Phenotypic compromise in the face of conflicting ecological demands: an example in red knots Calidris canutus

Phenotypic flexibility is a phenomenon where physiological functions in animals are reversibly adjusted in response to ecological constraints. Research usually focuses on effects of single constraints, but under natural conditions animals face a multitude of restrictions acting simultaneously, and potentially generating conflicting demands on the phenotype. We investigated the conflicting demands of low temperatures and a low quality diet on the phenotype of a shorebird, the red knot Calidris canutus . We tested the effects of switching diet from a high quality trout food to low quality hard-shelled bivalves in captive birds acclimated to temperatures reflecting natural winter conditions. Feeding on bivalves generated a digestive constraint forcing the birds to increase the height and width of their gizzard by 66% and 71%, respectively, over 30 days. The change in gizzard size was associated with an initial 15% loss of body mass and a reduction in size of the pectoral muscles by 11%. Because pectoral muscle size determines summit metabolic rate (M_sum, an indicator of cold endurance), measured M_sum declined by 9%. Therefore, although the birds were acclimated to cold, gizzard growth led to a loss of cold endurance. We propose that cold-acclimated knots facing a digestive constraint made a phenotypic compromise by giving-up cold hardiness for digestive capacity. Field studies suggest that phenotypic compromises occur in free-living red knots as well and help improve survival.     

Phenotypic flexibility in digestive system structure and function in migratory birds and its ecological significance

Birds during migration must satisfy the high energy and nutrient demands associated with repeated, intensive flight while often experiencing unpredictable variation in food supply and food quality. Solutions to such different challenges may often be physiologically incompatible. For example, increased food intake and gut size are primarily responsible for satisfying the high energy and nutrient demands associated with migration in birds. However, short-term fasting or food restriction during flight may cause partial atrophy of the gut that may limit utilization of ingested food energy and nutrients. We review the evidence available on the effects of long- and short-term changes in food quality and quantity on digestive performance in migratory birds, and the importance of digestive constraints in limiting the tempo of migration in birds. Another important physiological consequence of feeding in birds is the effect of diet on body composition dynamics during migration. Recent evidence suggests that birds utilize and replenish both protein and fat reserves during migration, and diet quality influences the rate of replenishment of both these reserves. We conclude that diet and phenotypic flexibility in both body composition and the digestive system of migratory birds are important in allowing birds to successfully overcome the often-conflicting physiological challenges of migration.     

Test of a digestion optimization model: effect of variable-reward feeding schedules on digestive performance of a migratory bird

Birds on migration often alternate between feeding and nonfeeding periods, in part because food resources may be patchily distributed and in part because birds on migration may adopt a risk-prone foraging strategy characterized by selection of variable rather than constant food rewards. Optimal digestion models predict that increases in intermeal interval like those encountered by some migratory birds should result in longer retention time of digesta and higher digestive efficiency if birds are maximizing their rate of energy intake. We tested these predictions by comparing residence time of digesta and extraction efficiency of lipid for captive yellow-rumped warblers (Dendroica coronata) feeding adlibitum and when we added intervals of time when the birds received no food. We increased the likelihood that the warblers were maximizing their rate of energy intake by increasing light levels during spring to induce hyperphagia (treatment birds (16L:8D light: dark cycle) ate 2.13 ± 0.14 g dry food day⁻¹ (n = 8) while control birds (10L:14D) ate 1.25 ± 0.03 g dry food day⁻¹ (n = 6)). Treatment birds offered food only every other 2–3 h ate 50% more during the 4-h test period than when they were always feeding adlibitum. Despite these differences in food intake, extraction efficiency of glycerol trioleate remained high and constant (93%), and mouth-to-anus total mean retention time (TMRT) did not change (overall mean: 54.8 ± 6.0 min). Residence time of lipid in the stomach increased whereas residence time of lipid in the intestine decreased when birds fed only every other 2–3 h compared to when birds always fed ad libitum. None of the results were consistent with the predictions of the optimal digestion model unless we assume that birds were minimizing their feeding time rather than maximizing their rate of energy gain. Furthermore, the ability of yellow-rumped warblers to maintain high extraction efficiency with no change in TMRT suggests some spare digestive capacity when food intake increases by as much as 50%. Received: 14 June 1997 / Accepted: 20 November 1997     

