Heterothermic responses in the speckled mousebird (Colius striatus)

We investigated normothermic thermoregulation and heterothermic responses to restricted food in the speckled mousebird Colius striatus, in the context of the widely accepted distinction between normothermia, rest-phase hypothermia, and torpor. Normothermic thermoregulation differed from typical endothermic patterns in that rest-phase body temperature (Tb) was not maintained with respect to a constant setpoint. Instead, Tb decreased during the course of the rest-phase, with the highest cooling rates observed at moderate ambient temperatures (Ta). Restricted food was associated with significant reductions in rest-phase Tb and metabolic rate. The lowest Tb recorded in a bird which was able to arouse spontaneously, was 18.2 degrees C. However, we were unable to clearly discern between normothermic, hypothermic and torpor Tb ranges. Furthermore, heterothermic responses did not accord with the patterns typically observed in birds and mammals. Metabolic suppression normally associated with entry into torpor and the defence of a torpor Tb setpoint was largely absent. The mousebirds significantly reduced their energy expenditure when heterothermic at moderate TaS only. We suggest that the observed patterns of thermoregulation in C. striatus, as well those previously reported in Colius colius, are associated with plesiomorphic clustering behaviour in the Coliiformes, and the tandem evolution of behavioural and metabolic thermoregulation.     

Is ‘nocturnal hypothermia’ a valid physiological concept in small birds?: a study on Bronze Mannikins Spermestes cucullatus

The thermoregulatory capacity and metabolic responses to light–dark cycles under various mild food-deprivation treatments were measured in Bronze Mannikins Spermestes cucullatus (10–11 g). We measured the response of minimum oxygen consumption to ambient temperature in order to determine the basal metabolic rate (BMR), thermal conductance and limits of thermoneutrality of the Mannikins. In addition, we measured oxygen consumption in response to light–dark cycles and three mild food-deprivation treatments. Bronze Mannikins have a low BMR (1.67 mlO₂/g/h) that is c. 50–60% of that predicted from phylogenetically independent allometric curves for all birds. A low BMR resulted in amplitudes of metabolism between the active and rest phases that were double those predicted allometrically from body mass. The reduced nocturnal metabolic rate did not represent torpor. Typically, Mannikins would need to reduce their metabolic rate during the rest phase to c. 17% of BMR to attain the average torpor metabolic rate of other birds. The data are, however, consistent with those of other group-living Afrotropical birds that benefit energetically from group huddling in environments in which moderate seasonality is accompanied by unpredictable climates – and thus unpredictable energy inputs in time and space. When food-deprived and placed under moderate cold stress (20 °C), Mannikins decreased their rest-phase metabolic rates to the same magnitude as several small Holarctic birds. We suggest that, in the context of the progress made to quantify and define proximate heterothermic responses in endotherms, such as torpor and hibernation, the term nocturnal hypothermia often applied to moderate nocturnal reductions in metabolic rate is vague, misleading and inappropriate.     

Thermoregulation and the energetic significance of clustering behavior in the white-backed mousebird (Colius colius)

Thermoregulation and the energetic significance of clustering behavior were assessed in the white-backed mousebird Colius colius. Basal metabolic rate was 40% below the predicted allometric values. Rest-phase body temperature (T(b)) was highly labile and as low as 26 degrees C. Rest-phase T(b) was not regulated with respect to a constant set point temperature, as occurs typically in endotherms. Rather, we observed periods of linear decreases in rest-phase T(b) at a rate dependent on ambient temperature (T(a)) and the number of individuals in a cluster. The apparent inability of individual mousebirds to maintain rest-phase homeothermy suggests that clustering behavior is obligatory in the defense of a rest-phase set point T(b). The low rest-phase body temperatures exhibited by single C. colius hence appear to represent a normothermic state rather than typical avian facultative hypothermia. The birds were able to make significant energy savings by means of clustering behavior. These energy savings were dependent on T(a) and the number of birds in the cluster. At a T(a) of 15 degrees C, the mean energy expenditure of each bird in a cluster of six was 50% of that of a single bird. The metabolic traits of C. colius are likely be adaptive in the arid habitats that this species inhabits.     

