López-Casado Gloria, Matas Antonio J, Domínguez Eva, Cuartero Jesús, Heredia Antonio
Departamento de Biología Molecular y Bioquímica, Universidad de Málaga, E-29071 Málaga, Spain.
J Exp Bot. 2007;58(14):3875-83. doi: 10.1093/jxb/erm233. Epub 2007 Nov 1.
The mechanical characteristics of the cuticular membrane (CM), a complex composite biopolymer basically composed of a cutin matrix, waxes, and hydrolysable polysaccharides, have been described previously. The biomechanical behaviour and quantitative contribution of cutin and polysaccharides have been investigated here using as experimental material mature green and red ripe tomato fruits. Treatment of isolated CM with anhydrous hydrogen fluoride in pyridine allowed the selective elimination of polysaccharides attached to or incrusted into the cutin matrix. Cutin samples showed a drastic decrease in elastic modulus and stiffness (up to 92%) compared with CM, which clearly indicates that polysaccharides incorporated into the cutin matrix are responsible for the elastic modulus, stiffness, and the linear elastic behaviour of the whole cuticle. Reciprocally, the viscoelastic behaviour of CM (low elastic modulus and high strain values) can be assigned to the cutin. These results applied both to mature green and red ripe CM. Cutin elastic modulus, independently of the degree of temperature and hydration, was always significantly higher for the ripe than for the green samples while strain was lower; the amount of phenolics in the cutin network are the main candidates to explain the increased rigidity from mature green to red ripe cutin. The polysaccharide families isolated from CM were pectin, hemicellulose, and cellulose, the main polymers associated with the plant cell wall. The three types of polysaccharides were present in similar amounts in CM from mature green and red ripe tomatoes. Physical techniques such as X-ray diffraction and Raman spectroscopy indicated that the polysaccharide fibres were mainly randomly oriented. A tomato fruit CM scenario at the supramolecular level that could explain the observed CM biomechanical properties is presented and discussed.
角质膜(CM)是一种复杂的复合生物聚合物,主要由角质基质、蜡质和可水解多糖组成,其机械特性此前已有描述。本文以成熟的绿色和红色成熟番茄果实为实验材料,研究了角质和多糖的生物力学行为及定量贡献。用吡啶中的无水氟化氢处理分离出的CM,可以选择性地去除附着在角质基质上或嵌入角质基质中的多糖。与CM相比,角质样品的弹性模量和刚度大幅下降(高达92%),这清楚地表明,掺入角质基质中的多糖决定了整个角质层的弹性模量、刚度和线性弹性行为。相反,CM的粘弹性行为(低弹性模量和高应变值)可归因于角质。这些结果适用于成熟的绿色和红色成熟CM。无论温度和水合程度如何,成熟样品的角质弹性模量总是显著高于绿色样品,而应变则较低;角质网络中的酚类物质含量是解释从成熟绿色角质到红色成熟角质硬度增加的主要因素。从CM中分离出的多糖家族有果胶、半纤维素和纤维素,它们是与植物细胞壁相关的主要聚合物。成熟绿色和红色成熟番茄的CM中,这三种多糖的含量相似。X射线衍射和拉曼光谱等物理技术表明,多糖纤维主要是随机取向的。本文提出并讨论了一个在超分子水平上的番茄果实CM模型,该模型可以解释观察到的CM生物力学特性。