Seasonal Dynamics of Shoot Growth in Forsythia ovata Nakai Plants: Rhythmicity of Apical and Radial Growth

S. A. Shavnin, A. A. Montile, L. A. Semkina, A. I. Montile
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Abstract

The aim of the study was to establish the nature (linear or nonlinear) and shape (presence of extremes and trends) of time dependences of apical and radial growth rates for shoots of two types (branching and formation) in woody plants using the example of an ornamental early-flowering shrub species introduced in the Middle Urals Forsythia ovata Nakai. The seasonal dynamics of morphometric parameters of shoots differing in growth intensity and origin (growing from the apical and dormant buds of the skeletal branch) were studied, and their relationship to air temperature and precipitation values was assessed. Both types of growth are accompanied by oscillations of increments, the duration of individual phases of which is about 7 days or more, which allows us to attribute this phenomenon to infradian growth rhythms. The weak relationship between the characteristics of the observed oscillations and weather conditions indicates the predominantly endogenous nature of the morphogenetic processes that determine them. The growth of formation shoots begins 4 weeks later than branching shoots, which is apparently associated with the time necessary for the activation of dormant bud growth. Seasonal dynamics curves of the rates and accelerations of apical and radial growth of shoots have the form of oscillations with phases decaying in amplitude and varying in duration that have not been observed before, which ends 1–2 months before the onset of unfavorable environmental conditions for growth. The processes that cause changes in the rate of shoot growth are apparently associated with the action of two or more differently directed factors that determine the characteristics and temporal stipulation for the increase in the volume of different shoot segments (including the different contribution of cell division and elongation to changes in the geometric dimensions of the apical meristem domains). In both types of shoots, there are differences in the seasonal dynamics of the apical and radial increments in terms of the growth duration (6–8 and 4 weeks, respectively), the number of extremes, and the amplitude and duration of individual phases of oscillations. The curves of the dynamics of the apical growth rates have two maxima, and those of the radial growth have only one. Each type of growth, despite the delay in its onset in the formation shoots, has the same total duration in different types of shoots, and the shapes of the curves of seasonal changes in rates and accelerations in different types of shoots are similar. The similarity of the curve shapes of the seasonal dynamics of individual growth types in different types of shoots indicates the presence of identical growth regulation programs in the shoots, ultimately determined by the genotype, which are manifested in the observed oscillations. The growth rhythmicity of both types of shoots during the season allows us to assume the existence of a temporal relationship between the rates of division and growth of individual cells located in functionally different groups of cells in the apex zone. The differences of dynamics in different types of shoots are that the amplitude of oscillations is higher in formation shoots than in branching shoots. This feature explains the well-known fact of higher values of their total increments.

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中井连翘嫩枝生长的季节动态:顶端和径向生长的节律性
摘要 该研究旨在以乌拉尔中部地区引进的观赏性早花灌木物种连翘(Forsythia ovata Nakai)为例,确定木本植物两种类型(分枝和形成)芽的顶端和径向生长率的时间依赖性的性质(线性或非线性)和形状(存在极端和趋势)。研究了不同生长强度和起源(从骨架枝的顶芽和休眠芽生长)的嫩枝形态计量参数的季节动态,并评估了它们与气温和降水值的关系。这两种生长类型都伴随着增量振荡,单个阶段的持续时间约为 7 天或更长,因此我们可以将这种现象归因于昼夜生长节律。观察到的振荡特征与天气条件之间的关系不大,这表明决定振荡的形态发生过程主要是内生性的。形成芽的生长开始时间比分枝芽晚 4 周,这显然与激活休眠芽生长所需的时间有关。嫩枝顶端和径向生长的速率和加速度的季节动态曲线呈振荡形式,振幅阶段性衰减,持续时间长短不一,这是以前从未观察到的,在不利于生长的环境条件开始前 1-2 个月结束。导致嫩枝生长速度变化的过程显然与两个或两个以上不同方向的因素的作用有关,这些因素决定了不同嫩枝段体积增加的特征和时间规定(包括细胞分裂和伸长对顶端分生组织区域几何尺寸变化的不同贡献)。在这两种类型的嫩枝中,顶端和径向增量的季节动态在生长持续时间(分别为 6-8 周和 4 周)、极值数量以及单个振荡阶段的振幅和持续时间方面存在差异。顶端增长率的动态曲线有两个最大值,而径向增长率的动态曲线只有一个最大值。尽管每种生长类型在形成芽中的起始时间都有延迟,但在不同类型的芽中,其总持续时间是相同的,而且不同类型芽中生长速率和加速度的季节变化曲线形状也很相似。不同类型芽中单个生长类型的季节动态曲线形状相似,表明芽中存在相同的生长调节程序,这些程序最终由基因型决定,表现为观察到的振荡。两类嫩枝在季节中的生长节律性使我们可以假定,位于顶区功能不同的细胞群中的单个细胞的分裂率和生长率之间存在时间关系。不同类型嫩枝的动态差异在于,形成期嫩枝的振荡幅度要高于分枝期嫩枝。这一特点解释了它们的总增量值更高这一众所周知的事实。
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