Cold Tolerance of Insects and Other Arthropods [and Discussion]
Arthropods, as poikilotherms, adapt to cold environments in a variety of ways that include extension of locomotory activity to low temperatures, enhancement of metabolic rate and maintenance of a positive energy balance whenever possible. The ecological implications for many such animals are extension of the life cycle and a requirement for an individual to overwinter several times. Prolonged sub-zero temperatures increase the risk of tissue freezing, and two main strategies have been evolved, first avoidance of freezing by supercooling, and secondly, tolerance of extracellular ice. In the first strategy, freezing is invariably lethal and extensive supercooling (to — 30 °C and below) occurs through elimination or masking of potential ice nucleators in the body and accumulation of cryoprotective substances such as polyhydric alcohols and sugars. Such species are termed freezing intolerant. The second strategy, freezing tolerance, is uncommon in arthropods and other invertebrates, and usually occurs in a single life stage of a species. Freezing of liquid in the extracellular compartment is promoted by proteinaceous ice nucleators. Freezing is therefore protective, and the lethal temperature is well below the supercooling point in freezing tolerant individuals, whereas in most freezing intolerant species it is close to or at the supercooling point. Proteins also act as antifreezes in insects of both strategies, producing a thermal hysteresis by lowering the freezing point of haemolymph in a non-colligative fashion while not affecting the melting point temperature. Recent studies and developments in arthropod cold tolerance are discussed against this background, and a broader approach than hitherto is advocated, which integrates ecological information with physiological data.