Freezing is a major environmental stress during an annual cycle of overwintering, temperate-zone perennials. The timing and extent of seasonal cold acclimation (development of freezing tolerance in the fall) and deacclimation (loss of acquired freezing tolerance in response to warm temperatures) are of critical importance for winter survival, particularly in view of the climate change, i.e., unpredictable extreme weather occurrences. For example, plants may acclimate inadequately if exposed to a milder fall climate and may be damaged by sudden frosts. Alternatively, they may deacclimate prematurely as a result of unseasonable, midwinter warm spells and be injured by the cold that follows. Efficient cold acclimation ability, high deacclimation resistance, and efficient reacclimation capacity are, therefore, important components of winter survival in overwintering perennials. These components should be evaluated separately for a successful breeding program focused on improving winter-hardiness. Another layer of complexity that should be carefully considered is that endodormant status (shallow versus deep) of the reproductive/vegetative apices can significantly impact these components of winter-hardiness. Winter survival, especially by woody evergreens, requires tolerance of light stress, which can result in photo-oxidative damage at cold temperatures when biochemistry of photosynthesis is somewhat compromised but light harvesting is unaffected. Accumulation of Elips (early light-induced proteins) in overwintering evergreens during winter represents a relatively novel strategy to cope with such light stress, and investigations on the precise cellular mechanism and genetic control of this strategy deserve research in the future. Investigations into the mechanisms for cold acclimation use laboratory-based, artificial acclimation protocols that often do not closely approximate conditions that plants are typically exposed to in nature. To draw meaningful conclusions about the biology of cold acclimation and ultimately improve freeze resistance under field conditions, one should also include in cold acclimation regimens parameters such as exposure to subfreezing temperatures and realistic diurnal temperature fluctuations and light levels to simulate natural conditions. One of the main objectives of this article is to highlight two areas of research that we believe are important in the context of plant cold-hardiness but, so far, have not received much attention. These are: 1) to understand the biology of deacclimation resistance and reacclimation capacity, two important components of freeze-stress resistance (winter-hardiness) in woody perennials; and 2) to investigate the cellular basis for various strategies used by broad-leaved evergreens for photoprotection during winter. Our emphasis, in this context, is on a family of proteins, called Elips. The second objective of this article is to draw attention of the cold-hardiness research community to the importance of using realistic cold acclimation protocols in controlled environments that will approximate natural/field conditions to be better able to draw meaningful conclusions about the biology of cold acclimation and ultimately improve freeze resistance. Results from our work with Rhododendron (deciduous azaleas and broad-leaved evergreens), blueberry, and that of other researchers are discussed to support these objectives.
Champions of winter survival: cold acclimation and molecular regulation of cold hardiness in evergreen conifers
