The logic of plant vascular patterning. Polarity, continuity and plasticity in the formation of the veins and of their networks

The problem of long-distance transport is solved in many multicellular organisms by tissue networks such as the vascular networks of plants. Because tissue networks transport from one tissue area to another, they are polar and continuous; most of them, including plant vascular networks, are also plastic. Surprisingly, the formation of tissue networks is in most cases just as polar, continuous and plastic. Available evidence suggests that the polarity, continuity and plasticity of plant vascular networks and their formation could be accounted for by a patterning process that combines: (i) excess of developmental alternatives competing for a limiting cell-polarizing signal; (ii) positive feedback between cell polarization and continuous, cell-to-cell transport of the cell-polarizing signal; and (iii) gradual restriction of differentiation that increasingly removes the cell-polarizing signal.