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í Friml31 Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9, Canada; 2 Department of Botany, University of Toronto, Toronto, Ontario M5S 3B2, Canada; 3 Center for Plant Molecular Biology (ZMBP), Tübingen University, D-72076 Tübingen, Germany
The formation of the leaf vascular pattern has fascinated biologists for centuries. In the early leaf primordium, complex networks of procambial cells emerge from homogeneous subepidermal tissue. The molecular nature of the underlying positional information is unknown, but various lines of evidence implicate gradually restricted transport routes of the plant hormone auxin in defining sites of procambium formation. Here we show that a crucial member of the AtPIN family of auxin-efflux-associated proteins, AtPIN1, is expressed prior to pre-procambial and procambial cell fate markers in domains that become restricted toward sites of procambium formation. Subcellular AtPIN1 polarity indicates that auxin is directed to distinct "convergence points" in the epidermis, from where it defines the positions of major veins. Integrated polarities in all emerging veins indicate auxin drainage toward pre-existing veins, but veins display divergent polarities as they become connected at both ends. Auxin application and transport inhibition reveal that convergence point positioning and AtPIN1 expression domain dynamics are self-organizing, auxin-transport-dependent processes. We derive a model for self-regulated, reiterative patterning of all vein orders and postulate at its onset a common epidermal auxin-focusing mechanism for major-vein positioning and phyllotactic patterning.
[Keywords: Arabidopsis; AtPIN genes; polar auxin transport; procambium; leaf development; vascular patterning]
Received December 19, 2005; revised version accepted February 16, 2006.
E-MAIL berleth{at}botany.utoronto.ca; FAX (416) 978-5878.
Supplemental material is available at http://www.genesdev.org.
Article and publication are at http://www.genesdev.org/cgi/doi/10.1101/gad.1402406
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Genes & Dev. 2006 20: 922-926.
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