Department of Biology, Nanjing University, Nanjing 210093, China.
Tree Physiol. 2013 Jul;33(7):753-62. doi: 10.1093/treephys/tpt053.
In theory, plants can alter the distribution of leaves along the lengths of their twigs (i.e., within-twig leaf distribution patterns) to optimize light interception in the context of the architectures of their leaves, branches and canopies. We hypothesized that (i) among canopy tree species sharing similar light environments, deciduous trees will have more evenly spaced within-twig leaf distribution patterns compared with evergreen trees (because deciduous species tend to higher metabolic demands than evergreen species and hence require more light), and that (ii) shade-adapted evergreen species will have more evenly spaced patterns compared with sun-adapted evergreen ones (because shade-adapted species are generally light-limited). We tested these hypotheses by measuring morphological traits (i.e., internode length, leaf area, lamina mass per area, LMA; and leaf and twig inclination angles to the horizontal) and physiological traits (i.e., light-saturated net photosynthetic rates, Amax; light saturation points, LSP; and light compensation points, LCP), and calculated the 'evenness' of within-twig leaf distribution patterns as the coefficient of variation (CV; the higher the CV, the less evenly spaced leaves) of within-twig internode length for 9 deciduous canopy tree species, 15 evergreen canopy tree species, 8 shade-adapted evergreen shrub species and 12 sun-adapted evergreen shrub species in a subtropical broad-leaved rainforest in eastern China. Coefficient of variation was positively correlated with large LMA and large leaf and twig inclination angles, which collectively specify a typical trait combination adaptive to low light interception, as indicated by both ordinary regression and phylogenetic generalized least squares analyses. These relationships were also valid within the evergreen tree species group (which had the largest sample size). Consistent with our hypothesis, in the canopy layer, deciduous species (which were characterized by high LCP, LSP and Amax) had more even leaf distribution patterns than evergreen species (which had low LCP, LSP and Amax); shade-adapted evergreen species had more even leaf distribution patterns than sun-adapted evergreen species. We propose that the leaf distribution pattern (i.e., 'evenness' CV, which is an easily measured functional trait) can be used to distinguish among life-forms in communities similar to the one examined in this study.
从理论上讲,植物可以改变其枝条上叶子的分布(即,枝内叶分布模式),以优化其叶子、枝条和树冠的架构下的光捕获。我们假设:(i)在具有相似光照环境的冠层树种中,落叶树的枝内叶分布模式会比常绿树更均匀(因为落叶树种比常绿树的代谢需求更高,因此需要更多的光),(ii)耐阴常绿树种的分布模式会比喜光常绿树种更均匀(因为耐阴树种通常受光限制)。为了验证这些假设,我们通过测量形态特征(即节间长度、叶面积、比叶面积、LMA;以及叶片和枝条与水平的倾斜角度)和生理特征(即光饱和净光合速率、Amax;光饱和点、LSP;和光补偿点、LCP),并计算枝内叶分布模式的“均匀性”,作为枝内节间长度的变异系数(CV;CV 越高,叶片的分布越不均匀),来测量 9 种落叶乔木、15 种常绿乔木、8 种耐阴常绿灌木和 12 种喜光常绿灌木在华东亚热带阔叶林的枝内分布。变异系数与大 LMA 和大叶片和枝条倾斜角度呈正相关,这两个特征共同指定了一个典型的适应低光捕获的特征组合,这在普通回归和系统发育广义最小二乘法分析中都得到了验证。这些关系在常绿树种组内也是有效的(该组具有最大的样本量)。与我们的假设一致,在冠层中,落叶树种(其特征是高 LCP、LSP 和 Amax)的叶分布模式比常绿树种更均匀(其特征是低 LCP、LSP 和 Amax);耐阴常绿树种的叶分布模式比喜光常绿树种更均匀。我们提出,叶片分布模式(即“均匀性”CV,这是一个易于测量的功能特征)可用于区分类似于本研究中所研究的群落中的生活型。