Vertical migration of rays leads to the development of a double-storied phenotype in the cambium of Aesculus turbinata

In the vascular cambium of Aesculus turbinata (Blume) the double-storied structure develops slowly. Initially, the arrangement of primary rays is nonstoried. New secondary rays are initiated during cambial expansion. Rays grow by addition of new initials at both ray margins and then split by the intrusive elongation of adjacent fusiform cells. The repetitive splits give rise to groups of several rays of common descent. Initially, the secondary rays are also nonstoried. Later, they become organized into horizontal tiers. This results from the vertical migration of ray initials in the vascular cambium. Controlled polar additions and eliminations of ray-cell initials at the opposite margins of the ray continue until it reaches the appropriate position within the storey of fusiform initials. We postulate that there are at least two mechanisms for the formation and maintenance of ray tiers in cambium. They are unrelated to cell inclination changes, which as described earlier, are known to sometimes induce a double-storied phenotype. The first of these mechanisms, involves initiation of secondary rays exactly within the storeys of fusiform initials, as in Hippophaë rhamnoides L. The second mechanism, present in A. turbinata, is based on the dynamic, controlled migration of rays.