Cherdantsev V G, Grigor'eva O V
Ontogenez. 2010 May-Jun;41(3):171-89.
The stages of the early morphogenesis of simple (unbranched) and complex (branched) unicellular trichomes are studied in two species of the genus Draba--D. sibirica (Pall.) Thell. and D. daurica DC. The geometry of morphogenesis is estimated by analyzing intraindividual variation of quantitative morphological characteristics of the developing leaf blade and peduncle trichomes. The surface of all types oftrichome cells first acquires a spherical shape, followed by a U-shaped configuration with cylindrical proximal and spherical distal regions. In the development of complex trichomes, the area of the distal zone grows at a higher rate, which leads to separation of its volume into individual spherical regions, the morphogenesis of which repeats the early morphogenetic stages of the overall trichome cell, forming simple (unbranched) or complex (branched) trichome rays. As a rule, the lateral polarity of a trichome cell coincides with the proximodistal polarity of the leaf. Quantitative morphological data make it possible to infer an algorithm of the changes in shape common for all trichome cells, namely, the growth cycle comprising alternation of the phases of increase and decrease in the curvature of the outer cell surface. This surface is an active membrane expanded by the internal pressure and concurrently capable of actively increasing its area by incorporation of new structural elements. A distinctive feature of the proposed model is the geometrical inhomogeneity of the surface movement, changing the radius of curvature and creating internal (active) mechanical stresses in this membrane. A decrease in the ratio of the membrane surface area to the volume deprives the spatially homogeneous shape of its stability; correspondingly, the transition from elastic resistance to internal pressure to active resistance with the help of curvature differentiation becomes more energetically favorable. The source for growth and morphogenesis of the active membrane is alternation of the phases of local curvature leveling, which "charges" the membrane with active mechanical stresses and "discharge" of these stresses, leading to differentiation of the membrane's local curvatures.
在葶苈属的两个物种——西伯利亚葶苈(Draba sibirica (Pall.) Thell.)和兴安葶苈(Draba daurica DC.)中,研究了简单(无分支)和复杂(有分支)单细胞毛状体早期形态发生的阶段。通过分析发育中的叶片和花葶毛状体定量形态特征的个体内变异来估计形态发生的几何特征。所有类型毛状体细胞的表面首先呈球形,随后呈U形结构,其近端为圆柱形,远端为球形。在复杂毛状体的发育过程中,远端区域的面积增长速度更快,这导致其体积分离为单个球形区域,其形态发生重复了整个毛状体细胞的早期形态发生阶段,形成简单(无分支)或复杂(有分支)的毛状体射线。通常,毛状体细胞的侧向极性与叶片的近远极性一致。定量形态学数据使得推断所有毛状体细胞共有的形状变化算法成为可能,即生长周期包括外细胞表面曲率增加和减少阶段的交替。这个表面是一个由内部压力扩张的活性膜,同时能够通过纳入新的结构元件来积极增加其面积。所提出模型的一个显著特征是表面运动的几何不均匀性,改变曲率半径并在该膜中产生内部(活性)机械应力。膜表面积与体积之比的降低使其空间均匀形状失去稳定性;相应地,借助曲率分化从弹性抵抗内部压力到活性抵抗的转变在能量上变得更有利。活性膜生长和形态发生的来源是局部曲率平整阶段的交替,这给膜“加载”活性机械应力并“释放”这些应力,导致膜局部曲率的分化。