东方蝾螈幼体热耐受性和游泳表现的热驯化响应

特定物种的热驯化能力决定着其是否能成功耐受环境温度的改变,在应对未来气候变暖的趋势中扮演重要角色。为评估有尾类两栖动物的热驯化反应,在3个恒定水温(15、20℃和25℃)中驯化东方蝾螈(Cynops orientalis)幼体4周,测定驯化后幼体在不同测试温度下的运动(游泳)表现、临界低温(CT_(Min))与临界高温(CT_(Max))。结果显示:驯化与测试温度均显著影响蝾螈的游泳速度;驯化温度亦影响蝾螈的CT_(Min)和CT_(Max),但不影响可耐受温度范围(TRR)。驯化与测试温度的交互作用对蝾螈泳速的影响显著,表明驯化温度可改变其游泳表现的热敏感性。经某一温度驯化后蝾螈泳速似乎在相同测试温度下表现最好,该结果可能支持驯化有益假说。CT_(Min)和CT_(Max)随驯化温度的升高而增加,表明:低温驯化可提高动物抗低温能力,而高温驯化提高其抗高温能力。两栖类动物热耐受性与运动表现热驯化反应的种间变异可能与栖息地热环境的差异有关。     

The effect of temperature and thermal acclimation on the sustainable performance of swimming scup

There is a significant reduction in overall maximum power output of muscle at low temperatures due to reduced steady-state (i.e. maximum activation) power-generating capabilities of muscle. However, during cyclical locomotion, a further reduction in power is due to the interplay between non-steady-state contractile properties of muscle (i.e. rates of activation and relaxation) and the stimulation and the length-change pattern muscle undergoes in vivo. In particular, even though the relaxation rate of scup red muscle is slowed greatly at cold temperatures (10°C), warm-acclimated scup swim with the same stimulus duty cycles at cold as they do at warm temperature, not affording slow-relaxing muscle any additional time to relax. Hence, at 10°C, red muscle generates extremely low or negative work in most parts of the body, at all but the slowest swimming speeds.Do scup shorten their stimulation duration and increase muscle relaxation rate during cold acclimation? At 10°C, electromyography (EMG) duty cycles were 18% shorter in cold-acclimated scup than in warm-acclimated scup. But contrary to the expectations, the red muscle did not have a faster relaxation rate, rather, cold-acclimated muscle had an approximately 50% faster activation rate. By driving cold- and warm-acclimated muscle through cold- and warm-acclimated conditions, we found a very large increase in red muscle power during swimming at 10°C. As expected, reducing stimulation duration markedly increased power output. However, the increased rate of activation alone produced an even greater effect. Hence, to fully understand thermal acclimation, it is necessary to examine the whole system under realistic physiological conditions.     

热驯化对鼎突多刺蚁热适应和运动行为的影响

为比较恒温和变温驯化对鼎突多刺蚁（Polyrhachis vicina Roger）热适应和运动行为的影响,将鼎突多翅蚁分别在15℃恒温和13.4～21.6℃变温下进行驯化,定量分析两种驯化温度对鼎突多翅蚁热适应和运动行为的影响.结果表明,恒温和变温驯化对鼎突多刺蚁的最适温度、逃避高温、运动中的停顿频率、疾跑速度、最大持续运动距离均有显著影响（p＜0.01）,而对逃避低温的影响不显著（p=0.343）;变温驯化后鼎突多刺蚁的最适温度仅有一个峰值,这不符合最适性模型的预测;恒温和变温驯化下鼎突多刺蚁的疾跑速度与最大持续运动距离都呈显著的正相关（分别为p=0.017和p＜0.001）,且经过变温驯化的鼎突多刺蚁的运动能力明显强于恒温驯化下个体的运动能力.     

