How phenology influences physiology in deciduous forest spring ephemerals.

Spring ephemerals of deciduous forest are adapted to take advantage of the high-light period available in early spring. They appear shortly after snow melt and complete their aboveground growth, including fruit production, within 2 months. After they produce new buds, they senesce and enter dormancy. Dormancy is not very deep in spring ephemerals and during summer differentiation occurs in the bud of the apparently resting organ. Low soil temperatures release dormancy, and the shoots and roots then grow slowly over autumn and winter. The goal of this paper is to show how this characteristic phenology influences many aspects of spring ephemerals' physiology, and the influences these different physiological parameters have on each other. Spring ephemerals have high photosynthetic rates that allow them to rapidly accumulate carbohydrates and complete their aboveground growth in a few weeks. To sustain high photosynthetic rates in early spring, the plants must be able to absorb water efficiently at low soil temperatures and to allocate large amounts of nutrients to the shoot to compensate for lower enzymatic activity at low temperatures. Nutrients are mainly absorbed in spring, although the root system is established in autumn. This means that a large amount of both carbohydrates and nutrients is translocated from the perennial organ to the developing shoot starting in autumn through early spring. Spring ephemerals have low nutrient absorption rates, but high resorption efficiency during leaf senescence. Nevertheless, their high nutrient needs restrict them to rich forest soils. The annual growth rate of spring ephemerals is very slow and this is more likely related to the inherent slow growth rate of the perennial organ than to their short leaf life. As soon as carbohydrate reserves are replenished in spring, sink limitation apparently builds up and induces leaf senescence. A better understanding of the factors controlling the growth rate of spring ephemerals is needed before we can predict these plants' response to climatic changes.