Thermoregulation in African Green Pigeons (Treron calvus) and a re-analysis of insular effects on basal metabolic rate and heterothermy in columbid birds

Columbid birds represent a useful model taxon for examining adaptation in metabolic and thermal traits, including the effects of insularity. To test predictions concerning the role of insularity and low predation risk as factors selecting for the use of torpor, and the evolution of low basal metabolic rate in island species, we examined thermoregulation under laboratory and semi-natural conditions in a mainland species, the African Green Pigeon (Treron calvus). Under laboratory conditions, rest-phase body temperature (T _b) was significantly and positively correlated with air temperature (T _a) between 0 and 35 °C, and the relationship between resting metabolic rate (RMR) and T _a differed from typical endothermic patterns. The minimum RMR, which we interpret as basal metabolic rate (BMR), was 0.825 ± 0.090 W. Green pigeons responded to food restriction by significantly decreasing rest-phase T _b, but the reductions were small (at most ~5 °C below normothermic values), with a minimum T _b of 33.1 °C recorded in a food-deprived bird. We found no evidence of the large reductions in T _b and metabolic rate and the lethargic state characteristic of torpor. The absence of torpor in T. calvus lends support to the idea that species restricted to islands that are free of predators are more likely to use torpor than mainland species that face the risk of predation during the rest-phase. We also analysed interspecific variation in columbid BMR in a phylogenetically informed framework and verified the conclusions of an earlier study which found that BMR is significantly lower in island species compared to those that occur on mainlands.     

Hypothermia versus torpor in response to cold stress in the native Australian mouse Pseudomys hermannsburgensis and the introduced house mouse Mus musculus

This study compared torpor as a response to food deprivation and low ambient temperature for the introduced house mouse (Mus musculus) and the Australian endemic sandy inland mouse (Pseudomys hermannsburgensis). The house mouse (mass 13.0+/-0.48 g) had a normothermic body temperature of 34.0+/-0.20 degrees C at ambient temperatures from 5 degrees C to 30 degrees C and a basal metabolic rate at 30 degrees C of 2.29+/-0.07 mL O2 g(-1) h(-1). It used torpor with spontaneous arousal at low ambient temperatures; body temperature during torpor was 20.5+/-3.30 degrees C at 15 degrees C. The sandy inland mouse (mass 11.7+/-0.16 g) had a normothermic T(b) of 33.0+/-0.38 degrees C between T(a) of 5 degrees C to 30 degrees C, and a BMR of 1.45+/-0.26 mL O2 g(-1) h(-1) at 30 degrees C. They became hypothermic at low T(a) (T(b) about 17.3 degrees C at T(a)=15 degrees C), but did not spontaneously arouse. They did, however, survive and become normothermic if returned to room temperature (23 degrees C). We conclude that this is hypothermia, not torpor. Consequently, house mice (Subfamily Murinae) appear to use torpor as an energy conservation strategy whereas sandy inland mice (Subfamily Conilurinae) do not, but can survive hypothermia. This may reflect a general phylogenetic pattern of metabolic reduction in rodents. On the other hand, this may be related to differences in the social structure of house mice (solitary) and sandy inland mice (communal).     

AMP does not induce torpor

Torpor, a state characterized by a well-orchestrated reduction of metabolic rate and body temperature (T(b)), is employed for energetic savings by organisms throughout the animal kingdom. The nucleotide AMP has recently been purported to be a primary regulator of torpor in mice, as circulating AMP is elevated in the fasted state, and administration of AMP causes severe hypothermia. However, we have found that the characteristics and parameters of the hypothermia induced by AMP were dissimilar to those of fasting-induced torpor bouts in mice. Although administration of AMP induced hypothermia (minimum T(b) = 25.2 +/- 0.6 degrees C) similar to the depth of fasting-induced torpor (24.9 +/- 1.5 degrees C), ADP and ATP were equally effective in lowering T(b) (minimum T(b): 24.8 +/- 0.9 degrees C and 24.0 +/- 0.5 degrees C, respectively). The maximum rate of T(b) fall into hypothermia was significantly faster with injection of adenine nucleotides (AMP: -0.24 +/- 0.03; ADP: -0.24 +/- 0.02; ATP: -0.25 +/- 0.03 degrees C/min) than during fasting-induced torpor (-0.13 +/- 0.02 degrees C/min). Heart rate decreased from 755 +/- 15 to 268 +/- 17 beats per minute (bpm) within 1 min of AMP administration, unlike that observed during torpor (from 646 +/- 21 to 294 +/- 19 bpm over 35 min). Finally, the hypothermic effect of AMP was blunted with preadministration of an adenosine receptor blocker, suggesting that AMP action on T(b) is mediated via the adenosine receptor. These data suggest that injection of adenine nucleotides into mice induces a reversible hypothermic state that is unrelated to fasting-induced torpor.     