The complex drivers of thermal acclimation and breadth in ectotherms

Thermal acclimation capacity, the degree to which organisms can alter their optimal performance temperature and critical thermal limits with changing temperatures, reflects their ability to respond to temperature variability and thus might be important for coping with global climate change. Here, we combine simulation modelling with analysis of published data on thermal acclimation and breadth (range of temperatures over which organisms perform well) to develop a framework for predicting thermal plasticity across taxa, latitudes, body sizes, traits, habitats and methodological factors. Our synthesis includes > 2000 measures of acclimation capacities from > 500 species of ectotherms spanning fungi, invertebrates, and vertebrates from freshwater, marine and terrestrial habitats. We find that body size, latitude, and methodological factors often interact to shape acclimation responses and that acclimation rate scales negatively with body size, contributing to a general negative association between body size and thermal breadth across species. Additionally, we reveal that acclimation capacity increases with body size, increases with latitude (to mid‐latitudinal zones) and seasonality for smaller but not larger organisms, decreases with thermal safety margin (upper lethal temperature minus maximum environmental temperatures), and is regularly underestimated because of experimental artefacts. We then demonstrate that our framework can predict the contribution of acclimation plasticity to the IUCN threat status of amphibians globally, suggesting that phenotypic plasticity is already buffering some species from climate change.     

The effect of temperature and acclimation period on repeat swimming performance in cutthroat trout

Hatchery cutthroat trout Oncorhynchus clarki clarki were used to examine the effects of 48 h and 3 week temperature acclimation periods on critical swimming speed ( U _crit). The U _crit was determined for fish at acclimation temperatures of 7, 14 and 18° C using two consecutive ramp‐ U _crit tests in mobile Brett‐type swim tunnels. An additional group was tested at the stock's ambient rearing temperature of 10° C. The length of the temperature acclimation period had no significant effect on either the first or the second U _crit( U _crit‐1 and U _crit‐2, respectively) or on the recovery ratio (the quotient of U _crit‐2  U _crit‐1⁻¹). As anticipated, there was a significant positive relationship between U _crit‐1 and temperature ( P  < 0·01) for both acclimation periods, and an increasing, though non‐significant, trend between U _crit‐2 and temperature ( P  = 0·10). Acclimation temperature had no significant effect ( P  = 0·71) on the recovery ratio. These results indicate that a 48 h acclimation to experimental temperatures within the range of −3 to +8° C of the acclimation temperature may be sufficient in studies of swimming performance with this species. This ability to acclimate rapidly is probably adaptive for cutthroat trout and other species that occupy thermally variable environments.     

Acclimation is beneficial at extreme test temperatures in the Danube crested newt, Triturus dobrogicus (Caudata, Salamandridae)

Despite many studies demonstrating the effect of acclimation on behavioural or physiological traits, considerable debate still exists about the evolutionary significance of this phenomenon. One of the unresolved issues is whether acclimation to warmer temperature is beneficial at treatment or at more extreme test temperatures. To answer this question, we assessed the effect of thermal acclimation on preferred body temperatures ( T _ps), maximum swimming and running speed, and critical thermal maximum ( CT _max) in the Danube crested newt ( Triturus dobrogicus ). Adult newts were kept at 15 °C (control) and 25 °C (treatment) for 8 weeks prior to measurements. We measured T _ps in an aquatic thermal gradient over 24 h, maximum speeds in a linear racetrack at six temperatures (5–33 °C), and CT _max in a continuously heated water bath. T _ps were higher in newts kept at 15 °C than in those kept at 25 °C. The maximum swimming speed did not acclimate. The maximum running speed at 30–33 °C was substantially higher in newts kept at 25 °C than in those kept at 15 °C. CT _max increased with the treatment temperature. Hence, we conclude that the acclimation response to warm temperature is beneficial not at treatment but at more extreme temperatures in newts.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 627–636.     

