Lipid composition of the stratum corneum in different regions of the body of Kuhl's pipistrelle from the Negev Desert, Israel.

The most superficial epidermal layer in endotherms is the stratum corneum (SC), which is composed of dead corneocytes embedded in a lipid matrix with free fatty acids, cholesterol, ceramides, and cerebrosides; the lipid composition of the SC determines its permeability to water vapor. Lipids that are more polar, have longer hydrocarbon chains, and are less bulky are often packed in more ordered phase states to slow cutaneous evaporative water loss (CEWL); these lipids also resist transitions to more disordered phases at high ambient temperatures (Ta). In bats, wing and tail membranes (wing patagia and tail uropatagium, respectively) allow powered flight, but increase surface area, and hence CEWL, with implications for survival in arid environments. We captured Pipistrellus kuhlii from an arid habitat and measured the lipid composition of the SC of the plagiopatagium in the wing, the uropatagium, and the non-membranous region (NMR) of the body using thin layer chromatography and reversed phase high performance liquid chromatography coupled with atmospheric pressure photoionization mass spectrometry. The patagia contained more cholesterol and shorter-chained ceramides, and fewer cerebrosides than the NMR, indicating that the lipid phase transition temperature in the patagia is lower than in the NMR. Thus, at moderate Ta the lipids in the SC in all body regions will remain in an ordered phase state, allowing water conservation; but as Ta increases, the lipids in the SC of the patagia will more easily transition into a disordered phase, resulting in increased CEWL from the patagia facilitating efficient heat dissipation in hot environments.

Evaporative water loss in Kuhl's pipistrelles declines along an environmental gradient, from mesic to hyperarid.

Intraspecific variation in animal energy and water balances may play an important role in local adaptation of populations to specific habitats such as deserts. We examined Kuhl's pipistrelle (Pipistrellus kuhlii), a common bat in Israel that ranges in distribution from mesic Mediterranean to hyperarid desert habitats, for intraspecific differences in metabolic rate (MR) and evaporative water loss (EWL) among populations along a climatic gradient. We tested the prediction that EWL, especially at high ambient temperatures is lower in Kuhl's pipistrelles from desert habitats than from mesic habitats. We measured MR and total evaporative water loss (TEWL) at four ambient temperatures (10 °C, 20 °C, 30 °C and 35 °C) in three groups of bats using open-flow respirometry. We fitted the bats with a mask to separate cutaneous water loss (CWL) from respiratory water loss (RWL) at 35 °C. At 35 °C, mean TEWL in the southernmost group, from the hyperarid location, was significantly lower than in the other two groups, with no apparent difference in mean MR. The source of difference TEWL was that the southern group had significantly lower CWL than the other two groups; RWL did not differ among them. This suggests that there are mechanisms that reduce EWL from the skin of the bats; a likely candidate is modification of the lipids in the outer layer of the dermis that make the skin possibly less permeable to water as has been described in birds and a few other species of bat.

The cutaneous lipid composition of bat wing and tail membranes: a case of convergent evolution with birds.

The water vapour permeability barrier of mammals and birds resides in the stratum corneum (SC), the outermost layer of the epidermis. The molar ratio and molecular arrangement of lipid classes in the SC determine the integrity of this barrier. Increased chain length and polarity of ceramides, the most abundant lipid class in mammalian SC, contribute to tighter packing and thus to reduced cutaneous evaporative water loss (CEWL). However, tighter lipid packing also causes low SC hydration, making it brittle, whereas high hydration softens the skin at the cost of increasing CEWL. Cerebrosides are not present in the mammalian SC; their pathological accumulation occurs in Gaucher's disease, which leads to a dramatic increase in CEWL. However, cerebrosides occur normally in the SC of birds. We tested the hypothesis that cerebrosides are also present in the SC of bats, because they are probably necessary to confer pliability to the skin, a quality needed for flight. We examined the SC lipid composition of four sympatric bat species and found that, as in birds, their SC has substantial cerebroside contents, not associated with a pathological state, indicating convergent evolution between bats and birds.

