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PubMedDate not supplied

Seasonal use of torpor by free-ranging Australian owlet-nightjars (Aegotheles cristatus).

With the exception of some data for common poorwills (Phalaenoptilus nuttallii) and anecdotal reports for a few other species, knowledge about the use of torpor by free-ranging birds is limited. Our study was designed to assess the use of torpor by free-ranging Australian owlet-nightjars (Aegotheles cristatus). We selected this species for study because of their relatively small body size (50 g), arthropod diet, nocturnal sedentary nature, taxonomic affiliation with other birds for whom the use of torpor is well documented, use of cavity roosts, and because of the cold winter (mean July minimum ambient temperature [T(a)] of approximately 0 degrees C) in the study area. We tracked 12 owlet-nightjars carrying temperature-sensitive transmitters for a total of 906 bird-days (range of 15-115 d per individual). Five different individuals entered torpor on 96 d in total. Torpor bouts occurred only between May 8 and September 8, the coldest period of the year. The lowest skin temperature (T(skin)) recorded for any bird was 19.6 degrees C, and the lowest core temperature was 22.4 degrees C. Surprisingly, torpor was rarely used at night because birds usually foraged then. Instead, torpor typically began near dawn, even on cold nights. Torpor bouts beginning at dawn lasted approximately 4 h. On 36% of days when torpor was used at dawn, birds reentered torpor later in the day. Torpor was not used during the breeding season, but this period also corresponds to the warm part of the year. There were no distinct daily minimum, maximum, or mean T(a) thresholds that could be used to reliably distinguish days when dawn torpor was used from those when it was not, although torpor was commonly employed when daily minimum T(a) fell below 3.9 degrees C. Our results show that even though Australia is typically thought of as a warm continent, at least some of the avifauna use torpor as a regular means of saving energy. We hypothesise that the reasons for this species' use of torpor include its ability to remain active all night foraging, either for terrestrial arthropods while walking or for flying insects taken on the wing, and/or its habit of roosting in cavities, which allows them to remain hidden in the daytime.

Adaptation, Physiological↗
PubMed1992

Time-dependent thresholds for torpor initiation in the rufous hummingbird (Selasphorus rufus).

Three models for torpor initiation were tested in rufous hummingbirds (Selasphorus rufus) during moult, when these birds appear to avoid the use of torpor. In model 1, the level of energy reserves at which torpor is initiated (the "threshold") remains constant throughout the night. In model 2, the threshold declines throughout the night, at a constant rate equivalent to the rate at which energy reserves are depleted during torpor. In model 3, the threshold declines at a rate equivalent to the rate of energy reserve depletion during torpor for most of the night, but at a higher rate (corresponding to the rate of energy expenditure during normothermia) during the final 2 h of the night, when these birds are usually normothermic. Model 1 predicts the most frequent and longest bouts of torpor, whereas model 3 predicts the fewest and shortest bouts. To determine the thresholds for each of 12 birds, food supply was manipulated to induce entry into torpor at different times on successive nights. Threshold slopes matched the predictions of model 3 most closely. Calculations comparing observed incidence of torpor with the predictions of model 1 show that the actual, time-dependent threshold for torpor initiation resulted in a 72% reduction in the number of torpor bouts compared with the number of torpor bouts that should have been initiated by a constant threshold. The advantage of a time-dependent threshold is that, although torpor is initiated when needed to prevent energy reserves from falling below a critical level, the amount of time spent in torpor can be minimized.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed2004

The role of polyunsaturated fatty acids in the expression of torpor by mammals: a review.