Resistance to temperature extremes in sub-Antarctic weevils: interspecific variation, population differentiation and acclimation

Much of the work on the responses of terrestrial arthropods to high and low temperatures has been done on model organisms such as Drosophila . However, considerable variation in thermotolerance is partitioned at the family level and above, raising questions about the broader applicability of this work to other taxa. Here we investigate resistance to high and low temperatures, following different temperature treatments, in ten species and 31 populations of weevils found on sub-Antarctic Heard Island and Marion Island, which have substantially different climates. In these weevils there is considerable interspecific and among-population variation in critical thermal minimum (CTmin) and critical thermal maximum (CTmax), but most of this variation in critical limits can be ascribed to phenotypic plasticity. We find no relationship between CTmin and CTmax at the species level, and this is true also of populations and of responses to the temperature treatments. In general, plastic (acclimation) changes in CTmin are larger than those in CTmax. Our data therefore provide support for the idea that resistance to heat and to cold are decoupled in terrestrial arthropods. Furthermore, our results suggest that investigations of physiological limits to species borders should incorporate the effects of phenotypic plasticity on physiological capabilities.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78, 401–414.     

Thermal acclimation in Arabidopsis lyrata: genotypic costs and transcriptional changes

Frost and heat events can be challenging for sessile organisms that cannot escape thermal extremes. However, adverse effects of thermal stress on fitness may be reduced by pre‐exposure to cold or heat, a process known as acclimation. To understand the ecological and evolutionary implications of acclimation, we investigated (1) the reduction in performance due to stress pre‐exposure, (2) the magnitude of increased leaf resistance to subsequent stress, (3) the costs of acclimation and (4) the genes differing in expression due to stress pre‐exposure. Plants of Arabidopsis lyrata were raised under three treatments of pre‐exposure: bouts of frost, bouts of heat or constant temperature. Resistance of leaves to subsequent frost and heat stress was then measured by electrolyte leakage. RNA‐seq analysis was performed to examine the genes differentially expressed between stress‐pre‐exposed and control plants. Pre‐exposure to stress during growth decreased plant size and increased leaf resistance to subsequent stress independent of whether pre‐exposure was to frost or heat. But the highest increase in leaf resistance to frost was found after pre‐exposure to frost (as a trend) and in leaf resistance to heat after pre‐exposure to heat. No evidence for costs of acclimation was detected. RNA‐sequencing suggested that acclimation by frost and heat pre‐exposure was caused by distinct mechanisms: modification of the chloroplast membrane and modification of the cell wall and membrane, respectively. Our results suggest that thermal resistance is a labile complex of traits, strongly affected by the previously experienced stress environment, with undetermined costs.     

Growth,stress, and acclimation responses to fluctuating temperatures in field and domesticated populations of Manduca sexta

Diurnal fluctuations in temperature are ubiquitous in terrestrial environments, and insects and other ectotherms have evolved to tolerate or acclimate to such fluctuations. Few studies have examined whether ectotherms acclimate to diurnal temperature fluctuations, or how natural and domesticated populations differ in their responses to diurnal fluctuations. We examine how diurnally fluctuating temperatures during development affect growth, acclimation, and stress responses for two populations of Manduca sexta: a field population that typically experiences wide variation in mean and fluctuations in temperature, and a laboratory population that has been domesticated in nearly constant temperatures for more than 300 generations. Laboratory experiments showed that diurnal fluctuations throughout larval development reduced pupal mass for the laboratory but not the field population. The differing effects of diurnal fluctuations were greatest at higher mean temperature (30°C): Here diurnal fluctuations reduced pupal mass and increased pupal development time for the laboratory population, but had little effect for the field population. We also evaluated how mean and fluctuations in temperature during early larval development affected growth rate during the final larval instar as a function of test temperature. At an intermediate (25°C) mean temperature, both the laboratory and field population showed a positive acclimation response to diurnal fluctuations, in which subsequent growth rate was significantly higher at most test temperatures. In contrast at higher mean temperature (30°C), diurnal fluctuations significantly reduced subsequent growth rate at most test temperatures for the laboratory population, but not for the field population. These results suggest that during domestication in constant temperatures, the laboratory population has lost the capacity to tolerate or acclimate to high and fluctuating temperatures. Population differences in acclimation capacity in response to temperature fluctuations have not been previously demonstrated, but they may be important for understanding the evolution of reaction norms and performance curves.     