Seasonal variation in resting metabolic rate and body temperature of streaked tenrecs, Hemicentetes nigriceps and H. Semispinosus (Insectivora: Tenrecidae)

Annual variations in resting metabolic rate (RMR) and body temperature (Tb) were measured in the streaked tenrecs Hemicentetes nigiceps and H. semispinosus. RMR in non-reproducing, non-torpid adults was lower than predicted by the Kleiber (1961) curve, supporting Genoud's (1990) hypothesis on BMR in hibernating mammals. Both species demonstrated a reduction in RMR and Tb during the austral winter and these changes were independent of body mass. H. semispinosus frequently roused from torpor to forage whereas H. nigriceps remained dormant throughout the winter, This contrast in activity patterns may result from altitudinal differences in species distributions. H. nigriceps may be an obligate hibernator whereas H. semispinosus, living at lower, warmer altitudes, may be a facultative hibernator. It is proposed that the ability of H. semispinosus to remain active during favourable climatic conditions allows winter breeding and increases annual reproductive output.     

Ecological factors affect the level and scaling of avian BMR

The basal rate of metabolism (BMR) in 533 species of birds, when examined with ANCOVA, principally correlates with body mass, most of the residual variation correlating with food habits, climate, habitat, a volant or flightless condition, use or not of torpor, and a highland or lowland distribution. Avian BMR also correlates with migratory habits, if climate and a montane distribution is excluded from the analysis, and with an occurrence on small islands if a flightless condition and migration are excluded. Residual variation correlates with membership in avian orders and families principally because these groups are behaviorally and ecologically distinctive. However, the distinction between passerines and other birds remains a significant correlate of avian BMR, even after six ecological factors are included, with other birds having BMRs that averaged 74% of the passerine mean. This combination of factors accounts for 97.7% of the variation in avian BMR. Yet, migratory species that belong to Anseriformes, Charadriiformes, Pelecaniformes, and Procellariiformes and breed in temperate or polar environments have mass-independent basal rates equal to those found in passerines. In contrast, penguins belong to an order of polar, aquatic birds that have basal rates lower than passerines because their flightless condition depresses basal rate. Passerines dominate temperate, terrestrial environments and the four orders of aquatic birds dominate temperate and polar aquatic environments because their high BMRs facilitate reproduction and migration. The low BMRs of tropical passerines may reflect a sedentary lifestyle as much as a life in a tropical climate. Birds have BMRs that are 30-40% greater than mammals because of the commitment of birds to an expensive and expansive form of flight.     

Water and energy economy of an omnivorous bird: population differences in the Rufous-collared Sparrow (Zonotrichia capensis)

We investigated the intraspecific variation in basal metabolic rate (BMR) and total evaporative water loss (TEWL) in the omnivorous passerine Zonotrichia capensis from two populations inhabiting regions with different precipitation regimes and aridity indices. Values of TEWL in birds from the semi-arid region were significantly lower than those found in sparrows from the mesic region. TEWL in birds from the semi-arid site was 74% of the expectation based on body mass for passerines from mesic areas and similar to the allometric expectation for passerines from arid environments. In sparrows from the mesic area, TEWL was higher than predicted by their body mass for passerines from arid environments (133%), but very close (97%) to the expectation for passerines from mesic areas. BMR values were 25% lower in sparrows from the semi-arid region. The lower TEWL and BMR of birds from the semi-arid region may be a physiological adjustment that allows them to cope with fewer resources and/or water. We propose that the lower endogenous heat production in birds from the semi-arid environment may decrease their water requirements.     