Heterothermic mammals increase the proportion of polyunsaturated fatty acids (PUFA) in their body fats prior to entering torpor. Because PUFA have low melting points, it is thought that they play an important role in maintaining the fluidity of depot fats and membrane phospholipids at low body temperatures. However, PUFA are more prone to autoxidation when exposed to reactive oxygen species (ROS) during torpor and during the periodic arousals that characterize hibernation. A lack of PUFA or an excess of PUFA may constrain the use of torpor by heterothermic mammals. We performed a mixed model meta-analysis of 17 controlled-feeding studies to test the effect of dietary PUFA on the depth and expression of torpor by daily heterotherms and hibernators. We also reviewed the literature on the PUFA content of the diet and depot fats of heterothermic mammals to address two principal topics: (1) Do low dietary levels of PUFA reduce the expression of torpor under laboratory conditions and, if so, are free-ranging animals constrained by a lack of PUFA? (2) Do high dietary levels of PUFA result in a reduction in the use, depth, and duration of torpor and, if so, do free-ranging animals seek to optimize rather than maximize PUFA intake? Low-PUFA diets consistently increase the lower setpoint for body temperature and minimum metabolic rate for both hibernators and daily heterotherms. Above the lower setpoint, low-PUFA diets usually increase body temperature and metabolic rate and decrease the duration of torpor bouts and this effect is similar for hibernators and daily heterotherms. Free-ranging rodent hibernators have dietary PUFA intakes that are far higher than those of the low-PUFA diets offered in controlled-feeding experiments, so these hibernators may never experience the constraints associated with a lack of PUFA. Diets of free-ranging insectivorous bats and echidnas have PUFA levels that are less than half as high as those offered in experimental low-PUFA diets, yet they exhibit deep and extended bouts of torpor. We argue that alternate mechanisms exist for maintaining the fluidity of body fats and that high-PUFA intake may not be a prerequisite for deep and extended bouts of torpor. Four studies indicate that animals that were fed high-PUFA diets are reluctant to enter torpor and show shallower and shorter torpor bouts. Although authors attribute this response to autoxidation, these animals did not have a higher PUFA content in their depot fats than animals where PUFA was shown to enhance torpor. We suggest that these contradictory results indicate inter-specific or inter-individual variation in the ability to control ROS and limit autoxidation of PUFA. High dietary levels of PUFA will constrain the expression of torpor only when the oxidative challenge exceeds the capacity of the antioxidant defence system. Studies of diet selection indicate that insectivorous species with low dietary PUFA levels seek to maximize PUFA intake. However, herbivorous species that have access to plants and plant parts of high-PUFA content do not appear to maximize PUFA intake. These data suggest that animals attempt to optimize rather than maximize PUFA intake. The effect of PUFA should be viewed in the light of a cost-benefit trade-off, where the benefit of high-PUFA intake is an easier access to low body temperatures and the cost is increased risk of autoxidation.

Journal Article↗
PubMed2002

Selective sleep deprivation after daily torpor in the Djungarian hamster.

Sleep, daily torpor and hibernation are no longer considered homologous processes. Animals emerging from these states spend most of their time in sleep. After termination of the torpor-associated hypothermia, there is an initial high electroenecephalogram (EEG) slow-wave activity (SWA; 0.75-4.0 Hz) and a subsequent monotonic decline. Both of these features are similar to the effects elicited by prolonged waking. It was previously shown that when hamsters are not allowed to sleep immediately after emerging from torpor, an additional SWA increase above the level reached after sleep deprivation (SD) alone occurs during the delayed recovery. A similar manipulation in hibernating ground squirrels abolished the subsequent SWA increase, shedding doubt on the similarity of the regulatory aspects following torpor and hibernation. To further investigate the extent to which SWA is homeostatically regulated after torpor, Djungarian hamsters were subjected to 1.5 h partial non-rapid eye movement (NREM) sleep deprivation (NSD) that either immediately followed the emergence from torpor (T + NSD) or 4-h SD (SD + NSD). The NSD was attained by disturbing the animals when they exhibited NREM sleep with high amplitude slow-waves. To investigate whether regional aspects of sleep homeostasis are similar after torpor and SD, the EEG was recorded from a parietal and frontal derivation after 4-h SD. An increase in SWA in NREM sleep occurred after all conditions in both EEG derivations. There was no significant difference in SWA during the initial 1.5-h recovery when torpor, T + NSD and SD + NSD were compared. During recovery from torpor and SD, SWA was higher in the frontal than in the parietal derivation. Our results provide further evidence that torpor and SD have similar effects on sleep. The SWA increase did not disappear after the NSD; therefore, SWA is homeostatically regulated after daily torpor. The frontal predominance of slow waves encountered both after torpor and SD indicates that waking and torpor induce similar regional changes in EEG SWA.

Animals↗
PubMed1998

Evolution of daily torpor and hibernation in birds and mammals: importance of body size.

