Department of Biological Sciences, Macquarie University, NSW 2109, Australia.
Ann Bot. 2011 Sep;108(3):419-28. doi: 10.1093/aob/mcr166.
A long-running debate centres on whether shade tolerance of tree seedlings is mainly a function of traits maximizing net carbon gain in low light, or of traits minimizing carbon loss. To test these alternatives, leaf display, light-interception efficiency, and simulated net daily carbon gain of juvenile temperate evergreens of differing shade tolerance were measured, and how these variables are influenced by ontogeny was queried.
The biomass distribution of juveniles (17-740 mm tall) of seven temperate rainforest evergreens growing in low (approx. 4 %) light in the understorey of a second-growth stand was quantified. Daytime and night-time gas exchange rates of leaves were also determined, and crown architecture was recorded digitally. YPLANT was used to model light interception and carbon gain.
An index of species shade tolerance correlated closely with photosynthetic capacities and respiration rates per unit mass of leaves, but only weakly with respiration per unit area. Accumulation of many leaf cohorts by shade-tolerant species meant that their ratios of foliage area to biomass (LAR) decreased more gradually with ontogeny than those of light-demanders, but also increased self-shading; this depressed the foliage silhouette-to-area ratio (STAR), which was used as an index of light-interception efficiency. As a result, displayed leaf area ratio (LAR(d) = LAR × STAR) of large seedlings was not related to species shade tolerance. Self-shading also caused simulated net daily carbon assimilation rates of shade-tolerant species to decrease with ontogeny, leading to a negative correlation of shade tolerance with net daily carbon gain of large (500 mm tall) seedlings in the understorey.
The results suggest that efficiency of energy capture is not an important correlate of shade tolerance in temperate rainforest evergreens. Ontogenetic increases in self-shading largely nullify the potential carbon gain advantages expected to result from low respiration rates and long leaf lifespans in shade-tolerant evergreens. The main advantage of their long-lived leaves is probably in reducing the costs of crown maintenance.
一个长期存在的争论集中在树木幼苗的耐荫性主要是最大限度地提高在低光照下净碳增益的特征的功能,还是最大限度地减少碳损失的特征。为了检验这些替代方案,测量了不同耐荫性的幼年温带常绿树种的叶片展示、光截获效率和模拟的净日碳增益,以及询问了这些变量如何受个体发育的影响。
在第二生长林林下低光照(约 4%)下生长的 7 种温带雨林常绿树种的幼树(17-740 毫米高)的生物量分布进行了量化。还测定了叶片的日间和夜间气体交换率,并数字化记录了树冠结构。使用 YPLANT 来模拟光截获和碳增益。
物种耐荫性指数与叶片的光合能力和呼吸速率密切相关,但与单位面积呼吸速率的相关性较弱。耐荫物种积累了许多叶群,这意味着它们的叶面积与生物量比(LAR)随个体发育的变化比需光者更缓慢,但也增加了自遮荫;这降低了叶形轮廓与面积比(STAR),它被用作光截获效率的指标。因此,大幼苗的显示叶面积比(LAR(d)=LAR×STAR)与物种耐荫性无关。自遮荫还导致耐荫物种的模拟日净碳同化率随个体发育而下降,导致林下大(500 毫米高)幼苗的耐荫性与净日碳增益呈负相关。
结果表明,能量捕获效率不是温带雨林常绿树种耐荫性的重要相关因素。自遮荫的个体发育增加在很大程度上抵消了耐荫常绿树种低呼吸速率和长叶片寿命所预期带来的潜在碳增益优势。它们长寿命叶片的主要优势可能在于降低树冠维护成本。