Optimal foraging of a herbivorous lizard,the green iguana in a seasonal environment

Food selection was studied in free living green iguanas (Iguana iguana) throughout the year in a semiarid environment, Curaçao (Netherlands Antilles). Food intake was determined by direct observations and converted into biomass intake. Comparison between intake and biomass availability of the various food items revealed that the lizards were selective, and that changes in seasonal food availability led to periodic switching of food plants. The extent to which nutrient constraints determine iguana feeding ecology was investigated. Potential constraints were the requirements for water, digestible crude protein, and metabolizable energy. By using a linear programming model that incorporates characteristics of the food (chemical composition, energy content, item size) and requirements and constraints of the green iguanas (nutrient and energy requirements digestive tract capacity, feeding rate) it was possible to identify which factors determine food choice over the year. During the dry period, when the iguanas had no access to drinking water they consumed flowers to increase water intake, though the amount of flowers consumed was too low to cover maintenance requirements for either energy or protein. After the young leaf flush, following the early rains in May, the biomass increased, free surface water was available during showers, and the linear programming solutions indicate that food selection conformed to the protein maximization criterion. Reproduction in green iguanas shows an annual cycle, in which oviposition takes place at the end of the dry season, when intake is below maintenance levels. Females show a 8–10 month gap between acquisition of most of the protein required for egg synthesis and the act of laying. Thus, as in avian and mammalian herbivores, food availability during a period prior to the energy and protein demanding reproductive season of iguanas determines reproductive success.     

The Effect of Digestive Capacity on the Intake Rate of Toxic and Non-Toxic Prey in an Ecological Context

Digestive capacity often limits food intake rate in animals. Many species can flexibly adjust digestive organ mass, enabling them to increase intake rate in times of increased energy requirement and/or scarcity of high-quality prey. However, some prey species are defended by secondary compounds, thereby forcing a toxin limitation on the forager’s intake rate, a constraint that potentially cannot be alleviated by enlarging digestive capacity. Hence, physiological flexibility may have a differential effect on intake of different prey types, and consequently on dietary preferences. We tested this effect in red knots (Calidris canutus canutus), medium-sized migratory shorebirds that feed on hard-shelled, usually mollusc, prey. Because they ingest their prey whole and crush the shell in their gizzard, the intake rate of red knots is generally constrained by digestive capacity. However, one of their main prey, the bivalve Loripes lucinalis, imposes a toxin constraint due to its symbiosis with sulphide-oxidizing bacteria. We manipulated gizzard sizes of red knots through prolonged exposure to hard-shelled or soft foods. We then measured maximum intake rates of toxic Loripes versus a non-toxic bivalve, Dosinia isocardia. We found that intake of Dosinia exponentially increased with gizzard mass, confirming earlier results with non-toxic prey, whereas intake of Loripes was independent of gizzard mass. Using linear programming, we show that this leads to markedly different expected diet preferences in red knots that try to maximize energy intake rate with a small versus a large gizzard. Intra- and inter-individual variation in digestive capacity is found in many animal species. Hence, the here proposed functional link with individual differences in foraging decisions may be general. We emphasize the potential relevance of individual variation in physiology when studying trophic interactions.     