1. The evolution of hibernation and daily torpor in mammals and birds remains a controversial subject. The original view was that use of torpor reflects a primitive thermoregulation, as it occurs in ancestral groups of mammals. 2. This view is no longer widely supported. However, the interpretation of a polyphyletic derivation of torpor also has been challenged because of the astonishing similarity of torpor patterns among various orders and even the two classes. 3. A recent argument is that mutations required for torpor and hibernation are unlikely to occur simultaneously and that torpor must be plesiomorphic (ancestral), although it is not functionally primitive. Homeothermy is interpreted as a loss of the ability to enter torpor in those groups that could survive without the requirement of heterothermic periods for energy conservation. 4. Interestingly, while torpor in mammals occurs in the phylogenetically old groups, lending support to the hypothesis of an ancestral derivation of torpor, the opposite is the case for birds. Modern bird groups and ancestral mammal groups contain mainly small species that often rely on fluctuating food supply, whereas modern mammalian orders and ancient bird orders contain the largest species with low energy requirements for maintenance and thermoregulation. 5. It is, therefore, possible that not phylogenetic position but size and diet determine the occurrence of heterothermy. Moreover, because endothermy and torpor in birds has apparently evolved separately from that in mammals and because it is possible that daily torpor and hibernation represent two distinct torpor patterns that evolved separately, a convergent evolution of torpor in endotherms cannot be excluded.

Animals↗
PubMed2000

Torpor, thermal biology, and energetics in Australian long-eared bats (Nyctophilus).

Previous studies have suggested that Australian long-eared bats (Nyctophilus) differ from northern-hemisphere bats with respect to their thermal physiology and patterns of torpor. To determine whether this is a general trait of Australian bats, we characterised the temporal organisation of torpor and quantified metabolic rates and body temperatures of normothermic and torpid Australian bats (Nyctophilus geoffroyi, 7 g and N. gouldi, 10 g) over a range of air temperatures and in different seasons. The basal metabolic rate of normothermic bats was 1.36 +/- 0.17 ml g(-1) h(-1) (N. geoffroyi) and 1.22 +/- 0.13 ml g(-1) h(-1) (N. gouldi), about 65% of that predicted by allometric equations, and the corresponding body temperature was about 36 degrees C. Below an air temperature of about 25 degrees C bats usually remained normothermic for only brief periods and typically entered torpor. Arousal from torpor usually occurred shortly after the beginning of the dark phase and torpor re-entry occurred almost always during the dark phase after normothermic periods of only 111 +/- 48 min (N. geoffroyi) and 115 +/- 66 min (N. gouldi). At air temperatures below 10 degrees C, bats remained torpid for more than 1 day. Bats that were measured overnight had steady-state torpor metabolic rates representing only 2.7% (N. geoffroyi) and 4.2% (N. gouldi) of the basal metabolic rate, and their body temperatures fell to minima of 1.4 and 2.3 degrees C, respectively. In contrast, bats measured entirely during the day, as in previous studies, had torpor metabolic rates that were up to ten times higher than those measured overnight. The steady-state torpor metabolic rate of thermoconforming torpid bats showed an exponential relationship with body temperature (r2 = 0.94), suggesting that temperature effects are important for reduction of metabolic rate below basal levels. However, the 75% reduction of metabolic rate between basal metabolic rate and torpor metabolic rate at a body temperature of 29.3 degrees C suggests that metabolic inhibition also plays an important role. Torpor metabolic rate showed little or no seasonal change. Our study suggests that Australian Nyctophilus bats have a low basal metabolic rate and that their patterns of torpor are similar to those measured in bats from the northern hemisphere. The low basal metabolic rate and the high proclivity of these bats for using torpor suggest that they are constrained by limited energy availability and that heterothermy plays a key role in their natural biology.

Adaptation, Physiological↗
PubMedDate not supplied

Torpor characteristics and energy requirements of furless Siberian hamsters.

