Pattern formation of plant leaf venation
Generating venation of plants is an intriguing pattern formation in
biological systems. Three hypotheses have been proposed to explain
the venation pattern formation: auxin canalization hypothesis,
activator-inhibitor-type reaction-diffusion hypothesis, and
substrate-depletion-type reaction-diffusion hypothesis which is
recently proposed by S. Tohya and A.M. The first hypothesis is based
on the assumption of the positive feedback regulation between auxin
flux and localization of PIN1 auxin efflux carrier on plasma
membrane. Auxin is a diffusible plant hormone of small molecule and
is thought to be important for vascular development. Here we
investigated a model based on the assumption of this feedback
regulation.
First, this model can generate auxin flow pathways with high
density, so-called “polar auxin transport”, from almost uniform
field. Using this feature we tried to generate various leaf venation
patterns. Venation patterns largely depend on the shape of leaf area.
For example, dicot pattern and Ginkgo pattern are generated from
different diamond-shaped leaves. Place and direction of cell division
also affects venation pattern crucially. In the case of cell division
at apical leaf part, secondary veins elongate preferably toward
apical side. In contrast cell division at basal part generates
secondary veins that elongate toward basal side like Arabidopsis.
Under the condition of cell division at the base of a leaf like
monocots, parallel vein pattern is generated. Moreover, anisotropy of
auxin diffusion has also large effect on pattern formation; veins
tend to elongate along more diffusible direction. These results
indicate that the positive feedback dynamics between auxin flow and
PIN1 localization generates diverse venation patterns of plants
depending on various conditions such as leaf shape, cell division and
anisotropy.