Similar burrow architecture of three arid-zone scorpion species implies similar ecological function.

Many animals reside in burrows that may serve as refuges from predators and adverse environmental conditions. Burrow design varies widely among and within taxa, and these structures are adaptive, fulfilling physiological (and other) functions. We examined the burrow architecture of three scorpion species of the family Scorpionidae: Scorpio palmatus from the Negev desert, Israel; Opistophthalmus setifrons, from the Central Highlands, Namibia; and Opistophthalmus wahlbergii from the Kalahari desert, Namibia. We hypothesized that burrow structure maintains temperature and soil moisture conditions optimal for the behavior and physiology of the scorpion. Casts of burrows, poured in situ with molten aluminum, were scanned in 3D to quantify burrow structure. Three architectural features were common to the burrows of all species: (1) a horizontal platform near the ground surface, long enough to accommodate the scorpion, located just below the entrance, 2-5 cm under the surface, which may provide a safe place where the scorpion can monitor the presence of potential prey, predators, and mates and where the scorpion warms up before foraging; (2) at least two bends that might deter incursion by predators and may reduce convective ventilation, thereby maintaining relatively high humidity and low temperature; and (3) an enlarged terminal chamber to a depth at which temperatures are almost constant (±2-4 °C). These common features among the burrows of three different species suggest that they are important for regulating the physical environment of their inhabitants and that burrows are part of scorpions' "extended physiology" (sensu Turner, Physiol Biochem Zool 74:798-822, 2000).

Metabolic rate, evaporative water loss and thermoregulatory state in four species of bats in the Negev desert.

Life in deserts is challenging for bats because of their relatively high energy and water requirements; nevertheless bats thrive in desert environments. We postulated that bats from desert environments have lower metabolic rates (MR) and total evaporative water loss (TEWL) than their mesic counterparts. To test this idea, we measured MR and TEWL of four species of bats, which inhabit the Negev desert in Israel, one species mainly restricted to hyper-arid deserts (Otonycteris hemprichii), two species from semi-desert areas (Eptesicus bottae and Plecotus christii), and one widespread species (Pipistrellus kuhlii). We also measured separately, in the same individuals, the two components of TEWL, respiratory water loss (RWL) and cutaneous evaporative water loss (CEWL), using a mask. In all the species, MR and TEWL were significantly reduced during torpor, the latter being a consequence of reductions in both RWL and CEWL. Then, we evaluated whether MR and TEWL in bats differ according to their geographic distributions, and whether those rates change with Ta and the use of torpor. We did not find significant differences in MR among species, but we found that TEWL was lowest in the species restricted to desert habitats, intermediate in the semi-desert dwelling species, and highest in the widespread species, perhaps a consequence of adaptation to life in deserts. Our results were supported by a subsequent analysis of data collected from the literature on rates of TEWL for 35 bat species from desert and mesic habitats.

Transient-state mechanisms of wind-induced burrow ventilation.

Burrows are common animal habitations, yet living in a burrow presents physiological challenges for its inhabitants because the burrow isolates them from sources and sinks for oxygen, carbon dioxide, water vapor and ammonia. Conventionally, the isolation is thought to be overcome by either diffusion gas exchange within the burrow or some means of capturing wind energy to power steady or quasi-steady bulk flows of air through it. Both are examples of what may be called 'DC' models, namely steady to quasi-steady flows powered by steady to quasi-steady winds. Natural winds, however, are neither steady nor quasi-steady, but are turbulent, with a considerable portion of the energy contained in so-called 'AC' (i.e. unsteady) components, where wind velocity varies chaotically and energy to power gas exchange is stored in some form. Existing DC models of burrow gas exchange do not account for this potentially significant source of energy for ventilation. We present evidence that at least two AC mechanisms operate to ventilate both single-opening burrows (of the Cape skink, Trachylepis capensis) and double-opening model burrows (of Sundevall's jird, Meriones crassus). We propose that consideration of the physiological ecology and evolution of the burrowing habit has been blinkered by the long neglect of AC ventilation.

