influence of host and pathogen traits on white-nose syndrome.

Hibernation consists of extended durations of torpor interrupted by periodic arousals. The ‘dehydration hypothesis’ proposes that hibernating mammals arouse to replenish water lost through evaporation during torpor. Arousals are energetically expensive, and increased arousal frequency can alter survival throughout hibernation. Yet we lack a means to assess the effect of evaporative water loss (EWL), determined by animal physiology and hibernation microclimate, on torpor bout duration and subsequent survival. White-nose syndrome (WNS), a devastating disease impacting hibernating bats, causes increased frequency of arousals during hibernation and EWL has been hypothesized to contribute to this increased arousal frequency. WNS is caused by a fungus, which grows well in humid hibernaculum environments and damages wing tissue important for water conservation. Here, we integrated the effect of EWL on torpor expression in a hibernation energetics model, including the effects of fungal infection, to determine the link between EWL and survival. We collected field data for Myotis lucifugus, a species that experiences high mortality from WNS, to gather parameters for the model. In saturating conditions, we predicted healthy bats experience minimal mortality. Infected bats, however, suffer high fungal growth in highly saturated environments, leading to exhaustion of fat stores before spring. Our results suggest that host adaptation to humid environments leads to increased arousal frequency from infection, which drives mortality across hibernaculum conditions. Our modified hibernation model provides a tool to assess the interplay between host physiology, hibernaculum microclimate, and diseases such as WNS on winter survival.

Alternate-day feeding leads to improved glucose regulation on fasting days without significant weight loss in genetically obese mice.

Alternate-day fasting (ADF) is effective for weight loss and increases insulin sensitivity in diet-induced obese rodents. However, the efficacy of ADF in genetic models of obesity has not been comprehensively studied. Mice that are deficient in leptin (ob/ob mice) are obese, diabetic, and prone to deep bouts of torpor when fasted. We tested the hypotheses that an ADF protocol in ob/ob mice would result in 1) induction of torpor on fasted days, 2) minimal body weight loss if the mice experienced torpor, and 3) no improvement in glucose control in the absence of weight loss. Female ob/ob mice and littermate controls were assigned to 1) an ad libitum regimen or 2) an ADF regimen, consisting of fasting every other day with ad libitum feeding between fasts. Over a 19-day period, littermate control mice on the ADF regimen consumed the same amount of food as littermate control mice on the ad libitum regimen, whereas the ADF ob/ob mice consumed 37% less food than ad libitum ob/ob mice. Fasting days, but not fed days, led to torpor in both genotypes. Fasting days, but not fed days, led to weight loss in both genotypes relative to ad libitum controls. Fasting days, but not fed days, produced enhanced insulin sensitivity in both genotypes and normalized circulating glucose in ob/ob mice. These data demonstrate improved glucose control on fasting days with the use of ADF in a genetic model of obesity in the face of minimal weight loss.

Frequent nocturnal torpor in a free-ranging Australian honeyeater, the noisy miner.

Torpor in birds is considered to be far less common than in mammals. This is particularly true for passerine birds for which knowledge of torpor expression is scarce, although almost all are small, have high energy expenditure and could profit energetically from using torpor. To assess whether the extent and diversity of avian and especially passerine torpor expression and heterothermy may be currently underestimated because of limited long-term data on free-ranging birds, core body temperature fluctuations were quantified over ~ 4.3months in a medium-sized honeyeater, the noisy miner (Manorina melanocephala, ~ 75g), in an open woodland during the cold season in eastern Australia. Miners used shallow nocturnal torpor frequently (63% of days), torpor bouts lasted on average for 6.5h (maximum 13.5h) and, unlike during hypothermia, torpor was terminated by endogenous heat production for rewarming. Body temperatures (Tb) ranged from a maximum of 43.5°C to a minimum of 33.0°C, often fell by 7°C at night, and the overall mean Tb was 38.7 ± 0.7°C. The data show that yet another passerine bird, widely viewed to be homeothermic, expresses torpor in the wild for energy conservation. Considering the size of miners, it seems probable that many other, especially smaller birds, use a similar approach at least in winter to enhance the chance of survival in the face of high energy expenditure and low food availability.

Thermoneutral temperature reduces liver volume but increases fat content in a mammalian hibernator.

Hibernators survive challenging winters by entering torpor, which lowers body temperature (Tb) to ∼5 °C for 12-14 days, followed by spontaneous arousals where Tb increases to ∼37 °C for 10-12 h before entering another torpor bout. This Tb cycle is accompanied by significant fluctuations in metabolic rate. Little is known about the role of the liver in lipid metabolism during hibernation. In this study we measured the effect of ambient temperature on liver volume and lipid content in 13-lined ground squirrels (Ictidomys tridecemlineatus). We housed animals at thermoneutral (25 °C) or cold (5 °C) ambient temperatures, with the same photoperiod (12 h light:12 h dark) for an entire year. We determined volume and water-fat ratio of the liver using magnetic resonance imaging (MRI). Ambient temperature significantly affected both liver volume and fat content. From October to August squirrels housed at 25 °C had 25% smaller livers compared to the squirrels housed at 5 °C, but their average lipid content (13.3%) was 37% higher. Because the squirrels housed at 25 °C appeared to continue feeding throughout the winter but did not enter extended torpor, more carbohydrates may have been diverted to lipid stores. By contrast, animals housed at 5 °C did not appear to feed, and carbohydrates would likely be preferentially stored in the liver as glycogen to supply glucose for brain metabolism. These results suggest that the fat burden caused by hibernators preparing for winter can lead to symptoms of metabolic syndrome, but that these symptoms are reversible in the spring.

