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Plant Cell Advance Online Publication
Published on April 22, 2005; 10.1105/tpc.105.031732

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Received February 11, 2005
Returned for revision February 11, 2005
Accepted March 18, 2005

COBRA, an Arabidopsis Extracellular Glycosylphosphatidylinositol-Anchored Protein, Specifically Controls Highly Anisotropic Expansion through Its Involvement in Cellulose Microfibril Orientation

François Roudier 1, Anita G. Fernandez 2, Miki Fujita 3, Regina Himmelspach 4, Georg H.H. Borner 5, Gary Schindelman 2, Shuang Song 1, Tobias I. Baskin 6, Paul Dupree 5, Geoffrey O. Wasteneys 3, and Philip N. Benfey 1*

1 Department of Biology, Duke University, Durham, North Carolina 27708
2 Department of Biology, New York University, New York, New York 10003
3 Department of Botany, University of British Columbia, Vancouver, British Columbia, Canada V6T 1Z4; Research School of Biological Sciences, Australian National University, Canberra ACT 2601, Australia
4 Department of Botany, University of British Columbia, Vancouver, British Columbia, Canada V6T 1Z4
5 Department of Biochemistry, University of Cambridge, CB2 1QW Cambridge, United Kingdom
6 Biology Department, University of Massachusetts, Amherst, Massachusetts 01003

* To whom correspondence should be addressed. E-mail: philip.benfey{at}duke.edu.

The orientation of cell expansion is a process at the heart of plant morphogenesis. Cellulose microfibrils are the primary anisotropic material in the cell wall and thus are likely to be the main determinant of the orientation of cell expansion. COBRA (COB) has been identified previously as a potential regulator of cellulose biogenesis. In this study, characterization of a null allele, cob-4, establishes the key role of COB in controlling anisotropic expansion in most developing organs. Quantitative polarized-light and field-emission scanning electron microscopy reveal that loss of anisotropic expansion in cob mutants is accompanied by disorganization of the orientation of cellulose microfibrils and subsequent reduction of crystalline cellulose. Analyses of the conditional cob-1 allele suggested that COB is primarily implicated in microfibril deposition during rapid elongation. Immunodetection analysis in elongating root cells revealed that, in agreement with its substitution by a glycosylphosphatidylinositol anchor, COB was polarly targeted to both the plasma membrane and the longitudinal cell walls and was distributed in a banding pattern perpendicular to the longitudinal axis via a microtubule-dependent mechanism. Our observations suggest that COB, through its involvement in cellulose microfibril orientation, is an essential factor in highly anisotropic expansion during plant morphogenesis.




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