Flea fitness is reduced by high fractional concentrations of CO2 that simulate levels found in their hosts' burrows.

Nidicolous ectoparasites such as fleas and gamasid mites that feed on small and medium-sized mammals spend much of their time in their hosts' burrows, which provide an environment for living, and often feeding, to their pre-imaginal and/or adult stages. Thus, these ectoparasites should be adapted to environmental conditions in burrows, including high fractional concentrations of CO2 (F(CO2)). We examined how a high F(CO2) (0.04) affected survival and reproductive success of a hematophagous ectoparasite of burrowing rodents using fleas Xenopsylla ramesis and Sundevall's jirds Meriones crassus. In the first experiment, fleas fed on hosts housed in high-CO2 (F(CO2) =0.04) or atmospheric-CO2 (F(CO2) ≈0.0004) air, and were allowed to breed. In a second experiment, fleas were maintained in high CO2 or CO2-free air with no hosts to determine how CO2 levels affect survival and activity levels. We found that at high F(CO2) fleas laid fewer eggs, reducing reproductive success. In addition, at high F(CO2), activity levels and survival of fleas were reduced. Our results indicate that fleas do not perform well in the F(CO2) used in this experiment. Previous research indicated that the type and intensity of the effects of CO2 concentration on the fitness of an insect depend on the F(CO2) used, so we advise caution when generalizing inferences drawn to insects exposed to other F(CO2). If, however, F(CO2) found in natural mammal burrows brings about reduced fitness in fleas in general, then burrowing hosts may benefit from reduced parasite infestation if burrow air F(CO2) is high.

Ventilation of multi-entranced rodent burrows by boundary layer eddies.

Rodent burrows are often assumed to be environments wherein the air has a high concentration of CO2. Although high burrow [CO2] has been recorded, many studies report burrow [CO2] that differs only slightly from atmospheric concentrations. Here, we advocate that one of the reasons for these differences is the penetration into burrows of air gusts (eddies), which originate in the turbulent boundary layer and prevent build-up of CO2. We have characterized the means by which burrows of Sundevall's jird, which are representative of the burrows of many rodent species with more than one entrance, are ventilated. Our results demonstrate that, even at low wind speeds, the random penetration of eddies into a burrow through its openings is sufficient to keep the burrow [CO2] low enough to be physiologically inconsequential, even in its deep and remote parts.

Age at weaning, immunocompetence and ectoparasite performance in a precocial desert rodent.

We studied the effects of early weaning on immunocompetence and parasite resistance in a precocial rodent Acomys cahirinus. We hypothesized that if parasite resistance is energetically expensive and nutritional and immunological support from mothers are necessary for the long-term health of offspring, then early weaned animals would be immunologically weaker and less able to defend themselves against parasites than later weaned animals. We weaned pups at 14, 21 or 28 days after birth and assessed their immunocompetence and resistance against fleas Parapulex chephrenis when they attained adulthood. Immunocompetence was assessed using leukocyte concentration (LC) and a phytohaemagglutinin injection assay (PHA test). To estimate resistance against fleas, we measured performance of fleas via the number of produced eggs and duration of development and resistance to starvation of the flea offspring. We found a significant positive effect of weaning age on the PHA response but not on LC. The effect of age at weaning on flea egg production was manifested in male but not female hosts, with egg production being higher if a host was weaned at 14 than at 28 days. Weaning age of the host did not affect either duration of development or resistance to starvation of fleas produced by mothers fed on these hosts. We conclude that even in relatively precocial mammals, weaning age is an important indicator of future immunological responses and the ability of an animal to resist parasite infestations. Hosts weaned at an earlier age make easier, less-resistant targets for parasite infestations than hosts weaned later in life.

Waking to drink: rates of evaporative water loss determine arousal frequency in hibernating bats.