Numerous cold arousals and rare arousal cascades as a hibernation strategy in European Myotis bats.

Hibernating bats optimise the duration of torpor bouts and arousals in relation to hibernaculum microclimatic conditions and fat reserves. Clustering has significant physiological and ecological benefits, promoting successful hibernation of individuals. Such aggregations may help maintain optimal temperatures, allowing better energy utilisation than in solitarily bats. However, aroused bats in a cluster could conceivably disturb those still hibernating, starting an energy-demanding arousal process. Our study was conducted over two winters in two different hibernacula (cave and mine) in the Czech Republic, where Greater mouse-eared bats (Myotis myotis) have previously been diagnosed with white-nose syndrome. In 118 arousal episodes we recorded 193 individual arousals in which a warming phase was observed, 135 (69.9%) being cold arousals, where bats ceased increasing their body temperatures at ≤ 10 °C. The remaining arousals were standard normothermic arousals, where body (fur) surface temperatures reached > 20 °C. Cold arousals occurred during the mid- and late hibernation periods, suggesting they were a response to disturbance by a neighbour in the same cluster. Arousal cascades, where bats aroused in series, were rare (12.7%) and reached a maximum in mid-January. Our data suggest that Myotis bats prolong their torpor bouts using numerous cold arousals but few arousal cascades. Upon arrival of a bat, the clustered bats show tolerance to disturbing by conspecifics.

Lipidomics Reveals Seasonal Shifts in a Large-Bodied Hibernator, the Brown Bear.

Prior to winter, heterotherms retain polyunsaturated fatty acids (“PUFA”), resulting in enhanced energy savings during hibernation, through deeper and longer torpor bouts. Hibernating bears exhibit a less dramatic reduction (2–5°C) in body temperature, but lower their metabolism to a degree close to that of small hibernators. We determined the lipid composition, via lipidomics, in skeletal muscle and white adipose tissues (“WAT”), to assess lipid retention, and in blood plasma, to reflect lipid trafficking, of winter hibernating and summer active wild Scandinavian brown bears (Ursus arctos). We found that the proportion of monounsaturated fatty acids in muscle of bears was significantly higher during winter. During hibernation, omega-3 PUFAs were retained in WAT and short-length fatty acids were released into the plasma. The analysis of individual lipid moieties indicated significant changes of specific fatty acids, which are in line with the observed seasonal shift in the major lipid categories and can be involved in specific regulations of metabolisms. These results strongly suggest that the shift in lipid composition is well conserved among hibernators, independent of body mass and of the animals’ body temperature.

Analysis of an Omega 6:3 Balanced Diet Impact on Hibernation Behavior and Cortisol of Captive Arctic Ground Squirrels

Arctic Ground Squirrels (AGS) are widely studied in laboratories for their capability to sustain low metabolic rates and temperatures without enduring debilitating bodily harm. A specific range of omega 6 polyunsaturated fatty acids (PUFAs) has been shown to improve captive hibernation by increasing torpor bout length and depth of hibernation across multiple species (Geiser 1993, Frank 2008). To our knowledge, only three studies have investigated the impact of omega 3 fatty acids on hibernation behavior with varying results (Giroud 2018, Frank 2004 and Hill and Florant 2000). Our prior findings show wild AGS plasma have a balanced omega 6 to 3 fatty acid profile (1.3:1), but no one, to our knowledge, has tested the impact of this specific PUFA ratio on captive AGS. We hypothesize that substituting a high omega 6:3 (8:1) diet for a balanced diet (1.3:1) would increase the length of torpor bouts, entrance timing into hibernation and decrease cortisol.Wild juvenile AGS (12 f, 12 m) were evenly distributed to two groups: a balanced omega 6:3 ratio diet (1.3:1) and a control diet of Mazuri rodent chow with an omega 6:3 ratio of (8:1). Animals were fed 47g daily of their diets until first torpor bout and food was withdrawn. Animals were instrumented with abdominal temperature I‐buttons in early August. White adipose tissue (WAT) and plasma were sampled longitudinally at fall (early August at the time of surgery) and winter (mid‐December at time of euthanasia). Animals were euthanized in three stages of the torpor bout cycle: early torpor, late torpor, and full arousal.WAT was analyzed a modified lipid dense tissue protocol using solid phase extraction. Cortisol was measured using RIA. Early August levels of cortisol were 1.4× higher in the animals supplemented with the balanced omega 6:3 diet (p&lt;0.0001). Preliminary temperature data indicates that the balanced diet and control diet AGS do not significantly differ in the number of hibernation days, entrance timing into hibernation or torpor bout length. Additionally, preliminary data indicates a significant increase in body weight of females on a balanced omega 6:3 diet during fall (p&lt;0.013) and winter (p&lt;0.049).Preliminary data indicates the balanced diet does not negatively impact hibernation behavior. The unexpected increase in cortisol in the balanced omega 6:3 ratio fed diet may reflect normal physiological levels rather than stress. Cortisol functions as a stress indicator, but also enhances fat deposition, which is an essential task prior to hibernation.Support or Funding InformationThis work was supported by NIH 20GM103395, UL1GM118991, TL4GM118992, or RL5GM118990. This work was performed in conformance with the FASEB Statement of Principles for animals used in research.This abstract is from the Experimental Biology 2019 Meeting. There is no full text article associated with this abstract published in The FASEB Journal.