Do northern bobwhite quail modulate intestinal nutrient absorption in response to dietary change? A test of an adaptational hypothesis

We acclimated northern bobwhite quail (Colinus virginianus) to either chow (high carbohydrate/low protein) or crickets (low carbohydrate/high protein) and tested predictions of hypotheses based on the premise of the economical design of animals. The adaptive modulation hypothesis predicts that d-glucose uptake would be higher and l-proline uptake lower in bobwhites acclimated to chow. The spare capacity hypothesis predicts that the capacity to absorb d-glucose actively will exceed the estimated nutrient load from daily food intake. There was no significant dietary effect on intestinal d-glucose (P = 0.8) and l-proline (P = 0.7) uptake rates measured in vitro using the everted sleeve technique. In chow eaters maximal mediated d-glucose uptake summed along the entire length of intestine (53 cm) was far too low (7.2 mmol/d) to explain observed rates of glucose absorption in vivo (>35 mmol/d). Hence, both predictions were falsified. In vitro uptake may not be an appropriate measure of the intestine's absorptive capacity because it does not measure possibly important pathways of passive absorption. There is increasing evidence that substantial passive glucose absorption occurs in some birds. If passive absorption predominates the adaptive modulation hypothesis might not apply.     

Diet quality and food limitation affect the dynamics of body composition and digestive organs in a migratory songbird (Zonotrichia albicollis)

Migrating songbirds interrupt their feeding to fly between stopover sites that may vary appreciably in diet quality. We studied the effects of fasting and food restriction on body composition and digestive organs in a migratory songbird and how these effects interacted with diet quality to influence the rate of recovery of nutrient reserves. Food limitation caused white-throated sparrows to reduce both lean and fat reserves, with about 20% of the decline in lean mass represented by a decline in stomach, small intestine, and liver. During refeeding on diets similar in nutrient composition to either grain or fruit, food-limited grain-fed birds ate 40% more than did control birds, and they regained body mass, with on average 60% of the increase in body mass composed of lean mass including digestive organs. In contrast, food-limited fruit-fed birds did not eat more than did control birds and did not regain body mass, suggesting that a digestive constraint limited their food intake. The interacting effects of food limitation and diet quality on the dynamics of body composition and digestive organs in sparrows suggest that the adequacy of the diet at stopover sites can directly influence the rate of recovery of body reserves in migrating songbirds and hence the pace of their migration.     

温度与光周期对白头鹎体质量、能量收支和消化道形态的影响

温度与光周期是环境季节性变化的最直接表现因子及时间变化指示标志,对动物的形态、生理及行为产生重要的影响.本文以白头鹎为研究对象,探讨了不同温度与光周期对其体质量、能量收支和消化道形态的影响,分析了能量收支与消化道形态特征的关系.将28只白头鹎(12雄16雌)分为4组:暖温长光组(30℃,16 L8 D;3雄4雌)、暖温短光组(30 ℃,8 L16 D;3雄4雌)、低温长光组(10 ℃,16 L8 D;3雄4雌)和低温短光组(10 ℃,8 L16 D;3雄4雌).结果表明: 低温与短光照可促进白头鹎的体质量、摄入能及同化能明显增加,同时温度与光周期的交互作用对白头鹎的摄入能及同化能影响显著.低温条件下,胃、小肠、直肠及总消化道的湿质量及干质量明显增加.残差分析表明,小肠与总消化道的长度及干质量与摄入能和同化能显著相关.表明低温与短光照下白头鹎通过增加体质量、能量摄入和改变消化道形态来应对严酷的环境条件.     

The Shift of Thermoneutral Zone in Striped Hamster Acclimated to Different Temperatures