Bats hibernate to cope with low ambient temperatures (T(a)) and low food availability during winter. However, hibernation is frequently interrupted by arousals, when bats increase body temperature (T(b)) and metabolic rate (MR) to normothermic levels. Arousals account for more than 85% of a bat's winter energy expenditure. This has been associated with variation in T(b), T(a) or both, leading to a single testable prediction, i.e. that torpor bout length (TBL) is negatively correlated with T(a) and T(b). T(a) and T(b) were both found to be correlated with TBL, but correlations alone cannot establish a causal link between arousal and T(b) or T(a). Because hydration state has also been implicated in arousals from hibernation, we hypothesized that water loss during hibernation creates the need in bats to arouse to drink. We measured TBL of bats (Pipistrellus kuhlii) at the same T(a) but under different conditions of humidity, and found an inverse relationship between TBL and total evaporative water loss, independent of metabolic rate, which directly supports the hypothesis that hydration state is a cue to arousal in bats.

Ambient temperature and nutritional stress influence fatty acid composition of structural and fuel lipids in Japanese quail (Coturnix japonica) tissues.

In birds, fatty acids (FA) serve as the primary metabolic fuel during exercise and fasting, and their composition affects metabolic rate and thus energy requirements. To ascertain the relationship between FAs and metabolic rate, a distinction should be made between structural and fuel lipids. Indeed, increased unsaturation of structural lipid FAs brings about increased cell metabolism, and changes in the FA composition of fuel lipids affects metabolic rate through selective mobilization and increasing availability of specific FAs. We examined the effects of acclimation to a low ambient temperature (Ta: 12.7±3.0°C) and nutritional status (fed or unfed) on the FA composition of four tissues in Japanese quail, Coturnix japonica. Differentiating between neutral (triglycerides) and polar (phospholipids) lipids, we tested the hypothesis that both acclimation to low Ta and nutritional status modify FA composition of triglycerides and phospholipids. We found that both factors affect FA composition of triglycerides, but not the composition of phospholipids. We also found changes in liver triacylglyceride FA composition in the low-Ta acclimated quail, namely, the two FAs that differed, oleic acid (18:1) and arachidonic acid (20:4), were associated with thermoregulation. In addition, the FAs that changed with nutritional status were all reported to be involved in regulation of glucose metabolism, and thus we suggest that they also play a role in the response to fasting.

Asthma and hemoglobinopathy: when is supplemental oxygen required?

Asthma is the most common reason for referral to the emergency department in childhood. In severe attacks, supplemental O2 is given when oxygen saturation level is <90%. Described herein is the case of a child with persistent low oxygen saturation as measured on pulse oximetry (S(p)O2) after full clinical recovery from an asthma attack. Simultaneously, P(a)O2 was normal. A diagnosis of abnormal hemoglobin with decreased oxygen affinity (hemoglobin Seattle) was made on hemoglobin electrophoresis and genetic analysis. To ascertain when supplemental oxygen was needed, an oxygen dissociation curve was plotted using the tonometer technique, and it was found that an S(p)O2 of 70% is parallel to a P(a)O2 of 60 mmHg. Plotting an oxygen dissociation curve is a simple reproducible method to determine when supplemental oxygen is required for a child with a hemoglobinopathy.

Physiological responses to food deprivation in the house sparrow, a species not adapted to prolonged fasting.