Environmental maternal effects on locomotor performance in the common lizard (Lacerta vivipara)

Environmental (i.e. non-genetic) maternal effects have the potential to associate the environmental conditions faced by mothers during gestation or before egg laying with the phenotype of their offspring. For this reason, maternal effects may play a major role in determining offspring phenotype independently of the genotype of the individuals, and can thus be considered a mechanistic basis of phenotypic plasticity. Despite the ecological and evolutionary implications of environmental maternal effects, few studies have experimentally investigated this phenomenon in reptiles. Here we report the results of an experimental laboratory study on the effects of maternal feeding rate and density on offspring locomotor performance in the common lizard (Lacerta vivipara). Lacerta vivipara is a viviparous lizard, and viviparity enhances the probability of a maternal influence on offspring phenotype. We focused on a particular phenotypic trait, maximal sprint running speed, because this trait is thought to be selectively important in squamates. Sprint speed was a repeatable trait, and it varied significantly among families. Maternal feeding rate significantly affected sprint speed, whereas density had no effect on this trait. The effect of maternal feeding rate differed according to the sex of the offspring and their body size, resulting in significant two-way and three-way interactions among these factors. In other words, the maternal feeding rate changed the shape of the allometric relationship between speed and size, but differently for males and females. The complexity of such effects makes it extremely difficult to offer an adaptive interpretation, but emphasizes the role played by the environment in shaping phenotypes among generations.     

Eggs in autumn: responses to declining incubation temperatures by the eggs of montane lizards

Facultative hatching in response to environmental cues may increase the viability of offspring, if the cue that stimulates hatching also predicts the negative consequences of delayed emergence. Declining incubation temperatures might provide such a cue for montane lizards, because eggs that fail to hatch before winter will perish in the nest. I tested this idea by incubating eggs of an alpine scincid lizard ( Bassiana duperreyi ) in the laboratory. For the first half of the incubation period the eggs were kept at nest temperatures typical of those experienced in summer in the field (daily cycle of 18 ± 7.5°C). I then transferred eggs at weekly intervals into cooler regimes (either 15 ± 7.5°C; or with daytime temperatures unchanged but dropping to 0°C overnight). Contrary to prediction, the eggs did not hatch early. However, transfer to lower temperatures caused only a relatively short delay in hatching, because of a virtual temperature-independence of developmental rates late (but not early) in incubation. Decreasing incubation temperatures also modified hatchling running speeds and post-hatching growth rates, even if the thermal decrease occurred only shortly before the usual time of hatching. These processes plausibly affect hatchling fitness in cold-climate reptiles, and might be adaptations to montane habitats. Alternatively, they may prove to be widespread in other (warmer-climate) reptile taxa, in which case no adaptive hypothesis need be proposed. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 71–77.     

Sexual differences in growth strategies of the wolf spider Hygrolycosa rubrofasciata

In invertebrates, the size at maturation is considered to be important for adult fitness. In the wolf spider Hygrolycosa rubrofasciata, however, it is only females that clearly benefit of larger size through augmented egg production, while male mating success is determined by display activity not related to size. Thus, we can expect conflicting growth patterns for the sexes. Additionally, populations differ greatly in adult size: individuals from dry habitats are smaller than those from wet habitats. To study the sexual differences in reaction norms of growth, we reared spiderlings from seven populations at two food levels under controlled laboratory conditions and compared size at sexual maturity. The shapes of reaction norms for adult size differed between the sexes. In females, the reaction norms were parallel, but individuals from dry habitats tended to grow larger at the given food levels. In males, there was a significant interaction between food level and population without any consistent differences between populations. Maturation time was a plastic character in both sexes with no genetic differences among populations. However, females on low food level matured later and significantly smaller in size than those on high food level. Males also matured later on low food level, but they were nearly of the same size as males that received more food. Female growth patterns reflected the strong selection for large size at maturity. However, the patterns for males were highly variable, which could be explained by the weak overall selection on male size, which means that any environmental factors can affect male growing patterns. This revised version was published online in November 2006 with corrections to the Cover Date.     

