To freeze or not to freeze? An evolutionary perspective on the cold-hardiness strategies of overwintering ectotherms

We address the question of whether freeze-tolerance, freeze-avoidance, or mixed strategy represents the best adaptation for overwintering ectotherms to endure severe winter. To this end, we develop an optimization fitness model that takes into account different physiological parameters such as energetic level, the physiological stress associated with each strategy, and climatic variables. The results show that the freeze-tolerance strategy is strongly dependent on a low sensitivity to the number of freezing days and on a capacity to reduce stress associated with freezing. This strategy is also favored when the initial energetic level is low compared to the freeze-avoidance strategy, which is favored by a high initial energetic level, a low stress associated with the supercooling, and a low sensitivity of this strategy to climatic conditions. From a theoretical point of view, the mixed strategy permits survival in harsher environments but requires the optimization of all parameters involved in both cold-hardiness strategies. However, the mixed strategy shows energetic advantages in variable environments allowing animals to resist the harshest periods. From the model results, it appears that the physiological processes developed by ectotherms to reduce these stresses might be a key to understanding the evolution of the cold-hardiness strategies.