Temperature affects all biological functions and will therefore modulate ecologically significant interactions between animals and their environment. Here, we examined the effect of ambient temperature (T_a) on the thermal biology and energy budget in striped hamsters acclimated to cold (5°C), warm (21°C) and hot temperatures (31°C). Thermoneutral zone (TNZ) was 22.5–32.5°C, 25–32.5°C and 30–32.5°C in the cold-, warm- and hot-acclimated hamsters, respectively. The cold acclimation decreased the lower critical temperature and made the TNZ wider, and hot exposure elevated the lower critical temperature, resulting in a narrow TNZ. Within the TNZ, cold-acclimated hamsters showed a significantly higher rate of metabolism and thermogenesis than those acclimated to hot temperature. Digestive enzymes activities, including intestinal sucrase, maltase, L-alanine aminopeptidase-N and leucine aminopeptidase were higher in the cold than in the hot. The changes in metabolic rate and thermogenesis at different temperatures were in parallel with cytochrome c oxidase activity and uncoupling protein 1 gene expression of brown adipose tissue. This suggests that the shift of the lower critical temperature of TNZ is possibly associated with the rate of metabolism and thermogenesis, as well as with the digestive capacity of the gastrointestinal tract at different T_a. The upper critical temperature of TNZ may be independent of the changes in T_a. The changes of lower critical temperature of TNZ are an important strategy in adaption to variations of T_a.     

Seasonal intake responses in the nectar-feeding bat Glossophaga soricina

Food intake in nectar-feeding animals is affected by food quality, their energetic demands, and the environmental conditions they face. These animals increase their food intake in response to a decrease in food quality, a behavior named “intake response”. However, their capacity to achieve compensatory feeding, in which they maintain a constant flux of energy, could be constrained by physiological processes. Here we evaluated how both a seasonal change in environmental conditions and physiological constraints affected the food ingestion in the bat Glossophaga soricina. We measured food intake rate during both the wet/warm and dry/cool seasons at sucrose solutions ranging from 146 to 1,022 mmol L⁻¹. We expected that food intake and metabolic demands would be greater during the dry/cool season. Bats ingested ~20% more food in the dry/cool than in the wet/warm season. Regardless of season, bats were unable to achieve a constant flux of energy when facing the different sugar concentrations that we used in our experiments. This suggests that the rate of food intake is physiologically constrained in G. soricina. Using the digestive capacity of bats we modeled their food intake. The analytic model we used predicts that digestive limitations to ingest energy should have an important effect on the ecology of this species.     

Freezing tolerance of citrus, spinach, and petunia leaf tissue : osmotic adjustment and sensitivity to freeze induced cellular dehydration

Seasonal variations in freezing tolerance, water content, water and osmotic potential, and levels of soluble sugars of leaves of field-grown Valencia orange (Citrus sinensis) trees were studied to determine the ability of citrus trees to cold acclimate under natural conditions. Controlled environmental studies of young potted citrus trees, spinach (Spinacia pleracea), and petunia (Petunia hybrids) were carried out to study the water relations during cold acclimation under less variable conditions. During the coolest weeks of the winter, leaf water content and osmotic potential of field-grown trees decreased about 20 to 25%, while soluble sugars increased by 100%. At the same time, freezing tolerance increased from lethal temperature for 50% (LT₅₀) of −2.8 to −3.8°C. In contrast, citrus leaves cold acclimated at a constant 10°C in growth chambers were freezing tolerant to about −6°C. The calculated freezing induced cellular dehydration at the LT₅₀ remained relatively constant for field-grown leaves throughout the year, but increased for leaves of plants cold acclimated at 10°C in a controlled environment. Spinach leaves cold acclimated at 5°C tolerated increased cellular dehydration compared to nonacclimated leaves. Cold acclimated petunia leaves increased in freezing tolerance by decreasing osmotic potential, but had no capacity to change cellular dehydration sensitivity. The result suggest that two cold acclimation mechanisms are involved in both citrus and spinach leaves and only one in petunia leaves. The common mechanism in all three species tested was a minor increase in tolerance (about −1°C) resulting from low temperature induced osmotic adjustment, and the second in citrus and spinach was a noncolligative mechanism that increased the cellular resistance to freeze hydration.     

