Buckley Thomas N, Roberts David W
Environmental Biology Group and CRC for Greenhouse Accounting, Research School of Biological Sciences, The Australian National University, Canberra, ACT 2601, Australia.
Tree Physiol. 2006 Feb;26(2):145-57. doi: 10.1093/treephys/26.2.145.
Conventional wisdom holds that the ratio of leaf area to sapwood area (L/S) should decline during height (H) growth to maintain hydraulic homeostasis and prevent stomatal conductance (g(s)) from declining. We contend that L/S should increase with H based on a numerical simulation, a mathematical analysis and a conceptual argument: (1) numerical simulation--a tree growth model, DESPOT (Deducing Emergent Structure and Physiology Of Trees), in which carbon (C) allocation is regulated to maximize C gain, predicts L/S should increase during most of H growth; (2) mathematical analysis--the formal criterion for optimal C allocation, applied to a simplified analytical model of whole tree carbon-water balance, predicts L/S should increase with H if leaf-level gas exchange parameters including g(s) are conserved; and (3) conceptual argument--photosynthesis is limited by several substitutable resources (chiefly nitrogen (N), water and light) and H growth increases the C cost of water transport but not necessarily of N and light capture, so if the goal is to maximize C gain or growth, allocation should shift in favor of increasing photosynthetic capacity and irradiance, rather than sustaining g(s). Although many data are consistent with the prediction that L/S should decline with H, many others are not, and we discuss possible reasons for these discrepancies.
传统观点认为,在树高(H)生长过程中,叶面积与边材面积之比(L/S)应下降,以维持水力平衡并防止气孔导度(g(s))下降。我们基于数值模拟、数学分析和概念论证认为,L/S应随H增加:(1)数值模拟——一个树木生长模型DESPOT(推导树木的新兴结构和生理学),其中碳(C)分配受调节以最大化C获取量,预测在大部分H生长过程中L/S应增加;(2)数学分析——应用于整株树木碳-水平衡简化分析模型的最优C分配形式标准预测,如果包括g(s)在内的叶级气体交换参数保持不变,L/S应随H增加;以及(3)概念论证——光合作用受几种可替代资源(主要是氮(N)、水和光)限制,H生长增加了水分运输的C成本,但不一定增加N和光捕获的C成本,因此,如果目标是最大化C获取量或生长,分配应转向有利于增加光合能力和辐照度,而不是维持g(s)。尽管许多数据与L/S应随H下降的预测一致,但也有许多数据并非如此,我们讨论了这些差异的可能原因。