Ray Peter M, Bret-Harte Marion Syndonia
Department of Biological Sciences, Stanford University, Stanford, CA, United States.
Institute of Arctic Biology, University of Alaska Fairbanks, Fairbanks, AK, United States.
Front Plant Sci. 2019 Feb 5;10:59. doi: 10.3389/fpls.2019.00059. eCollection 2019.
Tree and shrub branches subjected to cantilever loads such as intercepted snowfall undergo, in addition to the familiar instantaneous elastic bending, a conspicuous retarded-elastic bending, which is commonly 30-50% of their instantaneous bending and occasionally even more. The resultant bending creep that occurs after loading also often includes a slow, time-dependent irreversible bending. These phenomena occur quite generally among woody plants of different major biomes, taxonomic groups, and structural types. We give some of branch bending viscoelasticity's basic physical properties such as load dependence and stress relaxation. These properties belong to the secondary walls of branches' xylem (wood) cells; some properties differ notably from those reported for primary cell walls, a difference for which we propose explanations. A method for separating the overlapping time courses of retarded-elastic and time-dependent irreversible bending shows that multiple retarded-elastic ("Kelvin") elements of branches span a wide range of retardation times (a retardation spectrum, approximate examples of which we calculate), and that irreversible bending can occur in different cases either only in the first few h after loading, or more extensively through 24 h, or (rarely) for several days. A separate time-independent irreversible bending, permanent set, involving a substantial yield stress, also occurs. In three species of shrubs rapid irreversible bending began only several (up to 24) h after loading, implying an unusual kind of viscoelasticity. Deductions from the dynamics of bending suggest that retarded elasticity can help protect branches against breakage by wind gusts during storms. Irreversible bending probably contributes both to the form that tree and shrub crowns develop over the long term, involving progressive increase in the downward curvature and/or inclination of branches, and also to certain other, more specialized, developmental changes.
承受诸如积雪等悬臂载荷的树木和灌木枝条,除了常见的瞬时弹性弯曲外,还会发生明显的延迟弹性弯曲,这种弯曲通常占其瞬时弯曲的30%-50%,偶尔甚至更多。加载后发生的弯曲蠕变通常还包括缓慢的、随时间变化的不可逆弯曲。这些现象在不同主要生物群落、分类群和结构类型的木本植物中普遍存在。我们给出了枝条弯曲粘弹性的一些基本物理特性,如载荷依赖性和应力松弛。这些特性属于枝条木质部(木材)细胞的次生壁;有些特性与报道的初生细胞壁的特性明显不同,我们对此差异提出了解释。一种分离延迟弹性和随时间变化的不可逆弯曲的重叠时间过程的方法表明,枝条的多个延迟弹性(“开尔文”)元件跨越了很宽的延迟时间范围(我们计算了延迟谱的近似示例),并且不可逆弯曲在不同情况下可能仅在加载后的最初几个小时内发生,或者在24小时内更广泛地发生,或者(很少)持续几天。还会发生一种与时间无关的单独不可逆弯曲,即永久变形,它涉及相当大的屈服应力。在三种灌木中,快速不可逆弯曲仅在加载后几小时(最多24小时)开始,这意味着一种不同寻常的粘弹性。从弯曲动力学得出的推论表明,延迟弹性有助于保护枝条在风暴期间免受阵风破坏。不可逆弯曲可能既有助于树木和灌木树冠长期形成的形态,包括枝条向下曲率和/或倾斜度的逐渐增加,也有助于某些其他更特殊的发育变化。