Investigating thermal acclimation effects before and after a cold shock in Drosophila melanogaster using behavioural assays

Acclimation to environmental change can impose both costs and benefits to organisms. In this study we explored to what extent locomotor behaviour of Drosophila melanogaster is influenced by developmental temperature and adult temperature in both the laboratory and the field. Following development at 15, 25, or 31 °C, adult flies were tested for locomotor activity at all developmental temperatures in the laboratory before and after exposure to a cold shock and in the field for their ability to locate resources after a cold shock. Both test (15, 25, and 31 °C) and developmental temperatures strongly affected locomoter activity, with flies developed at 25 °C having the highest activity at all three test temperatures before the cold shock. After the cold shock flies developed at 15 °C had higher activity compared with flies developed at 25 and 31 °C when tested at 15 and 25 °C, and flies developed at 25 °C had the highest activity when tested at 31 °C. Furthermore, flies developed at 31 °C showed longer recovery times following the cold shock at test temperatures of 15 and 25 °C. However, flies acclimated at 15 °C during development did not recover faster at 15 and 25 °C compared with flies developed at 25 °C. There were no significant correlations between recovery time and locomotor activity at any of the test temperatures. Flies developed at 15 °C and exposed to a cold shock before release in the field were much more successful in locating a resource at low field temperatures compared with flies developed at 25 and 31 °C. Our results provide support for both the beneficial acclimation hypothesis and the optimal developmental temperature hypothesis, but the results are highly context dependent and change with the temperature experienced by the individual during its lifetime.     

外来捕食者克氏原螯虾对黑斑侧褶蛙不同发育阶段幼体表型与运动能力的影响

外来物种入侵是造成本土无尾两栖类种群下降和灭绝的最严重威胁之一。在中国南部丽水地区,克氏原螯虾(Procambarus clarkiion)是一种捕食本土无尾类蝌蚪的入侵物种。在实验室条件下,通过对黑斑侧褶蛙(Pelophylax nigromaculatus)受精卵和蝌蚪先后进行克氏原螯虾捕食风险的不同处理,检测孵出蝌蚪的形态特征、中期蝌蚪的形态特征、游泳表现及活动水平、变态时间与大小以及幼蛙跳跃表现,评估克氏原螯虾的捕食风险对黑斑侧褶蛙不同生长阶段幼体生长发育、运动能力及行为的综合影响。单因素方差分析表明,捕食者长期存在条件下黑斑侧褶蛙孵出蝌蚪的体长、尾长和尾肌宽均显著短于无捕食者的空白对照组和捕食者仅存在2 d的实验组,后两者处理下的蝌蚪体长差异显著,但尾长和尾肌宽差异不显著。长期捕食者存在条件下黑斑侧褶蛙蝌蚪的游泳总路程和平均游速均显著小于无捕食者和捕食者存在2 d的实验组,而蝌蚪体长、尾长、尾鳍高和尾肌高的组间差异均不显著。有捕食者存在下黑斑侧褶幼蛙的体长和体重均显著大于无捕食者和捕食者存在2 d的实验组的情况,而变态时间和幼蛙跳跃距离组间差异不显著。两因素方差分析表明,测试...     