After approximately 10 wk of exposure to decreasing day lengths, Siberian hamsters (Phodopus sungorus) begin to display spontaneous torpor bouts several times each week. Torpor is associated with reduced daily energy expenditure and lower food consumption and ameliorates the thermoregulatory challenges of winter. We tested the extent to which the energy savings conferred by daily torpor depend on the presence of an insulative pelage. Female hamsters were housed in a winter day length (8L:16D) at 5 degrees C; daily food intake and torpor characteristics were recorded for 5 wk in shaved (furless) or normal hamsters. Torpor-bout incidence decreased by 62% in furless hamsters, but the duration of individual bouts and the minimum body temperature attained during torpor were unaffected by loss of pelage. Body temperature declined more rapidly during entry into torpor and increased more slowly during arousal from torpor in furless than in control hamsters. Energy savings per torpor bout, assessed by the amount of food consumed on days that included a torpor bout, was substantially greater in normal than in furless hamsters (16.0% vs. 3.3%); this difference likely reflects the increased cost of thermoregulation during torpor, as well as the increased caloric expenditure incurred by furless hamsters during arousal from torpor. An insulative pelage may be a prerequisite for the energetic benefits derived from heterothermy in this species.

Animals↗
PubMedDate not supplied

Respiratory exchange and ventilation during nocturnal torpor in hummingbirds.

In mammals that undergo torpor and hibernation, a period of CO2 storage is often a prelude to, and may be required for, the onset of a bout of torpor. Storage of CO2 has been hypothesized to induce an energy-conserving metabolic suppression in torpid mammals. It is unclear whether CO2 storage also occurs in birds that undergo torpor. To study these questions, we measured rates of oxygen consumption (VO2) and carbon dioxide production (VCO2) during normothermy, entrance into torpor, steady-state torpor, and spontaneous arousal in Selasphorus hummingbirds. The respiratory exchange ratio (RER) declined from about 0.86 to about 0.71 in both normothermic and torpid individuals, suggesting a shift from carbohydrate oxidation to lipid oxidation during the night. Torpor metabolism ranged from 5% to 42% of normothermic values. In most individuals that became torpid, a brief period of CO2 storage occurred during entrance into torpor, beginning after VO2 had fallen considerably below normothermic rates. The volume of CO2 stored averaged 0.048 mL CO2 g-1, somewhat less than reported for mammalian hibernators. Some individuals entered torpor without measurable CO2 storage. There was no abrupt change in RER at the start of arousal, although mean arousal RER exceeded the RER during steady-state torpor. The change in metabolic rate between normothermia and torpor had a Q10 of 1.9-2.2 in birds that did not regulate body temperature in torpor. We conclude that a normal Q10 effect is sufficient to explain the metabolic rate of torpid hummingbirds and that transient CO2 storage may occur during entrance into torpor but is not a prerequisite for entrance. Similarly, CO2 release is not a prerequisite for arousal.

Animals↗
PubMed1991

Effects of temperature, steroids and castration on daily torpor in the Djungarian hamster (Phodopus sungorus).

1. In Phodopus sungorus the frequency and characteristics of daily torpor were not affected by a decrease in temperature. The seasonal cycle of daily torpor is essentially under photoperiodic control. 2. Testicular regression is a necessary condition for daily torpor occurrence. Testosterone and 5 alpha-dihydrotestosterone administration totally inhibited daily torpor in hamsters which were exposed to short days. The temporal inhibition of torpor by steroids had no effect on the seasonal time measuring mechanism governing daily torpor. 3. Castration influenced certain aspects of the daily torpor display. Castrated hamsters showed a delay in terminating torpor season lending further support to the hypothesis that recrudescence of the testis plays a role in terminating the daily torpor season. Castration, when performed before or on the day of transfer to short photoperiod led to an increase in the frequency of torpor bouts and reduce SP exposure duration which is necessary for the daily torpor season beginning.

Animals↗
PubMed2000

Metabolism and temperature regulation during daily torpor in the smallest primate, the pygmy mouse lemur (Microcebus myoxinus) in Madagascar.

Thermoregulation, energetics and patterns of torpor in the pygmy mouse lemur, Microcebus myoxinus, were investigated under natural conditions of photoperiod and temperature in the Kirindy/CFPF Forest in western Madagascar. M. myoxinus entered torpor spontaneously during the cool dry season. Torpor only occurred on a daily basis and torpor bout duration was on average 9.6 h, and ranged from 4.6 h to 19.2 h. Metabolic rates during torpor were reduced to about 86% of the normothermic value. Minimum body temperature during daily torpor was 6.8 degrees C at an ambient temperature of 6.3 degrees C. Entry into torpor occurred randomly between 2000 and 0620 hours, whereas arousals from torpor were clustered around 1300 hours within a narrow time window of less than 4 h. Arousal from torpor was a two-step process with a first passive climb of body temperature to a mean of 27 degrees C, carried by the daily increase of ambient temperature when oxygen consumption remained more or less constant, followed by a second active increase of oxygen consumption to further raise the body temperature to normothermic values. In conclusion, daily body temperature rhythms in M. myoxinus further reduce the energetic costs of daily torpor seen in other species: they extend to unusually low body temperatures and consequently low metabolic rates in torpor, and they employ passive warming to reduce the energetic costs of arousal. Thus, these energy-conserving adaptations may represent an important energetic aid to the pygmy mouse lemur and help to promote their individual fitness.

