Plant Cell Advance Online Publication Published on December 11, 2003; 10.1105/tpc.016550
Received August 22, 2003
Accepted October 21, 2003
Overexpression of an Arabidopsis Formin Stimulates Supernumerary Actin Cable Formation
from Pollen Tube Cell Membrane
Alice Y. Cheung 1* and Hen-ming Wu 1
1
Department of Biochemistry and Molecular Biology, Molecular and Cellular Biology
Program, Plant Biology Graduate Program, University of Massachusetts, Amherst, Massachusetts
01003
* To whom correspondence should be addressed. E-mail: acheung{at}biochem.umass.edu.
Formins, actin-nucleating proteins that stimulate the de novo polymerization of actin
filaments, are important for diverse cellular and developmental processes, especially
those dependent on polarity establishment. A subset of plant formins, referred to
as group I, is distinct from formins from other species in having evolved a unique
N-terminal structure with a signal peptide, a Pro-rich, potentially glycosylated
extracellular domain, and a transmembrane domain. We show here that overexpression
of the Arabidopsis formin AFH1 in pollen tubes induces the formation of arrays of
actin cables that project into the cytoplasm from the cell membrane and that its
N-terminal structure targets AFH1 to the cell membrane. Pollen tube elongation is
a polar cell growth process dependent on an active and tightly regulated actin cytoskeleton.
Slight increases in AFH1 stimulate growth, but its overexpression induces tube broadening,
growth depolarization, and growth arrest in transformed pollen tubes. These results
suggest that AFH1-regulated actin polymerization is important for the polar pollen
cell growth process. Moreover, severe membrane deformation was observed in the apical
region of tip-expanded, AFH1-overexpressing pollen tubes in which an abundance of
AFH1-induced membrane-associated actin cables was evident. These observations suggest
that regulated AFH1 activity at the cell surface is important for maintaining tip-focused
cell membrane expansion for the polar extension of pollen tubes. The cell surface-located
group-I formins may play the integrin-analogous role as mediators of external stimuli
to the actin cytoskeleton, and AFH1 could be important for mediating extracellular
signals from female tissues to elicit the proper pollen tube growth response during
pollination.
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