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Adaptive thermal plasticity allows organisms to adjust their physiology to cope with fluctuating environments. However, thermal plasticity is rarely studied in response to thermal variability and is often measured in a single life stage. Plasticity in response to thermal variability likely differs from responses to constant temperatures or acute stress. In addition, life stages likely differ in their plasticity and responses in one stage may be affected by the experiences in a previous stage. Increasing the resolution with which we understand thermal plasticity in response to thermal variation across ontogeny is crucial to understanding how organisms cope with the thermal variation in their environment and to estimating the capacity of plasticity to mitigate costs of rapid environmental change. We wanted to know if life stages differ in their capacity for thermal plasticity under temperature fluctuations. We reared Onthophagus taurus dung beetles in either low or high temperature fluctuation treatments and quantified thermal plasticity of metabolism of pupae and adults. We found that adults were thermally plastic and pupae were not. Next, we wanted to know if the plasticity observed in the adult life stage was affected by the thermal conditions during development. We again used low and high temperature fluctuation treatments and reared individuals in one condition through all egg to pupal stages. At eclosion, we switched half of the individuals in each treatment to the opposite fluctuation condition and, later, measured thermal plasticity of metabolism on adults. We found that temperature conditions experienced during the adult stage, but not egg to pupal stages, affects adult thermal plasticity. However, temperature fluctuations during development affect adult body size, suggesting that some aspects of the adult phenotype are decoupled from previous life stages and others are not. Our data demonstrate that life stages mount different responses to temperature variability and uniquely contribute to the adult phenotype. These findings emphasize the need to broadly integrate the life cycle into studies of phenotypic plasticity and physiology; doing so should enhance our ability to predict organismal responses to rapid global change and inform conservation efforts. 

Most studies exploring molecular and physiological responses to temperature have focused on constant temperature treatments. To gain a better understanding of the impacts of fluctuating temperatures, we investigated impacts of increased temperature variation on Phanaeus vindex dung beetles across levels of biological organization. Specifically, we hypothesized that increased temperature variation is energetically demanding. We predicted that thermal sensitivity of metabolic rate and energetic reserves would be reduced with increasing fluctuation. To test this, we examined responses of dung beetles exposed to constant (20°C), low fluctuation (20±5°C), or high fluctuation (20±12°C) temperature treatments using respirometry, energetic reserves, and HPLC-MS-based metabolomics. We found no significant differences in metabolic rates or energetic reserves, suggesting increased fluctuations were not energetically demanding. To understand why there was no effect of increased amplitude on energetics, we assembled and annotated a de novo transcriptome, finding non-overlapping transcriptomic and metabolomic responses of beetles exposed to different fluctuations. We found that 58 metabolites increased in abundance in both fluctuation treatments, but 15 only did so in response to high amplitude fluctuations. We found 120 transcripts were significantly upregulated following acclimation to any fluctuation, but 174 were upregulated only in beetles from the high amplitude fluctuation. Several differentially expressed transcripts were associated with post-translational modifications to histones that support a more open chromatin structure. Our results demonstrate that acclimation to different temperature fluctuations is distinct and may be supported by increasing transcriptional plasticity. Our results indicate for the first time that histone modifications may underlie rapid acclimation to temperature variation.