Animals↗
PubMedDate not supplied

Daily torpor in free-ranging rock elephant shrews, Elephantulus myurus: a year-long study.

Under laboratory conditions, rock elephant shrews, Elephantulus myurus, use daily torpor under both short and long photoperiod acclimation. However, use of heterothermy often differs under field and laboratory conditions. We investigated the use of torpor in free-ranging elephant shrews from May 2001 to May 2002. The elephant shrews were capable of daily torpor throughout the year, with torpor most prevalent during winter. We recorded two torpor bouts during early summer (November). We recorded a total of 467 torpor bouts during the year. The mean torpor minimum body temperature (Tbmin) for the whole year was 15.3 degrees +/-4.4 degrees C, and the mean bout length was 8.6+/-3.5 h. These values were in the range expected for daily heterotherms. However, there was some marginal overlap with hibernation characteristics; a few torpor bouts were longer than 24 h in duration, and Tbmin decreased below 10 degrees C. Torpor was highly correlated with low ambient temperature and photoperiod. Torpor was also correlated with invertebrate abundance after controlling for photoperiod effects. During the year in which this study was conducted, the rainfall was 14% below long-term average. Historical rainfall records show that summer rainfall during strong El Nino years is up to 40% below the long-term average. During these drought years, the frequency of summer torpor may be higher, highlighting the need for long-term physiological data in free-ranging animals.

Animals↗
PubMed2003

Sleep regulation in the Djungarian hamster: comparison of the dynamics leading to the slow-wave activity increase after sleep deprivation and daily torpor.

STUDY OBJECTIVES: Emerging from daily torpor, Djungarian hamsters (Phodopus sungorus) show an initial increase in electroencephalographic slow-wave activity (power density between 0.75 and 4.0 Hz) during sleep that gradually declines. This feature is typical for sleep following prolonged waking and supports the hypothesis that sleep pressure increases during daily torpor. After hamsters were subjected to sleep deprivation or partial non-rapid eye movement sleep deprivation immediately following torpor, slow-wave activity remained high and decreased only when sleep was allowed. An analysis of the dynamics of the process underlying the build-up of sleep pressure during episodes of waking and torpor may provide insights into the regulation of normal sleep and wakefulness. We have analyzed in more detail the timecourse of the process that is common for waking and daily torpor and that could account for the subsequent increase in slow-wave activity. DESIGN: Continuous 24-hour recordings of electroencephalography, electromyography, cortical temperature, and electroencephalographic spectral analysis were performed. Torpor data of 28 hamsters and sleep-deprivation data of diverse durations collected previously in 15 hamsters were analyzed. SETTING: N/A. PATIENTS OR PARTICIPANTS: N/A. INTERVENTIONS: Sleep deprivation. MEASUREMENTS AND RESULTS: Slow-wave activity invariably increased as a function of the duration of both prior waking and torpor. However, the time constant of the build-up of slow-wave activity was approximately 2.75 times slower during torpor compared to sleep deprivation. Brain temperature recorded during the torpor bouts was 10 degrees to 12 degrees C below euthermic brain temperature. Therefore, the temperature coefficient of the time constant for the slow-wave-activity increase is between 2.3 and 2.8, a range typical for biochemical processes. CONCLUSIONS: We conclude that the processes occurring during daily torpor in the Djungarian hamster are similar to those occurring during sleep deprivation, but the build-up of sleep pressure during torpor appears to be slowed down by the lower brain temperature.

Animals↗
PubMed2005

Development of thermoregulation and torpor in a marsupial: energetic and evolutionary implications.

