论文标题
叶片生长场的间歇性质
The Intermittent Nature of Leaf Growth Fields
论文作者
论文摘要
什么是允许组织或器官适当生长的一般原则?生长的叶子是这样一个系统的一个例子:它通过数量级来增加面积,并保持正确的(通常是平坦)的形状。没有中央控制单元,该如何实现?人们会认为,统一增长与粘弹性流变学的结合将允许这一点。在这里,我们表明完全相反的过程在作用:叶子表面的自然生长在时间和位置强烈波动。通过结合高分辨率测量和多尺度的统计分析,我们建议以叶片生长的方式变化范式。我们测量了3D拉格朗日坐标中烟叶的平面内组织生长,并研究了与张力生长场相关的三个标量场的统计数据:生长速率,其各向同性和方向性。我们确定生长速率波动的时间和长度尺度,并在高分辨率测量时捕获局部肿胀和收缩之间的大量切换。在较低的时空分辨率下,生长速率场变得光滑。相反,各向异性场随着时间的推移而增加。最后,我们发现白天和黑夜测得的生长之间存在显着差异。发现晚上的增长更加间歇性,相关长度较短,没有全球方向性。尽管其性质波动,但生长领域并不是随机的,因此携带有关生长调节的信息。实际上,机械分析表明,只有在调节/相关的测量波动时,生长的叶子才能保持平坦。我们的测量表明,应研究整个生长领域的统计数据,而不仅仅是其手段。特别是,应研究此类场的调节及其特征与叶片的全局几何形状之间的联系。
What are the general principles that allow proper growth of a tissue or an organ? A growing leaf is an example of such a system: it increases its area by orders of magnitude, maintaining a proper (usually flat) shape. How can this be achieved without a central control unit? One would think that a combination of uniform growth with viscoelastic rheology would allow that. Here we show that that the exact opposite process is in action: the natural growth of the leaf surface strongly fluctuates in time and position. By combining high resolution measurements and multi-scale statistical analysis, we suggest a paradigm-change in the way leaf growth is viewed. We measure the in-plane tissue growth of Tobacco leaves in Lagrangian coordinates in 3D and study the statistics of three scalar fields associated with the tensorial growth field: The growth rate, its isotropy and directionality. We identify the governing time and length scales of the fluctuations in the growth rate, and capture abundant switching between local area swelling and shrinking, when measured in high resolution. At lower spatio-temporal resolution the growth rate field becomes smooth. In contrast, the anisotropy field increases over time. Finally, we find significant differences between growth measured during day and night. The growth at night is found to be more intermittent, with shorter correlation lengths and no global directionality. Despite its fluctuative nature, growth fields are not random, thus carry information about growth regulation. Indeed, mechanical analysis shows that a growing leaf can stay flat only if the measured fluctuations are regulated/correlated. Our measurements suggest that the entire statistics of growth fields, and not just their means, should be studied. In particular, the regulation of such fields and the link between their characteristics and the global geometry of a leaf should be studied.