Allocation of resources within mountain birch canopy after simulated winter browsing

As a response to browsing, birches are known to produce fewer but larger, more nutritious leaves, with enhanced palatability for herbivores. We simulated winter browsing in ramets of mountain birch (Betula puhescens ssp. czerepanovii) to find out whether it decreases subsequent foliage biomass and alters the number and type of shoots. After removal of a considerable proportion of buds (up to 35%) in late winter, the birches were able to compensate for the lost leaf biomass in the following summer; there were no differences in total leaf biomass between winter-clipped and control ramets. This indicates that foliage growth was limited by the total amount of stored resources, not by the number of buds. Depending on the position of the buds removed, different mechanisms were responsible for the compensation. After removal of apical buds, the number of leaves decreased significantly but leaves were larger than in control ramets. Removal of the same mass of basal buds - containing similar amount of carbohydrates and proteins as in the treatment removing apical buds activated dormant buds, especially in apical locations, so that leaf number was similar as in the controls; consequently, size of individual leaves increased only slightly. Thus, while the total leaf biomass in a tree seems to be limited by resources from source organs, the distribution of resources among different canopy sections is controlled by their relative sink strengths. In terms of leaf biomass, epical Darts are able to compensate for bud loss by increasing shoot number, basal parts only by increasing leaf size.