Signals flowing from mature tissues to shoot apical meristem affect phyllotaxis in coniferous shoot.

Abstract

Axial homodromy in growing shoots of perennial plants with spiral phyllotaxis is the case when the chirality of phyllotactic pattern does not change in consecutive growth increments of the same axis. In conifers such as Picea or Abies this rule is strictly observed, except for the rare cases of discontinuous phyllotactic transitions. In Torreya, however, the chirality changes, at random, every year. The pattern of primordia packing, executed by vegetative shoot apical meristem (SAM), depends in Torreya on their identity. The primordia of bud scales are initiated in the decussate and those of needles in bijugate spiral pattern. The decussate, achiral i.e. neutral pattern always precedes the formation of new spiral pattern and thus facilitates random selection of its chiral configuration. Periodic change in organ identity cannot itself be responsible for the special behavior of Torreya, because in other conifers it also exists. There is, however, one important difference: in Torreya, when the initiation of bud scales begins at SAM, the distance between differentiated protoxylem and the initiation site gradually increases, while in other conifers it remains constant and small. In Torreya, at this phase of development, the rate of xylem differentiation and the rate of organogenesis become uncoupled. Closer anatomical examination shows that the decussate pattern in a bud scale zone develops slowly suggesting gradual decrease of the putative signal flowing acropetally from differentiated protoxylem, responsible for positioning of primordia. We hypothesize that in the absence of this signal SAM starts acting autonomously, distributing primordia according to their identity only. A constant presence of the signal in other conifers assures the continuation of the same phyllotactic pattern throughout the period of bud scale formation, despite the change in organ identity.