Growth and morphogenesis at the vegetative shoot apex of Anagallis arvensis L

A non-destructive replica method and a 3-D reconstruction algorithm are used to analyse the geometry and expansion of the shoot apex surface. Surface expansion in the central zone of the apex is slow and nearly isotropic while surface expansion in the peripheral zone is more intense and more anisotropic. Within the peripheral zone, the expansion rate, expansion anisotropy, and the direction of maximal expansion vary according to the age of adjacent leaf primordia. For each plastochron, this pattern of expansion is rotated around the apex by the Fibonacci angle. Early leaf primordium development is divided into four stages: bulging, lateral expansion, separation, and bending. These stages differ in their geometry and expansion pattern. At the bulging stage, the site of primordium initiation shows an intensified expansion that is nearly isotropic. The following stages develop sharp meridional gradients of expansion rates and anisotropy. The adaxial primordium boundary inferred from the surface curvature is shifting until the separation stage, when a crease develops between the primordium and the apex dome. The cells forming the crease, i.e. the future leaf axil, expand along the axil and contract across it. Thus they are arrested in this unique position.