Biological ice nucleation and ice distribution in cold-hardy ectothermic animals

For many ectotherms, overwintering survival depends on the avoidance or regulation of ice nucleation and growth within their body fluids. Freeze avoidance via supercooling plays an important role in the cold hardiness of many small species, particularly terrestrial arthropods, that do not survive the freezing of their body fluids. In contrast, mechanisms that limit supercooling and initiate freezing at relatively high temperatures promote survival of the few invertebrates and vertebrates that tolerate freezing. These mechanisms include inoculative freezing, which results from contact with ice in the environment, and various ice nucleating proteins, microbes, and crystalloid compounds. In freeze-tolerant ectotherms, cold hardiness is influenced by complex, seasonally changing interactions among physiological factors, ice nucleators, and the physical microenvironment. Extraorgan sequestration of ice is a major adaptation of freeze tolerance. For most freeze-tolerant species, ice growth is primarily restricted to extracellular compartments; however, intracellular freezing also occurs in some species.