Ice Nucleation, Propagation, and Deep Supercooling in Woody Plants

Summary The response of woody plants to freezing temperatures is complex. Species vary greatly in their ability to survive freezing temperatures and the resulting dehydrative and mechanical stresses that occur as a result from the presence of ice. Initially, this is presented by the ability to inhibit the formation of ice (ice nucleation) by supercooling. Significant questions exist about the role of internal and external ice nucleating agents in determining the extent to which any particular plant can supercool. Additionally, little is known about how plant structure can affect ice nucleation and propagation. In this review, the ability of high-resolution infrared thermography to reveal significant details about the freezing process is demonstrated. In general, the presence of effective, intrinsic nucleators appear to be common in woody plants. The nucleators appear to be as effective as external ice nucleators and induce stems to freeze at warm, subzero temperatures. Barriers appear to exist, however, that prevent ice propagation into lateral appendages such as buds, or newly extended primary tissues. Deep supercooling represents a unique adaptation of woody plants to avoid freezing injury by dramatically suppressing ice formation in specific tissues. The extent of suppression is limited by the homogeneous nucleation temperature of water (-38°C) and therefore deep supercooling is characteristic of moderately hardy woody plants. In contrast, it has been proposed that the most cold-hardy woody plants have the ability to form glassed solutions. These solutions are very stable as long as the cell remains below the melting temperature of the glass and so allows tissues to become relatively impervious to the stresses associated with extremely low temperatures.