Early ontogeny defines the diversification of primary vascular bundle systems in angiosperms

Abstract
Understanding vascular bundle systems in angiosperms is a challenge. On the one hand, the vascular tissue is extremely important in reconstructing the evolution and survivability of plants, but on the other hand, it forms a complicated three-dimensional system of controversially discussed phylogenetic and ontogenetic origin. To increase clarity, in this paper we briefly summarize histological, phylogenetic and molecular aspects of primary vascular bundle formation in angiosperms, adding histological data on early stages of vascular bundle formation, proposing a concept combining the different views and providing simplified illustrations to improve the understanding of primary vascular systems in angiosperms. Based on the auxin hypothesis, vascular bundle formation is stimulated by the development of leaf primordia. Provascular domains appear at the base of the leaf primordia and develop into two directions (leaf, internode). The low vs. high number of internodal bundles, their circular vs. scattered arrangement and the open vs. closed bundle construction in eudicots vs. monocots can be deduced to be due to processes below the shoot apical meristem. The most important processes distinguishing monocots from eudicots are the isolated bundle initiation outside the primary meristem, the enormous expansion of the leaf bases associated with a high number of vascular bundles and the early onset of primary thickening passively dislocating vascular bundles.