The multiscale nature of leaf growth fields

Abstract

A growing leaf is a prototypical active solid, as its active units, the cells, locally deform during the out-of-equilibrium process of growth. During this local growth, leaves increase their area by orders of magnitude, yet maintain a proper shape, usually flat. How this is achieved in the lack of a central control, is unknown. Here we measure the in-plane growth tensor of Tobacco leaves and study the statistics of growth-rate, isotropy and directionality. We show that growth strongly fluctuates in time and position, and include multiple shrinkage events. We identify the characteristic scales of the fluctuations. We show that the area-growth distribution is broad and non-Gaussian, and use multiscale statistical methods to show how growth homogenizes at larger/longer scales. In contrast, we show that growth isotropy does not homogenize in time. Mechanical analysis shows that with such growth statistics, a leaf can stay flat only if the fluctuations are regulated/correlated.