Many wild birds fast during reproduction, molting, migration, or because of limited food availability. Species that are adapted to fasting sequentially oxidize endogenous fuels in three discrete phases. We hypothesized that species not adapted to long fasts have truncated, but otherwise similar, phases of fasting, sequential changes in fuel oxidization, and similar changes in blood metabolites to fasting-adapted species. We tested salient predictions in house sparrows (Passer domesticus biblicus), a subspecies that is unable to tolerate more than ~32 h of fasting. Our main hypothesis was that fasting sparrows sequentially oxidize substrates in the order carbohydrates, lipids, and protein. We dosed 24 house sparrows with [(13)C]glucose, palmitic acid, or glycine and measured (13)CO(2) in their breath while they fasted for 24 h. To ascertain whether blood metabolite levels reflect fasting-induced changes in metabolic fuels, we also measured glucose, triacylglycerides, and ß-hydroxybutyrate in the birds' blood. The results of both breath (13)CO(2) and plasma metabolite analyses did not support our hypothesis; i.e., that sparrows have the same metabolic responses characteristic of fasting-adapted species, but on a shorter time scale. Contrary to our main prediction, we found that recently assimilated (13)C-tracers were oxidized continuously in different patterns with no definite peaks corresponding to the three phases of fasting and also that changes in plasma metabolite levels accurately tracked the changes found by breath analysis. Notably, the rate of recently assimilated [(13)C]glycine oxidization was significantly higher (P < 0.001) than that of the other metabolic tracers at all postdosing intervals. We conclude that the inability of house sparrows to fast for longer than 32 h is likely related to their inability to accrue large lipid stores, separately oxidize different fuels, and/or spare protein during fasting.

Effects of host diet and thermal state on feeding performance of the flea Xenopsylla ramesis.

We examined feeding performance of the flea Xenopsylla ramesis on three different hosts: its natural, granivorous, rodent host, Sundevall's jird (Meriones crassus); the frugivorous Egyptian fruit bat (Rousettus aegyptiacus); and an insectivorous bat, Kuhl's pipistrelle (Pipistrellus kuhlii). Because these fleas are not known to occur on bats, we hypothesized that the fleas' feeding performance (i.e. feeding and digestion rates) would be higher when feeding on their natural host than on either of the bats that they do not naturally parasitize. We found that mass-specific blood-meal size of both male and female fleas was significantly lower when feeding on Kuhl's pipistrelles than on the other two species, but was not different in female fleas feeding on fruit bats or on jirds at all stages of digestion. However, more male fleas achieved higher levels of engorgement if they fed on Sundevall's jirds than if they fed on Egyptian fruit bats. The fleas digested blood of fruit bats and jirds significantly faster than blood of Kuhl's pipistrelle. In addition, after a single blood meal, the survival time of fleas fed on normothermic Kuhl's pipistrelles was significantly shorter than that of fleas fed on Sundevall's jirds and even lower when male fleas fed on Egyptian fruit bats. Thus, our prediction was partially supported: normothermic Kuhl's pipistrelles were inferior hosts for fleas compared with Sandevall's jirds and Egyptian fruit bats. Interestingly, the proportion of engorged fleas that fed on torpid Kuhl's pipistrelles was significantly higher than the proportion of the fleas that fed on normothermic individuals, indicating that becoming torpid might be a liability, rather than an effective defense against parasites.

Allocation of endogenous and dietary protein in the reconstitution of the gastrointestinal tract in migratory blackcaps at stopover sites.

During migratory flight, the mass of the gastrointestinal tract (GIT) and its associated organs in small birds decreases in size by as much as 40%, compared with the preflight condition because of the catabolism of protein. At stopover sites, birds need 2-3 days to rebuild their GIT so that they can restore body mass and fat reserves to continue migration. The source of protein used to rebuild the GIT may be exogenous (from food ingested) or endogenous (reallocated from other organs) or both. Because the relative contribution of these sources to rebuild the GIT of migratory birds is not yet known, we mimicked in-flight fasting and then re-feeding in two groups of blackcaps (Sylvia atricapilla), a Palearctic migratory passerine. The birds were fed a diet containing either 3% or 20% protein to simulate different refueling scenarios. During re-feeding, birds received known doses of (15)N-(l)-leucine before we measured the isotope concentrations in GIT and associated digestive organs and in locomotory muscles. We then quantified the extent to which blackcaps rebuilt their GIT with endogenous and/or dietary protein while refeeding after a fast. Our results indicate that blackcaps fed the low-protein diet incorporated less exogenous nitrogen into their tissues than birds fed the 20% protein diet. They also allocated relatively more exogenous protein to the GIT than to pectoral muscle than those birds re-fed with the high-protein diet. However, this compensation was not sufficient for birds eating the low-protein diet to rebuild their intestine at the same rate as the birds re-fed the high-protein diet. We concluded that blackcaps must choose stopover sites at which they can maximize protein intake to minimize the time it takes to rebuild their GIT and, thus, resume migration as soon as possible.

