Early stages of initiation of two types of leaves in Thuja occidentalis (eastern white cedar)

Abstract

The developmental morphology of shoots of Thuja occidentalis L. (eastern white cedar) was investigated using scanning electron microscopy to determine the pattern of initiation of two types of leaves characteristic of higher (third and above) order branches. The shoots of eastern white cedar are bilateral in symmetry and bear leaves in an orthogonal decussate phyllotactic pattern. The shoot system is further characterized by the presence of two alternating and morphologically different pairs of leaves that constitute the basic repeating pattern of the shoot apical meristem (SAM). At maturity the dimorphism between leaf types is marked. Leaves in one plane are wide and flat in comparison with narrower and cup-shaped leaves growing in a plane perpendicular to the other leaf type. The early stages of development of each of the two types of leaves were compared using scanning electron microscopy. During the earliest visible stages of initiation (primordial crest), cup-shaped and flat leaves are very similar in morphology. As individual leaf primordia become more easily delimited as structures by the presence of a furrow between the SAM and the leaf, they differ in terms of width. As they develop further and begin to cover the SAM, the two leaf types are distinguishable morphologically (flat vs. cup shaped). Quantitative parameters such as diameter of the SAM, angle of insertion of individual leaves, and size of leaf primordia (in both a tangential and perpendicular plane) were measured on three categories of leaves: stage 1, earliest visible stage of initiation; stage 2, delineation of leaf primordium from SAM by furrowing; stage 3, leaf primordium partially covering SAM. These measurements corroborate our morphological observations, which show that during early stages of development, flat and cup-shaped leaves are morphologically similar and they diverge in their pattern of development postinitiation, especially as far as leaf width and thickness are concerned. Our results also suggest that the size and shape of the apex goes through a "repeating" cycle and is related to the type of primordium that will be initiated.Key words: Thuja occidentalis, eastern white cedar, leaf development, shoot apical meristem, phyllotaxy, leaf dimorphism.