Growth synchrony between leaves and stems during twig development differs among plant functional types of subtropical rainforest woody species.

Abstract

During the development of woody twigs, the growth in leaf may or may not be proportional to the growth in stem. The presence or absence of a synchronicity between these two phenologies may reflect differences in life history adaptive strategies concerning carbon gain. We hypothesized that sun-adapted species are more likely to be less synchronous between growths in total leaf area (TLA) and stem length compared with shade-adapted species, with a bias in growth in stem length, and that shade-adapted species are more likely to be more synchronous between increases in individual leaf area (ILA) (leaf size) and leaf number (LN) during twig development compared with sun-adapted species, giving priority to growth of leaf size. We tested these two hypotheses by recording the phenologies of leaf emergence, leaf expansion and stem elongation during twig development for 19 evergreen woody species (including five shade-adapted understory species, six sun-adapted understory species and eight sun-adapted canopy species) in a subtropical evergreen broad-leaved forest in eastern China. We constructed indices to characterize the synchronicity between TLA and stem length (αLS) and between leaf size and leaf number (αSN) and we derived the α values from logistic functions taking the general form of A = A(max)/[1 + exp(β - αB)] (where A is the TLA or average ILA, B is the corresponding stem length or LN at a specific time, and A(max) is the maximum TLA or the maximum ILA of a twig; the higher the numerical value of α, the less synchronous the corresponding phenologies). Consistent with our hypotheses, sun-adapted species were higher both in α(LS) and α(SN), showing less synchronous patterns in the growths of TLA vs stem length and leaf size vs LN during twig development. Moreover, α(LS) and α(SN) were significantly positively correlated with relative growth rates of LN and leaf size across species, as indicated by both analyses of ordinary regression and phylogenetic generalized least squares. The across-species synchronies during twig development show that the temporal dynamics of the leaf size-twig size spectrum is of adaptive significance in plants. We suggest that temporal dynamics of plant functional traits should be extensively studied to characterize plant life history.