Givnish Thomas J, Kriebel Ricardo
Department of Botany, University of Wisconsin-Madison, Madison, Wisconsin 53706
Department of Botany, University of Wisconsin-Madison, Madison, Wisconsin 53706.
Am J Bot. 2017 Mar;104(3):354-366. doi: 10.3732/ajb.1600287. Epub 2017 Feb 22.
A recent commentary by Edwards et al. (. 103: 975-978) proposed that constraints imposed by the packing of young leaves in buds could explain the positive association between non-entire leaf margins and latitude but did not thoroughly consider alternative explanations.
We review the logic and evidence underlying six major hypotheses for the functional significance of marginal teeth, involving putative effects on (1) leaf cooling, (2) optimal support and supply of the areas served by major veins, (3) enhanced leaf-margin photosynthesis, (4) hydathodal function, (5) defense against herbivores, and (6) bud packing.
Theoretical and empirical problems undermine all hypotheses except the support-supply hypothesis, which implies that thinner leaves should have non-entire margins. Phylogenetically structured analyses across angiosperms, the El Yunque flora, and the genus all demonstrate that non-entire margins are indeed more common in thinner leaves. Across angiosperms, the association of leaf thickness with non-entire leaf margins is stronger than that of latitude.
We outline a synthetic model showing how biomechanics, hydraulics, vein geometry, rates of leaf expansion, and length of development within resting buds, all tied to leaf thickness, drive patterns in the distribution of entire vs. non-entire leaf margins. Our model accounts for dominance of entire margins in the tropics, Mediterranean scrub, and tundra, non-entire margins in cold temperate deciduous forests and tropical vines and early-successional trees, and entire leaf margins in monocots. Spinose-toothed leaves should be favored in short-statured evergreen trees and shrubs, primarily in Mediterranean scrub and related semiarid habitats.
爱德华兹等人最近的一篇评论文章(《植物学报》103卷:975 - 978页)提出,芽中幼叶的排列方式所施加的限制可以解释非全缘叶边缘与纬度之间的正相关关系,但未充分考虑其他解释。
我们回顾了关于叶缘齿功能意义的六个主要假说的逻辑和证据,这些假说涉及对以下方面的假定影响:(1)叶片降温;(2)主要叶脉所服务区域的最佳支撑和供应;(3)增强叶缘光合作用;(4)排水器功能;(5)抵御食草动物;(6)芽的排列。
理论和实证问题削弱了除支撑 - 供应假说之外的所有假说,该假说意味着较薄的叶片应该具有非全缘边缘。对被子植物、埃尔云克植物区系以及该属进行的系统发育结构分析均表明,非全缘边缘在较薄叶片中确实更为常见。在被子植物中,叶片厚度与非全缘叶边缘的关联比与纬度的关联更强。
我们概述了一个综合模型,展示了生物力学、水力学、叶脉几何形状、叶片扩展速率以及休眠芽内的发育时长,所有这些都与叶片厚度相关,如何驱动全缘叶边缘与非全缘叶边缘分布模式的形成。我们的模型解释了全缘叶边缘在热带、地中海灌丛和冻原中的优势,非全缘叶边缘在寒温带落叶林、热带藤本植物和早期演替树木中的情况,以及单子叶植物中全缘叶边缘的情况。具刺齿叶应在矮生常绿树和灌木中更受青睐,主要在地中海灌丛及相关半干旱生境中。