Phenotypic plasticity of abdominal pigmentation in Drosophila kikkawai: multiple interactions between a major gene, sex, abdomen segment and growth temperature

Drosophila kikkawai is known to be polymorphic for a single autosomal locus controlling abdomen pigmentation in females. Two strains homozygous at this locus (Abdomen pigmentation, Abp) were established from a polymorphic Indian population: one was homozygous (DD) for the dark allele, the other (LL) for the light allele. A Mendelian analysis of crosses at 25°C confirmed the occurrence of a major locus, with dominance of the D allele. Phenotypic variation of pigmentation according to growth temperature was then analyzed in DD and LL male and female flies, and in reciprocal F1. A slight difference was found between reciprocal F1 females from a dark mother were darker but not at all temperatures. In females, the D allele exhibited an antero‐posterior gradient of increasing expression from segment 27, with dominance over L and an increased expression at low temperatures. In males, abdomen pigmentation was uniformly light in segments 25, the D allele being repressed by the sex genotype. In segment 6, the D allele was expressed but only at low temperatures, and was either recessive to L or codominant. Phenotypic plasticity that is, amount of change induced by different growth temperatures, was variable according to genotype and segment. It always corresponded to a darkening of the fly at lower temperatures, but was generally much less than in D. melanogaster. In D. kikkawai, climatic adaptation might occur more by changing the frequency of the D allele than by phenotypic plasticity. This revised version was published online in July 2006 with corrections to the Cover Date.     

Predictability rather than amplitude of temperature fluctuations determines stress resistance in a natural population of Drosophila simulans

The adaptability of organisms to novel environmental conditions depends on the amount of genetic variance present in the population as well as on the ability of individuals to adjust their phenotype through phenotypic plasticity. Here, we investigated the phenotypic plasticity induced by a single generation's exposure to three different temperature regimes with respect to several life‐history and stress‐resistance traits in a natural population of Drosophila simulans. We studied a constant as well as a predictably and an unpredictably fluctuating temperature regime. We found high levels of phenotypic plasticity among all temperature regimes, suggesting a strong influence of both temperature fluctuations and their predictability. Increased heat tolerance was observed for flies developed in both types of fluctuating thermal environments compared with flies developed in a constant environment. We suggest that this was due to beneficial hardening when developing in either fluctuating temperature environment. To our surprise, flies that developed in constant and predictably changing environments were similar to each other in most traits when compared to flies from the unpredictably fluctuating environment. The unpredictably changing thermal environment imposed the most stressful condition, resulting in the lowest performance for stress‐related traits, even though the absolute temperature changes never exceeded that of the predictably fluctuating environment. The overall decreased stress resistance of flies in the unpredictably fluctuating environment may be the consequence of maladaptive phenotypic plasticity in this setting, indicating that the adaptive value of plasticity depends on the predictability of the environment.     

中华倒刺鲃不同生理性能对降温的响应速率

为了考查长期高温驯化的中华倒刺鲃(Spinibarbus sinensis)幼鱼快速降温后不同生理功能对低温环境的响应速率,将实验鱼分为2组分别在15℃和25℃条件下长期驯养(8周),随后25℃组快速降温至15℃(降温组),同时15℃组(恒低温对照组)温度保持不变;在降温后的第1、2、4和8周分别测定和比较降温组和恒温对照组实验鱼的温度耐受能力、游泳能力、自发活动水平、摄食代谢和生长性能等相关生理指标。结果显示:降温后,低温耐受能力(CT_(min))1周时就达到稳定与对照组不再有差异,而高温耐受能力(CT_(max))2周时才达到稳定;与对照组相比,降温处理导致实验鱼自发活动水平(运动时间百分比,PTM和运动总距离,TDM)显著上升,且此差异持续整个驯化周期(P0.05);降温组的临界游泳速度(Critical swimming speed U_(crit))在降温的1—2周均显著小于对照组(P0.05),直到4周才达到稳定;降温组实验鱼特殊动力作用(Specific dynamic action SDA)持续时间和摄食后代谢峰值(Peak metabolic rate PMR)与对照组在降温1—4周均没有显著差异;尽管降温组有更高的摄食率,其食物转化效率和增重率均显著低于对照组(P0.05)。研究表明:中华倒刺鲃在水体温度下降后,不同生理功能的稳定速率存在差异,这可能与不同生理功能的生态关联及内在机制的不同有关;降温处理导致机体生长受阻,其主要原因可能是由于应对环境温度变动的逆境胁迫、生理功能调整以及自发活动增加所导致的能量消耗上升所致。