Altricial mammals and birds become endothermic at about half the size of adults and presumably would benefit energetically from entering torpor at that time. Because little is known about torpor during development in endotherms, we investigated whether after the establishment of endothermic thermoregulation (i.e. the ability to maintain a high body temperature during cold exposure), Sminthopsis macroura, a small (approximately 25 g) insectivorous marsupial, is capable of entering torpor and whether torpor patterns change with growth. Endothermic thermoregulation was established when the nest young reached a body mass of approximately 10 g, and they were capable of entering torpor early during development at approximately 10-12 g, lending some support to the view that torpor is a phylogenetically old mammalian trait. Torpor bout length shortened significantly and the minimum metabolic rate during torpor increased as juveniles approached adult size, and consequently total daily energy expenditure increased steeply with age. Relationships between total daily energy expenditure and body mass during development of S. macroura (slope approximately 1.3) differed substantially from the relationship between basal metabolism and body mass in adult endotherms (slope approximately 0.75) suggesting that the energy expenditure-size relationship during the development differs substantially from that in adults under thermo-neutral conditions. Our study shows that while torpor can substantially reduce energy expenditure during development of endotherms and hence is likely important for survival during energy bottlenecks, it also may enhance somatic growth when food is limited. We therefore hypothesize that torpor during the development in endotherms is far more widespread than is currently appreciated.

Animals↗
PubMed2006

Daily torpor alters multiple gene expression in the suprachiasmatic nucleus and pineal gland of the Djungarian hamster (Phodopus sungorus).

Circadian rhythms are still expressed in animals that display daily torpor, implying a temperature compensation of the pacemaker. Nevertheless, it remains unclear how the clock works in hypothermic states and whether torpor itself, as a temperature pulse, affects the circadian system. To reveal changes in the clockwork during torpor, we compared clock gene and neuropeptide expression by in situ hybridization in the suprachiasmatic nucleus (SCN) and pineal gland of normothermic and torpid Djungarian hamsters (Phodopus sungorus). Animals from light-dark (LD) 8ratio16 were sacrificed at 8 time points throughout 24 h. To investigate the effect of a previous torpor episode on the clock, we sacrificed a group of normothermic hamsters 1 day after torpor. In normothermic animals, Per1 peaked at zeitgeber time (ZT)4; whereas, Bmal1 reached maximal expression between ZT16 and ZT19. AVP mRNA in the SCN showed highest levels at ZT7. On the day of torpor, the levels of all mRNAs investigated, except for AVP mRNA, were increased during the torpor bout. Moreover, the Bmal1 rhythm was advanced. On the day after the hypothermia, Bmal1 and AVP rhythms showed severely depressed amplitude. Those distinct amplitude changes of Bmal1 and AVP on the day after a torpor episode expression suggests that torpor affects the circadian system, probably by altered translational processes that might lead to a modified protein feedback on gene expression. In the pineal gland, an important clock output, Aanat expression, peaked between ZT16 and ZT22 in normothermic animals. Aanat levels were significantly advanced on the day of hypothermia, an effect which was still visible 1 day afterward. In summary, this study showed that daily torpor affects the phase and amplitude of rhythmic clock gene and clock-controlled gene expression in the SCN. Furthermore, the rhythmic gene expression in a peripheral oscillator, the pineal gland, is also affected.

ARNTL Transcription Factors↗
PubMed2004

Sleep and circadian rhythms in mammalian torpor.

Sleep and circadian rhythms are the primary determinants of arousal state, and torpor is the most extreme state change that occurs in mammals. The view that torpor is an evolutionary extension of sleep is supported by electrophysiological studies. However, comparisons of factors that influence the expression of sleep and torpor uncover significant differences. Deep sleep immediately following torpor suggests that torpor is functionally a period of sleep deprivation. Recent studies that employ post-torpor sleep deprivation, however, show that the post-torpor intense sleep is not homeostatically regulated, but might be a reflection of synaptic loss and replacement. The circadian system regulates sleep expression in euthermic mammals in such a way that would appear to preclude multiday bouts of torpor. Indeed, the circadian system is robust in animals that show shallow torpor, but its activity in hibernators is at least damped if not absent. There is good evidence from some species, however, that the circadian system plays important roles in the timing of bouts of torpor.

Animals↗
PubMed2000

Corticosterone and nocturnal torpor in the rufous hummingbird (Selasphorus rufus).