Digestive response to restricted feeding in migratory yellow-rumped warblers

Smaller guts and slow initial mass gains at stopover sites have led to the idea that digestive physiology limits refueling rates in migrating birds. We tested the digestive-limitation hypothesis in yellow-rumped warblers using food restriction to simulate infrequent feeding during migration, which may cause a reduction in alimentary tract mass. Restricted birds had small intestine, pancreas, and liver masses 18%-22% lower than ad lib.-fed controls. Total activities of sucrase, maltase, aminopeptidase, and amylase were significantly lower in restricted birds, while those of trypsin and chymotrypsin were not. Only aminopeptidase mass-specific activity was significantly lower in restricted birds. Previously restricted birds were able to feed and digest at a high rate immediately following return to ad lib. feeding. Digestive efficiency did not differ between groups. These results suggest that before migration yellow-rumped warblers have some spare digestive capacity to compensate for declines in their digestive organ masses during migration.     

Out of the frying pan into the air—emersion behaviour and evaporative heat loss in an amphibious mangrove fish (Kryptolebias marmoratus)

Amphibious fishes often emerse (leave water) when faced with unfavourable water conditions. How amphibious fishes cope with the risks of rising water temperatures may depend, in part, on the plasticity of behavioural mechanisms such as emersion thresholds. We hypothesized that the emersion threshold is reversibly plastic and thus dependent on recent acclimation history rather than on conditions during early development. Kryptolebias marmoratus were reared for 1 year at 25 or 30°C and acclimated as adults (one week) to either 25 or 30°C before exposure to an acute increase in water temperature. The emersion threshold temperature and acute thermal tolerance were significantly increased in adult fish acclimated to 30°C, but rearing temperature had no significant effect. Using a thermal imaging camera, we also showed that emersed fish in a low humidity aerial environment (30°C) lost significantly more heat (3.3°C min⁻¹) than those in a high humidity environment (1.6°C min⁻¹). In the field, mean relative humidity was 84%. These results provide evidence of behavioural avoidance of high temperatures and the first quantification of evaporative cooling in an amphibious fish. Furthermore, the avoidance response was reversibly plastic, flexibility that may be important for tropical amphibious fishes under increasing pressures from climatic change.     

Optimal foraging and community structure: implications for a guild of generalist grassland herbivores

Summary A particular linear programming model is constructed to predict the diets of each of 14 species of generalist herbivores at the National Bison Range, Montana. The herbivores have body masses ranging over seven orders of magnitude and belonging to two major taxa: insects and mammals. The linear programming model has three feeding constraints: digestive capacity, feeding time and energy requirements. A foraging strategy that maximizes daily energy intake agrees very well with the observed diets. Body size appears to be an underlying determinant of the foraging parameters leading to diet selection. Species that possess digestive capacity and feeding time constraints which approach each other in magnitude have the most generalized diets. The degree that the linear programming models change their diet predictions with a given percent change in parameter values (sensitivity) may reflect the observed ability of the species to vary their diets. In particular, the species which show the most diet variability are those whose diets tend to be balanced between monocots and dicots. The community-ecological parameters of herbivore body-size ranges and species number can possibly be related to foraging behavior.     

Effects of acclimation and acute temperature change on specific dynamic action and gastric processing in the green shore crab, Carcinus maenas

