Physiological, morphological and allocational plasticity in understory deciduous trees: importance of plant size and light availability

In a 4-year study, we investigated changes in leaf physiology, crown morphology and whole-tree biomass allocation in seedlings and saplings of shade-tolerant sugar maple (Acer saccharum Marsh.) and intermediate shade-tolerant yellow birch (Betula alleghaniensis Britt.) growing in natural understory light (0.5 to 35% of full sunlight) or in understory light reduced by 50% with shade nets to simulate the effect of gap closure. Leaf physiological parameters were mainly influenced by the light gradient, whereas crown morphological and whole-tree allocational parameters were mainly influenced by tree size. No single physiological, morphological or allocational trait was identified that could explain the difference in shade tolerance between the species. Yellow birch had higher growth rates, biomass allocation to branches and leaf physiological plasticity and lower crown morphological plasticity in unmodified understory light than sugar maple. Sugar maple did not display significant physiological plasticity, but showed variation with tree size in both crown morphology and whole-tree biomass allocation. When sugar maple was small, a greater proportion of whole-tree biomass was allocated to roots. However, physiological differences between the species decreased with decreasing light and most morphological and allocational differences tended to disappear with increasing tree size, suggesting that many species differences in shade-tolerance are expressed mainly during the seedling stage. Understory trees of both species survived for 4 years under shade nets, possibly because of higher plasticity when small and the use of stored reserves when taller