Zwieniecki Maciej A, Boyce C Kevin, Holbrook N Michele
Department of Organismic and Evolutionary Biology, Harvard University, 16 Divinity Ave, Cambridge, MA 02138, USA.
Ann Bot. 2004 Oct;94(4):507-13. doi: 10.1093/aob/mch173. Epub 2004 Aug 19.
Morphological diversity of leaves is usually quantified with geometrical characters, while in many cases a simple set of biophysical parameters are involved in constraining size and shape. One of the main physiological functions of the leaf is transpiration and thus one can expect that leaf hydraulic parameters can be used to predict potential morphologies, although with the caveat that morphology in turn influences physiological parameters including light interception and boundary layer thickness and thereby heat transfer and net photosynthesis.
An iterative model was used to determine the relative sizes and shapes that are functionally possible for single-veined leaves as defined by their ability to supply the entire leaf lamina with sufficient water to prevent stomatal closure. The model variables include the hydraulic resistances associated with vein axial and radial transport, as well as with water movement through the mesophyll and the leaf surface.
The four parameters included in the model are sufficient to define a hydraulic functional design space that includes all single-veined leaf shapes found in nature, including scale-, awl- and needle-like morphologies. This exercise demonstrates that hydraulic parameters have dissimilar effects: surface resistance primarily affects leaf size, while radial and mesophyll resistances primarily affect leaf shape.
These distinctions between hydraulic parameters, as well as the differential accessibility of different morphologies, might relate to the convergent evolutionary patterns seen in a variety of fossil lineages concerning overall morphology and anatomical detail that frequently have evolved in linear and simple multi-veined leaves.
叶片的形态多样性通常用几何特征来量化,而在许多情况下,一组简单的生物物理参数也参与了对叶片大小和形状的限制。叶片的主要生理功能之一是蒸腾作用,因此可以预期叶片水力参数可用于预测潜在的形态,不过需要注意的是,形态反过来也会影响包括光截获和边界层厚度在内的生理参数,进而影响热传递和净光合作用。
使用一个迭代模型来确定单脉叶片在功能上可能的相对大小和形状,单脉叶片的定义是其有能力为整个叶片提供足够的水分以防止气孔关闭。模型变量包括与叶脉轴向和径向运输相关的水力阻力,以及水分通过叶肉和叶片表面的移动阻力。
模型中包含的四个参数足以定义一个水力功能设计空间,该空间涵盖了自然界中发现的所有单脉叶片形状,包括鳞片状、锥状和针状形态。这项研究表明水力参数有不同的影响:表面阻力主要影响叶片大小,而径向和叶肉阻力主要影响叶片形状。
水力参数之间的这些差异,以及不同形态的不同可达性,可能与多种化石谱系中在整体形态和解剖细节方面常见的趋同进化模式有关,这些模式常常在线性和简单多脉叶片中进化而来。