The effects of temperature acclimation and acute temperature change were investigated in postprandial green shore crabs, Carcinus maenas. Oxygen uptake, gut contractions and transit rates and digestive efficiencies were measured for crabs acclimated to either 10 °C or 20 °C and subsequently exposed to treatment temperatures of 5, 15, or 25 °C. Temperature acclimation resulted in a partial metabolic compensation in unfed crabs, with higher oxygen uptake rates measured for the 10 °C acclimated group exposed to acute test temperatures. The Q₁₀ values were higher than normal, probably because the acute temperature change prevented crabs from fully adjusting to the new temperature. Both the acclimation and treatment temperature altered the characteristics of the specific dynamic action (SDA). The duration of the response was longer for 20 °C acclimated crabs and was inversely related to the treatment temperature. The scope (peak oxygen consumption) was also higher for 20 °C acclimated crabs with a trend towards an inverse relationship with treatment temperature. Since the overall SDA (energy expenditure) is a function of both duration and scope, it was also higher for 20 °C acclimated crabs, with the highest value measured at the treatment temperature of 15 °C. The decline in total SDA after acute exposure to 5 and 25 °C suggests that both cold stress and limitations to oxygen supply at the temperature extremes could be affecting the SDA response. The contractions of the pyloric sac of the foregut region function to propel digesta through the gut, and contraction rates increased with increasing treatment temperature. This translated into faster transit rates with increasing treatment temperatures. Although pyloric sac contractions were higher for 20 °C acclimated crabs, temperature acclimation had no effect on transit rates. This suggests that a threshold level in pyloric sac contraction rates needs to be reached before it manifests itself on transit rates. Although there was a correlation between faster transit times and the shorter duration of the SDA response with increasing treatment temperature, transit rates do not make a good proxy for calculating the SDA characteristics. The digestive efficiency showed a trend towards a decreasing efficiency with increasing treatment temperature; the slower transit rates at the lower treatment temperatures allowing for more efficient nutrient absorption. Even though metabolic rates of 10 °C acclimated crabs were higher, there was no effect of acclimation temperature on digestive efficiency. This probably occurred because intracellular enzymes and digestive enzymes are modulated through different control pathways. These results give an insight into the metabolic and digestive physiology of Carcinus maenas as it makes feeding excursions between the subtidal and intertidal zones.     

Dynamic digestive responses to increased energy demands in the leaf-eared mouse (<Emphasis Type="Italic">Phyllotis darwini</Emphasis>)

A major area of interest in comparative physiology has been to understand how animals cope with changing environmental demands in time and space. The digestive system has been identified as one of the more sensitive systems to changes in environmental conditions. However, most research on this topic has evaluated these effects during peak energetic demands, which do not allow for evaluation of the dynamics of the digestive response along a more natural continuous gradient of environmental conditions. We examined phenotypic flexibility in digestive responses of the leaf-eared mouse Phyllotis darwini to increments in total energy demands (via sequential exposure to 26, 12 and 0°C). Additionally, we evaluated the effect of a moderate energy demand (12°C) over three different time periods (7, 17 and 27 days) on digestive traits. Moderate increases in energy demand were associated with changes in the distribution of digesta in the gut, whereas higher increases in energy demand involved increases in the tissue mass of digestive organs. Time-course analysis showed that at 12°C practically all digestive variables reached stable values within 7 days, which is in agreement with empirical data and theoretical deductions from cellular turnover rates. We conclude that although the input of energy and nutrients into the digestive tract is typically periodic, many aspects of digestive physiology are likely to be flexible in response to environmental variability over both short-term (daily) and long-term (seasonal) time scales.     

Thermal performance of the Chagas disease vector,Triatoma infestans,under thermal variability

Vector-borne diseases (VBD) are particularly susceptible to climate change because most of the diseases’ vectors are ectotherms, which themselves are susceptible to thermal changes. The Chagas disease is one neglected tropical disease caused by the protozoan parasite, Trypanosoma cruzi. One of the main vectors of the Chagas disease in South America is Triatoma infestans, a species traditionally considered to be restricted to domestic or peridomestic habitats, but sylvatic foci have also been described along its distribution. The infestation of wild individuals, together with the projections of environmental changes due to global warming, urge the need to understand the relationship between temperature and the vector’s performance. Here, we evaluated the impact of temperature variability on the thermal response of T. infestans. We acclimated individuals to six thermal treatments for five weeks to then estimate their thermal performance curves (TPCs) by measuring the walking speed of the individuals. We found that the TPCs varied with thermal acclimation and body mass. Individuals acclimated to a low and variable ambient temperature (18°C ± 5°C) exhibited lower performances than those individuals acclimated to an optimal temperature (27°C ± 0°C); while those individuals acclimated to a low but constant temperature (18°C ± 0°C) did not differ in their maximal performance from those at an optimal temperature. Additionally, thermal variability (i.e., ± 5°C) at a high temperature (30°C) increased performance. These results evidenced the plastic response of T. infestans to thermal acclimation. This plastic response and the non-linear effect of thermal variability on the performance of T. infestans posit challenges when predicting changes in the vector’s distribution range under climate change.     