Reduced body mass gain in small passerines during migratory stopover under simulated heat wave conditions.

For birds that migrate long distances, maximizing the rate of refueling at stopovers is advantageous, but ambient conditions may adversely influence this vital process. We simulated a 3-day migratory stopover for garden warblers (Sylvia borin) and compared body temperatures (T(b)) and rates of refueling under conditions of a heat wave (T(a)=40 °C by day, and 15 °C at night) with those under more moderate conditions (T(a)=27 °C by day, and 15 °C at night). We measured T(b) with implanted thermo-sensitive radio transmitters. Birds had significantly lower rates of body mass gain on the first day of stopover (repeated measures mixed model ANOVA, p=0.002) affecting body mass during the entire stopover (p=0.034) and higher maximum T(b) during the day when exposed to high T(a) than when exposed to moderate T(a) (p=0.002). In addition, the birds exposed to high T(a) by day had significantly lower minimum T(b) at night than those exposed to moderate daytime T(a) (p=0.048), even though T(a) at night was the same for both groups. We interpret this lower nighttime T(b) to be a means of saving energy to compensate for elevated daytime thermoregulatory requirements, while higher T(b) by day may reduce protein turnover. All effects on T(b) were significantly more pronounced during the first day of stopover than on days two and three, which may be linked to the rate of renewal of digestive function during stopover. Our results suggest that environmental factors, such as high T(a), constrain migratory body mass gain. Extreme high T(a) and heat waves are predicted to increase due to global climate change, and thus are likely to pose increasing constraints on regaining body mass during stopover and therefore migratory performance in migratory birds.

Air Temperature and Humidity at the Bottom of Desert Wolf Spider Burrows Are Not Affected by Surface Conditions.

Burrows are animal-built structures that can buffer their occupants against the vagaries of the weather and provide protection from predators. We investigated whether the trapdoors of wolf spider (Lycosa sp.; temporary working name "L. hyraculus") burrows in the Negev Desert serve to maintain favorable environmental conditions within the burrow by removing trapdoors and monitoring the ensuing temperature and relative humidity regime within them. We also monitored the behavioral responses of "L. hyraculus" to trapdoor removal at different times of the day and in different seasons. "L. hyraculus" often spun silk mesh in their burrow entrances in response to trapdoor removal during the day, possibly to deter diurnal predators. The frequency of web-spinning peaked on summer mornings, but spiders began spinning webs sooner after trapdoor removal later in the day. In addition, we monitored temperature and relative humidity in artificial burrows in the summer during the morning and at midday. At noon, air temperature (Ta) at the bottom of open burrows increased by <1 °C more than in covered burrows, but water vapor pressure in burrows did not change. The relatively small increase in Ta in uncovered burrows at midday can probably be ascribed to the penetration of direct solar radiation. Thus, air temperature and humidity at the bottom of the burrow are apparently decoupled from airflow at the surface.

Sucrose digestion capacity in birds shows convergent coevolution with nectar composition across continents.

The major lineages of nectar-feeding birds (hummingbirds, sunbirds, honeyeaters, flowerpiercers, and lorikeets) are considered examples of convergent evolution. We compared sucrose digestion capacity and sucrase enzymatic activity per unit intestinal surface area among 50 avian species from the New World, Africa, and Australia, including 20 nectarivores. With some exceptions, nectarivores had smaller intestinal surfaces, higher sucrose hydrolysis capacity, and greater sucrase activity per unit intestinal area. Convergence analysis showed high values for sucrose hydrolysis and sucrase activity per unit intestinal surface area in specialist nectarivores, matching the high proportion of sucrose in the nectar of the plants they pollinate. Plants pollinated by generalist nectar-feeding birds in the Old and New Worlds secrete nectar in which glucose and fructose are the dominant sugars. Matching intestinal enzyme activity in birds and nectar composition in flowers appears to be an example of convergent coevolution between plants and pollinators on an intercontinental scale.