Cleavage site of a major yolk protein (MYP) determined by cDNA isolation and amino acid sequencing in sea urchin,Hemicentrotus pulcherrimus

The grey mouse lemur (Microcebus murinus) is a small nocturnal primate exhibiting daily torpor. In constant ambient temperature (22-24 degrees C), body temperature (Tb) and locomotor activity were monitored by telemetry in animals exposed to short (SP: 10 h light/day) or long (LP: 14 light/day) photoperiods. They were first fed ad libitum for 8 days and then subjected to 80% restricted feeding for 8 more days. During ad libitum feeding, locomotor activity was significantly lower in SP-exposed animals than in LP-exposed animals. Whatever the photoperiod, animals entered daily hypothermia within the first hours following the light onset. Depth of daily hypothermia increased irregularly under SP exposure, whereas minimal daily Tb was constantly above 35 degrees C under LP exposure. After the transfer from long photoperiod to short photoperiod corresponding to the induction of seasonal fattening, locomotor activity and depth of controlled daily hypothermia did not change significantly. In contrast, food restriction led to a significant increase in locomotor activity and in frequency of daily torpor (Tb<33 degrees C) and body temperature reached minimum values averaging 25 degrees C. However, SP-exposed animals exhibited lower minimal daily Tb and higher torpor duration than LP exposed animals. Therefore, daily torpor appears as a rapid response to food restriction occurring whatever the photoperiod, although enhanced by short photoperiod.     

Peripheral ghrelin deepens torpor bouts in mice through the arcuate nucleus neuropeptide Y signaling pathway

Many small mammals have the ability to enter torpor, characterized by a controlled drop in body temperature (Tb). We hypothesized that ghrelin would modulate torpor bouts, because torpor is induced by fasting in mice coincident with elevated circulating ghrelin. Female National Institutes of Health (NIH) Swiss mice were implanted with a Tb telemeter and housed at an ambient temperature (Ta) of 18 degrees C. On fasting, all mice entered a bout of torpor (minimum Tb: 23.8+/-2.0 degrees C). Peripheral ghrelin administration (100 microg) during fasting significantly deepened the bout of torpor (Tb minimum: 19.4+/-0.5 degrees C). When the arcuate nucleus (ARC) of the hypothalamus, a ghrelin receptor-rich region of the brain, was chemically ablated with monosodium glutamate (MSG), fasted mice failed to enter torpor (minimum Tb=31.6+/-0.6 degrees C). Furthermore, ghrelin administration had no effect on the Tb minimum of ARC-ablated mice (31.8+/-0.8 degrees C). Two major pathways that regulate food intake reside in the ARC, the anorexigenic alpha-melanocyte stimulating hormone (alpha-MSH) pathway and the orexigenic neuropeptide Y (NPY) signaling pathway. Both Ay mice, which have the alpha-MSH pathway blocked, and Npy-/-mice exhibited shallow, aborted torpor bouts in response to fasting (Tb minimum: 29.1+/-0.6 degrees C and 29.9+/-1.2 degrees C, respectively). Ghrelin deepened torpor in Ay mice (Tb minimum: 22.8+/-1.3 degrees C), but had no effect in Npy-/-mice (Tb minimum: 29.5+/-0.8 degrees C). Collectively, these data suggest that ghrelin's actions on torpor are mediated via NPY neurons within the ARC.     

Monosodium glutamate-induced arcuate nucleus damage affects both natural torpor and 2DG-induced torpor-like hypothermia in Siberian hamsters

Siberian hamsters (Phodopus sungorus) have the ability to express daily torpor and decrease their body temperature to approximately 15 degrees C, providing a significant savings in energy expenditure. Daily torpor in hamsters is cued by winterlike photoperiods and occurs coincident with the annual nadirs in body fat reserves and chronic leptin concentrations. To better understand the neural mechanisms underlying torpor, Siberian hamster pups were postnatally treated with saline or MSG to ablate arcuate nucleus neurons that likely possess leptin receptors. Body temperature was studied telemetrically in cold-acclimated (10 degrees C) male and female hamsters moved to a winterlike photoperiod (10:14-h light-dark cycle) (experiments 1 and 2) or that remained in a summerlike photoperiod (14:10-h light-dark cycle) (experiment 3). In experiment 1, even though other photoperiodic responses persisted, MSG-induced arcuate nucleus ablations prevented the photoperiod-dependent torpor observed in saline-treated Siberian hamsters. MSG-treated hamsters tended to possess greater fat reserves. To determine whether reductions in body fat would increase frequency of photoperiod-induced torpor after MSG treatment, hamsters underwent 2 wk of food restriction (70% of ad libitum) in experiment 2. Although food restriction did increase the frequency of torpor in both MSG- and saline-treated hamsters, it failed to normalize the proportion of MSG-treated hamsters undergoing photoperiod-dependent torpor. In experiment 3, postnatal MSG treatments reduced the proportion of hamsters entering 2DG-induced torpor-like hypothermia by approximately 50% compared with saline-treated hamsters (38 vs. 72%). In those MSG-treated hamsters that did become hypothermic, their minimum temperature during hypothermia was significantly greater than comparable saline-treated hamsters. We conclude that 1) arcuate nucleus mechanisms mediate photoperiod-induced torpor, 2) food-restriction-induced torpor may also be reduced by MSG treatments, and 3) arcuate nucleus neurons make an important, albeit partial, contribution to 2DG-induced torpor-like hypothermia.     

