The PXY-CLE41 receptor ligand pair defines a multifunctional pathway that controls the rate and orientation of vascular cell division

Controlling the orientation of cell division is fundamental to the development of complex body plans. This is particularly apparent in plants, where development is determined by differential growth that results solely from changes in cell expansion and orientation of the cell division plane. Despite the fundamental importance of cell division orientation to plant development, the mechanisms regulating this process remain almost completely unknown. During vascular development, the meristematic cambial cells divide down their long axis in a highly orientated manner to generate clear files of cells. The receptor kinase PXY has previously be shown to be essential for this orientation. Here, we demonstrate that the division plane is determined by the interactions of PXY and its peptide ligand, CLE41. PXY is expressed within dividing meristematic cells of the procambium, whereas CLE41 localises to the adjacent phloem cells. Altering the pattern of CLE41 expression leads to a loss of cell division orientation and a dramatic loss of ordered vascular tissue development. By contrast, increasing phloem-specific expression of CLE41 results in more cell divisions, but the orientation of cell division is retained, leading to both increased and well-ordered vascular development. We demonstrate that PXY signalling is down-regulated by CLE41. This feedback mechanism is crucial in integrating the different roles of PXY signalling in controlling xylem differentiation, regulating the rate of vascular cell division and determining the orientation of cell division. Parallels with animal systems indicate that localised signalling from adjacent cells is a general mechanism for defining the plane of cell division.