Size-dependent leaf area ratio in plant twigs: implication for leaf size optimization

Although many hypotheses have been proposed to explain variation in leaf size, the mechanism underlying the variation remains not fully understood. To help understand leaf size variation, the cost/benefit of twig size was analysed, since, according to Corner's rule, twig size is positively correlated with the size of appendages the twig bears. An extensive survey of twig functional traits, including twig (current-year shoots including one stem and few leaves) and leaf size (individual leaf area and mass), was conducted for 234 species from four broadleaved forests. The scaling relationship between twig mass and leaf area was determined using standardized major axis regression and phylogenetic independent comparative analyses. Leaf area was found to scale positively and allometrically with both stem and twig mass (stem mass plus leaf mass) with slopes significantly smaller than 1 center dot 0, independent of life form and habitat type. Thus, the leaf area ratio (the ratio of total leaf area to stem or twig mass) decreases with increasing twig size. Moreover, the leaf area ratio correlated negatively with individual leaf mass. The results of phylogenetic independent comparativeanalyses were consistent with the correlations. Based on the above results, a simple model for twig size optimization was constructed, from which it is postulated that large leaf size-twig size may be favoured when leaf photosynthetic capacity is high and/or when leaf life span and/or stem longevity are long. The model's predictions are consistent with leaf size variation among habitats, in which leaf size tends to be small in poor habitats with a low primary productivity. The model also explains large variations in leaf size within habitats for which leaf longevity and stem longevity serve as important determinants. The diminishing returns in the scaling of total leaf area with twig size can be explained in terms of a very simple model on twig size optimization.