Torpor in an African caprimulgid, the freckled nightjar Caprimulgus tristigma

Recent data suggest that facultative hypothermic responses such as torpor are more important in the energy balance of birds from tropical and sub-tropical regions than previously thought. We used telemetric measurements of skin temperature (T_skin) for five individuals on 151 bird-nights to investigate the occurrence of torpor during winter in an 81 g African caprimulgid, the freckled nightjar Caprimulgus tristigma . We found that freckled nightjars have the capacity to enter torpor, with a minimum observed T_skin of 12.8°C. During the torpor bouts we observed, complete rewarming typically occurred after sunrise, and coincided with the availability of solar radiation. There was considerable inter-individual variability in the frequency and depth of torpor bouts, with one female nightjar exhibiting particularly frequent and deep torpor. Our results confirm the ability to use torpor by a nocturnal aerial insectivore from the Afrotropics, and reiterate the variability in patterns of torpor that can exist within a population.     

Metabolic consequences of overlapping food restriction and cell‐mediated immune response in a long‐distance migratory shorebird,the little ringed plover Charadrius dubius

Investment in immunity is commonly viewed as an energetically costly activity in birds. Although several studies have focused on the energy cost of mounting an immune response and its concomitant physiological trade‐offs, nothing is known about the metabolic adjustments experienced by immunochallenged birds under resource limitation, or about the basal metabolism cost of mounting cell‐mediated immune (CMI) responses in bird species other than non‐migratory passerines. Here we measured the basal metabolic rate (BMR), inflammatory response, and body mass in ad libitum fed and food‐restricted little ringed plovers Charadrius dubius challenged with phytohemagglutinin (PHA) in order to assess the energy cost, the strength, and the time course of the CMI response in a long‐distance migratory bird in different nutritional states. We found that ad libitum birds injected with PHA significantly increased both mass‐independent BMR and inflammatory response, whereas birds with an induced food restriction‐immune response overlap experienced a mass‐independent BMR downregulation and decreased inflammatory response relative to ad libitum birds. We suggest that both the BMR downregulation and the diminished inflammatory response observed in birds facing such an overlap could be energy‐saving mechanisms to maintain the body mass above a critical level and maximize fitness.     

Reproductive activity influences thermoregulation and torpor in pouched mice, Saccostomus campestris

The Afrotropical pouched mouse Saccostomus campestris displays sexual disparity in the use of daily torpor; males reluctantly enter torpor. We tested the hypothesis that males may compensate for a limited heterothermic capacity with lower basal and resting metabolic rates relative to females. We also investigated the association between gonadal activity (testosterone) and the propensity for daily torpor. Body temperature and oxygen consumption were measured at various ambient temperatures and were compared between sexes under ad libitum and restricted-diet treatments. Whereas no significant sex differences were observed in body temperature and oxygen consumption under ad libitum treatment, there were pronounced differences in heterothermic responses under food restriction. Females employed torpor more frequently and also had lower minimum torpor body temperatures (ca. 25 degrees C) than males (ca. 29 degrees C). Testosterone inhibited torpor in males, whereas the majority of saline-treated animals employed torpor under both ad libitum and restricted-diet treatments. This study demonstrated that the limited capacity of male S. campestris to enter torpor is a consequence of reproductive activity and that opportunistic breeding and the absence of seasonal testes regression compromises the capacity to conserve energy through daily torpor.     

