Structural integration at the shoot apical meristem: Models, measurements, and experiments

The shoot apical meristem (SAM) produces stem and initiates leaves. Its structure is maintained despite a continuous flow of cells basipetally from the distal portion of the meristem. The apoplasm and symplasm are the obvious means of cell integration, and their role in chemical cell-to-cell signaling is known. However, the cell wall apoplasm is most likely also involved in a mechanical integration mode, in which mechanical stress and strains (elastic and plastic strain, i.e., growth) are putative signaling factors. Shoot apex cells grow symplastically and their growth is in general anisotropic. Therefore tensor of growth rates that depends on the displacements caused by growth is the most suitable physical entity to describe growth. The tensor approach introduces the concept of principal directions of growth, i.e., the directions in which growth rates attain extremal values. Because of the symplastic mode of growth, the cell wall pattern within the shoot apical meristem informs us about the sequence and planes of cell divisions and about the deformation of existing walls. In consequence, within the meristem, periclines and anticlines can be recognized, both representing the principal directions of growth.