Fasting triggers hypothermia, and ambient temperature modulates its depth in Japanese quail Coturnix japonica.

We tested three hypotheses regarding the cues that elicit facultative hypothermia in Japanese quail (Coturnix japonica): H(1)) Ambient temperature (T(a)), alone, influences the onset and depth of hypothermia; H(2)) Fasting, alone, influences the onset and depth of hypothermia; H(3)) T(a) acts synergistically with fasting to shape the use of hypothermia. Eight quail were maintained within their thermoneutral zone (TNZ) at 32.6+/-0.2 degrees C, and eight below their lower critical temperature (T(lc)) at 12.7+/-3.0 degrees C. All quail entered hypothermia upon food deprivation, even quail kept within their TNZ. Body temperature (T(b)) decreased more (38.36+/-0.53 degrees C vs. 39.57+/-0.57 degrees C), body mass (m(b)) loss was greater (21.0+/-7.20 g vs.12.8+/-2.62g), and the energy saved by using hypothermia was greater (25.18-45.01% vs. 7.98-28.06%) in low the T(a) treatment than in TNZ treatment. Interestingly, the depth of hypothermia was positively correlated with m(b) loss in the low T(a) treatment, but not in TNZ treatment. Our data support H(3), that both thermoregulatory costs and body energy reserves are proximate cues for entry into hypothermia in quail. This outcome is not surprising below the T(lc). However, the quail kept at their TNZ also responded to food deprivation by entering hypothermia with no apparent dependence on m(b) loss. Therefore inputs, other than thermoregulatory costs and body condition, must serve as cues to enter hypothermia. Consequently, we address the role that tissue sparing may play in the physiological 'decision' to employ hypothermia.

Heterothermy in small, migrating passerine birds during stopover: use of hypothermia at rest accelerates fuel accumulation.

For small endothermic animals, heterothermy serves as an energy-saving mechanism for survival in challenging environments, but it may also accelerate fat accumulation in individuals preparing for fuel-demanding activities. This is the first study to demonstrate adaptive hypothermic responses in migrating passerines. While monitoring body temperature (T(b)) of eight blackcaps (Sylvia atricapilla) by radiotelemetry, we found that during daytime T(b)=42.5+/-0.4 degrees C (mean +/- s.d.); at night T(b) decreased to a minimum between 33 and 40 degrees C. We determined the lower limit for normothermy at 37.4 degrees C and found that on 12 out of 34 bird-nights of observations under semi-natural conditions blackcaps reduced their T(b) below normothermic resting levels with minimum values of 33 and 34.5 degrees C compared with rest-phase normothermic T(b) of 38.8+/-0.8 degrees C. In birds of body mass (m(b)) <16.3 g, minimum T(b) at night correlated with the individual's m(b) (r=0.67, P<0.01, N=17), but this was not the case in birds with m(b)>16.3 g. Minimum nocturnal T(b) did not correlate with night-time air temperature (T(a)). Measurements of metabolic rate in birds subjected to a T(a) of 15 degrees C showed that hypothermia of this magnitude can lead to a reduction of some 30% in energy expenditure compared with birds remaining normothermic. Our data suggest that by reducing the T(b)-T(a) gradient, blackcaps accelerate their rate of fuel accumulation at a stopover. When body energy reserves are low blackcaps may achieve this reduction by entering hypothermia. Since hypothermia, as seen in blackcaps, may lead to significant energy savings and facilitate body mass gain, we predict that it is common among small migrating passerines.