Basal metabolic rate and risk‐taking behaviour in birds

Basal metabolic rate (BMR) constitutes the minimal metabolic rate in the zone of thermo‐neutrality, where heat production is not elevated for temperature regulation. BMR thus constitutes the minimum metabolic rate that is required for maintenance. Interspecific variation in BMR in birds is correlated with food habits, climate, habitat, flight activity, torpor, altitude, and migration, although the selective forces involved in the evolution of these presumed adaptations are not always obvious. I suggest that BMR constitutes the minimum level required for maintenance, and that variation in this minimum level reflects the fitness costs and benefits in terms of ability to respond to selective agents like predators, implying that an elevated level of BMR is a cost of wariness towards predators. This hypothesis predicts a positive relationship between BMR and measures of risk taking such as flight initiation distance (FID) of individuals approached by a potential predator. Consistent with this suggestion, I show in a comparative analysis of 76 bird species that species with higher BMR for their body mass have longer FID when approached by a potential predator. This effect was independent of potentially confounding variables and similarity among species due to common phylogenetic descent. These results imply that BMR is positively related to risk‐taking behaviour, and that predation constitutes a neglected factor in the evolution of BMR.     

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.     

Fasting metabolism and thermoregulatory competence of the star-nosed mole, Condylura cristata (Talpidae: Condylurinae).

Metabolic and body temperature (Tb) responses of star-nosed moles (Condylura cristata) exposed to air temperatures ranging from 0 to 33 degrees C were investigated. The thermoneutral zone of this semi-aquatic mole extended from 24.5 to 33 degrees C, over which its basal rate of metabolism averaged 2.25 ml O2 g-1 h-1 (45.16 J g-1 h-1). This rate of metabolism is higher than predicted for terrestrial forms, and substantially higher than for other moles examined to date. Minimum thermal conductance was nearly identical to that predicted for similar-sized eutherians and may represent a compromise between the need to dissipate heat while digging and foraging in subterranean burrows, and the need to conserve heat and avoid hypothermia during exposure to cold. C. cristata precisely regulated Tb (mean +/- SE = 37.7 +/- 0.05 degrees C) over the entire range of test temperatures. Over three separate 24-h periods, Tb of a radio-implanted mole varied from 36.6 to 38.8 degrees C, and generally tracked level of activity. No obvious circadian variation in Tb and activity was apparent, although cyclic 2-4 h intervals of activity punctuated by periods of inactivity lasting 3-5 h were routinely observed. We suggest that the elevated basal metabolic rate and relatively high Tb of star-nosed moles may reflect the semi-aquatic habits of this unique talpid.     

Bird species numbers in an archipelago of reeds at Lake Velence, Hungary

1 Bird species numbers were studied on 109 reed islands at Lake Velence, Hungary, in the 1993 and 1994 breeding seasons. The aim was to describe and account for the abundance and distribution patterns of the bird species. 2 It was expected that an exponential model would fit the calculated species–area curves. However, for the 1993 data, both the power function (LogS ~ LogArea) and the exponential (S ~ LogArea) models did so, while the power function, exponential and linear (S ~ A) models fitted the curves for the 1994 data. 3 The results showed that the pattern was not random: a collection of small islands held more species than a few large islands with the same total area. 4 The relative species richness of small islands is a result of the preference of most common passerine bird species for the edges of reed islands. Most individuals were found in the first 5 m of the reedbed, and no edge avoidance was detected on a local spatial scale. Large, rarer species (e.g. Great White Egret), however, were found to be dependent on large reed islands. 5 Comparison of results with two other studies on bird communities of reed islands revealed that the type of landscape matrix (e.g. deep water, shallow water or agricultural lands) among reed patches significantly influences bird communities. Deep water was dominated by grebes and coot, shallow water by reed‐nesting passerines, and farmed areas by reed‐ and bush‐nesting passerines.     

Translational initiation is uncoupled from elongation at 18 degrees C during mammalian hibernation

Cellular and organismal homeostasis must be maintained across a body temperature (Tb) range of 0 to 37 degrees C during mammalian hibernation. Hibernators depress biosynthetic activities including protein synthesis, concordant with limited energy availability and temperature effects on reaction rates. We used polysome analysis to show that initiation of protein synthesis ceases during entrance into torpor in golden-mantled ground squirrels (Spermophilus lateralis) when Tb reaches 18 degrees C. Elongation of preinitiated polypeptides continues slowly throughout the torpor bout. As Tb begins to rise, initiation resumes even at temperatures below 18 degrees C, although the euthermic polysome pattern is not reestablished. At precisely 18 degrees C, there is a large increase in initiation events and a complete restoration of euthermic polysome distribution patterns. These data indicate a role for both passive and active depression of translation during torpor and are consistent with a requirement for new protein biosynthesis during each interbout arousal.