Dormancy cycling at the shoot apical meristem: transitioning between self-organization and self-arrest.

To survive winter deciduous perennials of the temperate zones cease growth and acquire a cold-acclimated state. Timing of these events is guided by sensory systems in the leaves that register critical alterations in photoperiod. Growth cessation on its own is not sufficient to develop adequate freezing tolerance, which requires entry of the shoot apical meristem (SAM) into dormancy. To fully appreciate perennial dormancy as a precondition for cold acclimation it is necessary to assess how it is brought about in a timely fashion, what the nature of it is, and how it is released. Short day (SD) exposure results in growth cessation, bud set, dormancy establishment at the SAM, and a moderate to high level of freezing tolerance. Subsequent chilling releases the SAM from dormancy and enhances freezing tolerance further. Recent investigations indicate that dormancy is a state of self-arrest that is brought about by an enzyme-based system which disrupts the intrinsic signal network of the SAM. Release from this state requires a complimentary enzyme-based system that is preformed during SD and mobilized by chilling. These findings are in agreement with the paradigm of dormancy cycling, which defines the seasonal alternations at the SAM as transitions between states of self-organization and self-arrest. The success of this survival strategy is based on the adequate scheduling of a complex array of events. The appreciation is growing that this involves signal cascades that are, mutatis mutandis, also recruited in floral evocation in many annuals, including Arabidopsis. A heuristic model is presented of dormancy cycling at the SAM, which depicts crucial molecular and cellular events that drive the cycle.