Insect cold hardiness : Metabolic , gene , and protein adaptation 1

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.