The Effect of Temperature Variability on Biological Responses of Ectothermic Animals-A Meta-Analysis

Climate change is altering temperature means and variation, and both need to be considered in predictions underpinning conservation. However, there is no consensus in the literature regarding the effects of temperature fluctuations on biological functions. Fluctuations may affect biological responses because of inequalities from non-linear responses, endocrine regulation or exposure to damaging temperatures. Here we establish the current state of knowledge of how temperature fluctuations impact biological responses within individuals and populations compared to constant temperatures with the same mean. We conducted a meta-analysis of 143 studies on ectothermic animals (1492 effect sizes, 118 species). In this study, 89% of effect sizes were derived from diel cycles, but there were no significant differences between diel cycles and shorter (<8 h) or longer (>48 h) cycles in their effect on biological responses. We show that temperature fluctuations have little effect overall on trait mean and variance. Nonetheless, temperature fluctuations can be stressful: fluctuations increased 'gene expression' in aquatic animals, which was driven mainly by increased hsp70. Fluctuating temperatures also decreased longevity, and increased amplitudes had negative effects on population responses in aquatic organisms. We conclude that mean temperatures and extreme events such as heat waves are important to consider, but regular (particularly diel) temperature fluctuations are less so.