RESUMEN

Many ectotherms species grow faster but attain a smaller body size when reared under warmer conditions, a phenomenon known as the Temperature-Size Rule (TSR). This rule appears to be stronger in aquatic ectotherms than in terrestrial ectotherms. The difference could be related to difficulties for oxygen uptake in water, whereas on land, adaptive responses in body size may relate to seasonal time constraints. To assess the role of seasonal time constraints in temperature size response of terrestrial ectotherms, we reared the small copper Lycaena phlaeas at three temperatures (18 ˚C, 23˚C and 28˚C) and two photoperiods (16L: 8D and 12L: 12D). We examined whether differences in body size across treatments was related to (1) differences in growth and development, (2) differences in breakpoints during growth trajectories, or (3) differences in ommatidia size (as a proxy for cell size). We found a weak inverse relationship between developmental temperature and the body size of adult butterflies; adult size decreased by approximately 1% for every degree warmer. Under warmer temperatures, caterpillars developed more quickly and had higher growth rates but reached a smaller body size. Under a short photoperiod, both growth and development were slower, especially at the two lower temperatures, but the body size resulting from slow growth over a longer developmental period did not vary with photoperiod. Breakpoints in growth trajectories occurred when larvae reached ∼40% of their maximum mass and these breakpoints were strongly correlated with the size of the resulting adults, suggesting that adult size is predetermined at an early stage. Temperature did not appear to cause reductions in body size through reductions in cell size. Butterflies were largely able to buffer their body size by modulating larval growth and development in tandem. They appear to use photoperiod as a cue to gauge the availability of time (with 12L: 12D indicating less time available) while temperature speeds up growth and development and as such governs the amount of time they need to complete a developmental cycle. Temperature and photoperiod thus induce changes in voltinism to fit a discrete number of generations into a growing season.

RESUMEN

The ability of organisms to cope with environmental stressors depends on the duration and intensity of the stressor, as well as the type of stress. For aquatic organisms, oxygen limitation has been implicated in limiting heat tolerance. Here we examine how starvation affects heat tolerance in the amphipod Gammarus fossarum (Koch, 1836) and whether observed changes can be explained from alterations in oxidative metabolism, depletion of energy reserves, upregulation of heat shock proteins or susceptibility to oxygen limitation. Starved amphipods showed impaired survival compared to fed amphipods during prolonged exposure to mild heat. In contrast, under acute, high-intensity heat exposure they actually showed improved survival. We observed a lower demand for oxygen in starved amphipods which could make them less susceptible to oxygen limitation. Such a role for oxygen in limiting heat tolerance was verified as hypoxia impaired the heat tolerance of amphipods, especially starved ones. Fed amphipods likely rely more on anaerobic metabolism to maintain energy status during heat stress, whereas for starved amphipods aerobic metabolism appears to be more important. The depletion of their energy reserves constrains their ability to maintain energy status via anaerobic metabolism. We did not find evidence that alterations in heat tolerance following starvation were related to the upregulation of heat shock proteins. In conclusion, starvation can have opposite effects on heat tolerance, acting via pathways that are operating on different time scales.

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RESUMEN

Temperature has a profound impact on ectotherms. Warming increases the metabolic oxygen demand of ectotherms, which could result in a mismatch between their oxygen demand and their ability to extract and deliver sufficient oxygen to meet demand. This hypothesis has been mainly tested using short-term exposure to intense thermal stress. However, the thermal responses of organisms can be different on longer timescales, where physiological acclimation becomes increasingly important. Such thermal acclimation effects may reduce the vulnerability of ectotherms to warming on the long term. Thus, responses to intense, short-term thermal stress may be different from responses to moderate, prolonged thermal stress. Here, we examine the effect of thermal acclimation on heat tolerance and metabolism in the aquatic ectotherm Gammarus fossarum (Koch, 1836). Amphipods were acclimated to either 11.1 ±â€¯0.1 °C or 19.8 ±â€¯0.1 °C and after thermal acclimation we measured both their metabolism and their survival time at different temperatures. Our results show that metabolism strongly increased with increasing temperatures in the cold-acclimated group, but less so in the warm-acclimated group. Cold-acclimated amphipods were also more sensitive to thermal stress, especially during prolonged exposure. Thus, the differences between both thermal acclimation groups support the idea of oxygen-limited heat tolerance: cold-acclimated amphipods showed increased oxygen consumption and decreased thermal tolerance. However, across individuals, those that sharply increased oxygen consumption with increasing temperature did not differ in heat tolerance from individuals whose metabolism was much less sensitive to temperature. Thus, acclimation to different temperatures appeared to be beneficial, but a role for oxygen limitation could not be demonstrated unambiguously. Beneficial effect of acclimation were much larger during prolonged exposure, with the acclimation response ratio (ARR) ranging from 0.03 to over 0.5 depending on the time scale (minutes to months). Thus, the acclimatory capacity may have been underestimated by short-term experimental studies.

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