Insect cold hardiness: metabolic, gene, and protein adaptation1This review is part of a virtual symposium on recent advances in understanding a variety of complex regulatory processes in insect physiology and endocrinology, including development, metabolism, cold hardiness, food intake and digestion, and diuresis, through the use of omics technologies in the postgenomic era.

Winter survival for thousands of species of insects relies on adaptive strategies for cold hardiness. Two basic mechanisms are widely used (freeze avoidance by deep supercooling and freeze tolerance where insects endure ice formation in extracellular fluid spaces), whereas additional strategies (cryoprotective dehydration, vitrification) are also used by some polar species in extreme environments. This review assesses recent research on the biochemical adaptations that support insect cold hardiness. We examine new information about the regulation of cryoprotectant biosynthesis, mechanisms of metabolic rate depression, role of aquaporins in water and glycerol movement, and cell preservation strategies (chaperones, antioxidant defenses and metal binding proteins, mitochondrial suppression) for survival over the winter. We also review the new information coming from the use of genomic and proteomic screening methods that are greatly widening the scope for discovery of genes and proteins that support winter survival.

A rapid cold-hardening response protecting against cold shock injury in Drosophila melanogaster

In studies of insect cold-hardiness, the supercooling point (SCP) is defined as the temperature at which spontaneous nucleation of body fluids occurs. Despite having an SCP of −20 degrees C, adults of Drosophila melanogaster did not survive exposure to −5 degrees C, which suggests that cold shock causes lethal injury that is not associated with freezing. If, however, flies were chilled at 5 degrees C, for as little as 30 min, approximately 50% of the flies survived exposure to −5 degrees C for 2h. This capacity to cold-harden rapidly was greatest in 3- and 5-day-old adults. The rapid cold-hardening response was also observed in larvae and pupae: no larvae survived 2 h of exposure to −5 degrees C, whereas 63% pupariated if chilled at 5 degrees C before subzero exposure. Similarly, although exposure of pupae to −8 degrees C was lethal, if pre-chilled at 5 degrees C 22% eclosed. This extremely rapid cold-hardening response may function to allow insects to enhance cold-tolerance in response to diurnal or unexpected seasonal decreases in environmental temperature.