Three experiments were designed to investigate whether corticosterone (CORT), known to have a role in restoration of energy homeostasis, regulates nocturnal torpor, an energy conservation state used by some small mammals and birds to offset environmental challenges to energy balance. In two experiments, one during autumn migration and one during early spring molt, captive rufous hummingbirds (Selasphorus rufus) were fed control and dilute (85% strength) nectar on alternate days. In migratory birds, torpor occurred more frequently over all, and nectar dilution resulted in increased torpor duration and increased concentration of CORT in evening but not midday cloacal fluid (CF) samples. In molting birds, torpor occurred infrequently on both control and food dilution days, but, although there was a significant increase in evening CF CORT on food dilution days, torpor duration did not increase significantly in response and there was no correlation between torpor duration and CF CORT at either time of day. Daily CF CORT patterns showed an increase from midday to evening during migration, but the reverse pattern during the molt. In a third experiment, CORT administered in the nectar elevated the use of torpor and depressed food intake. The results of these three experiments support the hypothesis that CORT is involved in the regulation of torpor, but suggest that some feature of the CORT signal other than concentration per se may be required to fully explain seasonal changes in the relations among energy challenge, CORT, and nocturnal torpor in hummingbirds.

Animals↗
PubMed1988

Reduction of metabolism during hibernation and daily torpor in mammals and birds: temperature effect or physiological inhibition?

The present study addresses the controversy of whether the reduction in energy metabolism during torpor in endotherms is strictly a physical effect of temperature (Q10) or whether it involves an additional metabolic inhibition. Basal metabolic rates (BMR; measured as oxygen consumption, VO2), metabolic rates during torpor, and the corresponding body temperatures (Tb) in 68 mammalian and avian species were assembled from the literature (n = 58) or determined in the present study (n = 10). The Q10 for change in VO2 between normothermia and torpor decreased from a mean of 4.1 to 2.8 with decreasing Tb from 30 to less than 10 degrees C in hibernators (species that show prolonged torpor). In daily heterotherms (species that show shallow, daily torpor) the Q10 remained at a constant value of 2.2 as Tb decreased. In hibernators with a Tb less than 10 degrees C, the Q10 was inversely related to body mass. The increase of mass-specific metabolic rate with decreasing body mass, observed during normothermia (BMR), was not observed during torpor in hibernators and the slope relating metabolic rate and mass was almost zero. In daily heterotherms, which had a smaller Q10 than the hibernators, no inverse relationship between the Q10 and body mass was observed, and consequently the metabolic rate during torpor at the same Tb was greater than that of hibernators. These findings show that the reduction in metabolism during torpor of daily heterotherms and large hibernators can be explained largely by temperature effects, whereas a metabolic inhibition in addition to temperature effects may be used by small hibernators to reduce energy expenditure during torpor.

Animals↗
PubMed2003

Defining torpor in free-ranging bats: experimental evaluation of external temperature-sensitive radiotransmitters and the concept of active temperature.

A variety of definitions involving body temperature (Tb), metabolic rate and behavior have been used to define torpor in mammals and birds. This problem is confounded in some studies of free-ranging animals that employ only skin temperature (Tsk), a measure that approximates but may not precisely reflect Tb. We assess the accuracy of Tsk in the context of a recent definition for torpor called active temperature. We compared the active temperatures of individual big brown bats (Eptesicus fuscus), which aggregate in cavities, with solitary, foliage-roosting hoary bats (Lasiurus cinereus). In captive big brown bats, we compared Tsk and core Tb at a range of ambient temperatures for clustered and solitary roosting animals, compared Tsk and Tb during arousal from torpor, and quantified the effect of flight on warming from torpor. Hoary bats had significantly lower active temperatures than big brown bats despite having the same normothermic Tsk. Tsk was significantly lower than Tb during normothermia but often greater than Tb during torpor. Flight increased the rate of warming from torpor. This effect was more pronounced for Tsk than Tb. This suggests that bats could rely on heat generated by flight muscles to complete the final stages of arousal. Using active temperature to define torpor may underestimate torpor due to ambient cooling of external transmitters or animals leaving roosts while still torpid. Conversely, active temperature may also overestimate shallow torpor use if it is recorded during active arousal when shivering and non-shivering thermogenesis warm external transmitters. Our findings illuminate the need for laboratory studies that quantify the relationship between metabolic rate and Tsk over a range of ambient temperatures.

Acclimatization↗