Horbens Melanie, Feldner Alexander, Höfer Monika, Neinhuis Christoph
Institute of Botany, Technische Universität Dresden, Zellescher Weg 20b, D-01217 Dresden, Germany.
Ann Bot. 2014 Jan;113(1):105-18. doi: 10.1093/aob/mct262. Epub 2013 Nov 27.
Apple (Malus) fruit peduncles are highly modified stems with limited secondary growth because fruit ripening lasts only one season. They must reliably connect rather heavy fruits to the branch and cope with increasing fruit weight, which induces dynamic stresses under oscillating wind loads. This study focuses on tissue modification of these small, exposed structures during fruit development.
A combination of microscopic, static and dynamic mechanical tests, as well as Raman spectroscopy, was used to study structure-function relationships in peduncles of one cultivar and 12 wild species, representatively chosen from all sections of the genus Malus. Tissue differentiation and ontogenetic changes in mechanical properties of Malus peduncles were observed throughout one growing season and after successive removal of tissues.
Unlike in regular stems, the vascular cambium produces mainly phloem during secondary growth. Hence, in addition to a reduced xylem, all species developed a centrally arranged sclerenchyma ring composed of fibres and brachysclereids. Based on differences in cell-wall thickness, and proportions and arrangement of sclereids, two types of peduncle construction could be distinguished. Fibres provide an increased maximum tensile strength and contribute most to the overall axial rigidity of the peduncles. Sclereids contribute insignificantly to peduncle strength; however, despite being shown to have a lower elastic modulus than fibres, they are the most effective tissue in stiffening peduncles against bending.
The experimental data revealed that sclereids originating from cortical parenchyma act as 'accessory' cells to enhance proportions of sclerenchyma during secondary growth in peduncles. The mechanism can be interpreted as an adaptation to continuously increasing fruit loads. Under oscillating longitudinal stresses, sclereids may be regarded as regulating elements between maintenance of stiffness and viscous damping, the latter property being attributed to the cortical parenchyma.
苹果(Malus)果梗是高度特化的茎,次生生长有限,因为果实成熟仅持续一个季节。它们必须可靠地将较重的果实连接到树枝上,并应对不断增加的果实重量,这会在振荡风荷载下产生动态应力。本研究聚焦于这些小型外露结构在果实发育过程中的组织变化。
结合显微镜观察、静态和动态力学测试以及拉曼光谱,研究了从苹果属所有组中代表性选取的一个栽培品种和12个野生种的果梗结构与功能的关系。在一个生长季节以及连续去除组织后,观察苹果果梗的组织分化和力学性能的个体发育变化。
与正常茎不同,维管形成层在次生生长过程中主要产生韧皮部。因此,除了木质部减少外,所有物种都发育出了由纤维和石细胞组成的中央排列的厚壁组织环。基于细胞壁厚度、石细胞比例和排列的差异,可以区分出两种类型的果梗结构。纤维提供了更高的最大拉伸强度,对果梗的整体轴向刚度贡献最大。石细胞对果梗强度的贡献不大;然而,尽管其弹性模量低于纤维,但它们是增强果梗抗弯曲能力最有效的组织。
实验数据表明,源自皮层薄壁组织的石细胞在果梗次生生长过程中作为“辅助”细胞,以增加厚壁组织的比例。该机制可解释为对不断增加的果实负荷的一种适应。在振荡纵向应力下,石细胞可被视为维持刚度和粘性阻尼之间调节元件,后者归因于皮层薄壁组织。