Thermal and digestive constraints to foraging behaviour in marine mammals

While foraging models of terrestrial mammals are concerned primarily with optimizing time/energy budgets, models of foraging behaviour in marine mammals have been primarily concerned with physiological constraints. This has historically centred on calculations of aerobic dive limits. However, other physiological limits are key to forming foraging behaviour, including digestive limitations to food intake and thermoregulation. The ability of an animal to consume sufficient prey to meet its energy requirements is partly determined by its ability to acquire prey (limited by available foraging time, diving capabilities and thermoregulatory costs) and process that prey (limited by maximum digestion capacity and the time devoted to digestion). Failure to consume sufficient prey will have feedback effects on foraging, thermoregulation and digestive capacity through several interacting avenues. Energy deficits will be met through catabolism of tissues, principally the hypodermal lipid layer. Depletion of this blubber layer can affect both buoyancy and gait, increasing the costs and decreasing the efficiency of subsequent foraging attempts. Depletion of the insulative blubber layer may also increase thermoregulatory costs, which will decrease the foraging abilities through higher metabolic overheads. Thus, an energy deficit may lead to a downward spiral of increased tissue catabolism to pay for increased energy costs. Conversely, the heat generated through digestion and foraging activity may help to offset thermoregulatory costs. Finally, the circulatory demands of diving, thermoregulation and digestion may be mutually incompatible. This may force animals to alter time budgets to balance these exclusive demands. Analysis of these interacting processes will lead to a greater understanding of the physiological constraints within which the foraging behaviour must operate.     

Coping with Daily Thermal Variability: Behavioural Performance of an Ectotherm Model in a Warming World

Global climate change poses one of the greatest threats to species persistence. Most analyses of the potential biological impacts have focused on changes in mean temperature, but changes in thermal variance will also impact organisms and populations. We assessed the effects of acclimation to daily variance of temperature on dispersal and exploratory behavior in the terrestrial isopod Porcellio laevis in an open field. Acclimation treatments were 24±0, 24±4 and 24±8°C. Because the performance of ectotherms relates nonlinearly to temperature, we predicted that animals acclimated to a higher daily thermal variation should minimize the time exposed in the centre of open field, – i.e. increase the linearity of displacements. Consistent with our prediction, isopods acclimated to a thermally variable environment reduce their exploratory behaviour, hypothetically to minimize their exposure to adverse environmental conditions. This scenario as well as the long latency of animals after releases acclimated to variable environments is consistent with this idea. We suggested that to develop more realistic predictions about the biological impacts of climate change, one must consider the interactions between the mean and variance of environmental temperature on animals'' performance.     

Effect of cold acclimation on bulk tissue electrical impedance: I. Measurements with birdsfoot trefoil at subfreezing temperatures

The resistive and reactive components of electrical impedance were measured for birdsfoot trefoil (Lotus corniculatus L.) stems at freezing temperatures to −8°C. As temperature decreased the specific resistance at frequencies between 49 hertz and 1.11 megahertz of stems from cold acclimated plants increased more rapidly than from nonacclimated plants. This temperature dependence of specific resistance could be characterized by an Arrhenius activation energy; cold acclimated stems had a larger Arrhenius activation energy than nonacclimated stems. The low frequency resistance is believed to characterize the extracellular region of the stems and the high frequency resistance is believed to characterize the intracellular region of the stems. Cold acclimation increased the intracellular but not the extracellular resistance at nonfreezing temperatures. Cold acclimated stems were not injured by freezing to −8°C and thawing, but nonacclimated stems were injured by freezing to temperatures between −2.2 and −5.6°C and thawing. Injury to nonacclimated stems at freezing temperatures below −2.2°C was indicated by a decrease in the ratio of resistance at 49 Hz to that at 1.11 megahertz.