Evaluating seasonal changes of cone photoreceptor structure in the 13-lined ground squirrel

Cone photoreceptors of the 13-lined ground squirrel (13-LGS) undergo reversible structural changes during hibernation, including cone outer segment disc degeneration and inner segment mitochondria depletion. Here, we evaluated cone structure with adaptive optics scanning light ophthalmoscopy (AOSLO) before, during, and after hibernation. Also, intra-animal comparisons of cone structure were made at distinct physiological states (pre-hibernation, torpor, interbout euthermia, and post-hibernation) with AOSLO and transmission electron microscopy. Our results indicate that the 13-LGS cone mosaic is only transiently affected by structural remodeling during hibernation. Outer segment remodeling starts during torpid states during a period of fall transition in room temperature, with more severe structural changes during bouts of torpor in cold temperature. Cones return to euthermic-like structure during brief periods of interbout euthermia and recover normal waveguiding properties as soon as 24 h post-hibernation. Cone structure is visible with split-detector AOSLO throughout hibernation, providing evidence that intact outer segments are not necessary to visualize cones with this technique. Despite the changes to cone structure during hibernation, cone density and packing remained unchanged throughout the seasonal cycle. Pairing non-invasive imaging with ultrastructural assessment may provide insight to the biological origins of cone photoreceptor signals observed with AOSLO.

Metabolomic Study of Hibernating Syrian Hamster Brains: In Search of Neuroprotective Agents.

Syrian hamsters undergo a reversible hyperphosphorylation of protein τ during hibernation, providing a unique natural model that may unveil the physiological mechanisms behind this critical process involved in the development of Alzheimer's disease and other tauopathies. The hibernation cycle of these animals fluctuates between a pair of stages: 3-4 days of torpor bouts interspersed with periods of euthermia called arousals that last several hours. In this study, we investigated for the first time the metabolic changes in brain tissue during hibernation. A total of 337 metabolites showed statistically significant differences during hibernation. Based on these metabolites, several pathways were found to be significantly regulated and, therefore, play a key role in the regulation of hibernation processes. The increase in the levels of ceramides containing more than 20 C atoms was found in torpor animals, reflecting a higher activity of CerS2 during hibernation, linked to neurofibrillary tangle generation and structural changes in the Golgi apparatus. Our results open up the debate about the possible significance of some metabolites during hibernation, which may possibly be related to τ phosphorylation and dephosphorylation events. In general, this study may provide insights into novel neuroprotective agents because the alterations described throughout the hibernation process are reversible.

Hibernation and daily torpor in Australian and New Zealand bats: does the climate zone matter?

We aim to summarise what is known about torpor use and patterns in Australian and New Zealand (ANZ) bats from temperate, tropical/subtropical and arid/semiarid regions and to identify whether and how they differ. ANZ bats comprise ~90 species from 10 families. Members of at least nine of these are known to use torpor, but detailed knowledge is currently restricted to the pteropodids, molossids, mystacinids, and vespertilionids. In temperate areas, several species can hibernate (use a sequence of multiday torpor bouts) in trees or caves mostly during winter and continue to use short bouts of torpor for the rest of the year, including while reproducing. Subtropical vespertilionids also use multiday torpor in winter and brief bouts of torpor in summer, which permit a reduction in foraging, probably in part to avoid predators. Like temperate-zone vespertilionids they show little or no seasonal change in thermal energetics during torpor, and observed changes in torpor patterns in the wild appear largely due to temperature effects. In contrast, subtropical blossom-bats (pteropodids) exhibit more pronounced daily torpor in summer than winter related to nectar availability, and this involves a seasonal change in physiology. Even in tropical areas, vespertilionids express short bouts of torpor lasting ~5 h in winter; summer data are not available. In the arid zone, molossids and vespertilionids use torpor throughout the year, including during desert heat waves. Given the same thermal conditions, torpor bouts in desert bats are longer in summer than in winter, probably to minimise water loss. Thus, torpor in ANZ bats is used by members of all or most families over the entire region, its regional and seasonal expression is often not pronounced or as expected, and it plays a key role in energy and water balance and other crucial biological functions that enhance long-term survival by individuals.