Expression of shoot features in early leaf development of Murraya paniculata (Rutaceae)

Abstract

The early development of the pinnately compound leaves in Murraya paniculata was studied using both epi-illumination and scanning electron microscopy as well as semithin plastic sectioning of the same specimens that were illustrated by means of epi-illumination. It is shown that morphological conclusions may be influenced by technical approaches such as the plane of sectioning. If the developing leaves are sectioned in the (median) sagittal plane, they appear to be rather different from stems and shoots. If, however, they are sectioned in the frontal plane, perpendicular to the sagittal plane, they appear more shoot-like in early development. Their apex could be described in terms of a tunica-corpus organization and the leaflet primordia are initiated like leaf primordia on a shoot tip with distichous phyllotaxy sensu lato. Subsequently, due to differential growth, reorientation of the leaflets occurs in one plane. Thus, the planar structure of the pinnate leaf is ontogenetically secondary. From a phylogenetic perspective, at least two conclusions are possible for plants with pinnate leaves such as those of Murraya: (i) if the ancestor of a pinnate taxon had simple leaves, the pinnate condition arose through homeosis, i.e., the expression of shoot features in leaf sites; (ii) if the ancestor of a pinnate taxon did not have simple leaves, the shoot-like early development of the pinnate leaves may indicate a common evolutionary basis of shoots and pinnate leaves in primitive branching systems. Since it is generally thought that the most primitive angiosperms have simple leaves, the homeotic hypothesis appears to be the preferred hypothesis for the origin of compound leaves in flowering plants. Key words: leaf development, comparative morphogenesis, shoot–leaf relationships, partial shoot theory of the leaf, homeosis.