The mechanism underlying overwintering death in poplar: the cumulative effect of effective freeze–thaw damage

We analyzed the relationships linking overwintering death and frost cracking to temperature and sunlight as well as the effects of low temperatures and freeze–thaw cycles on bud-burst rates, relative electrical conductivity, and phloem and cambial ultrastructures of poplar. Overwintering death rates of poplar were not correlated with negative accumulated temperature or winter minimum temperature. Freeze–thaw cycles caused more bud damage than constant exposure to low temperatures. Resistance to freeze–thaw cycles differed among clones, and the bud-burst rate decreased with increasing exposure to freeze–thaw cycles. Cold-resistant clones had the lowest relative electrical conductivity. Chloroplasts exhibited the fastest and the most obvious reaction to freeze–thaw damage, whereas a single freeze–thaw cycle caused little damage to cambium ultrastructure. Several such cycles resulted in damage to plasma membranes, severe damage to organelles, dehydration of cells and cell death. We conclude that overwintering death of poplar is mainly attributed to the accumulation of effective freeze–thaw damage beyond the limits of freeze–thaw resistance.