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Foldable structures and the natural design of pollen grains
Edited by José N. Onuchic, University of California San Diego, La Jolla, CA, and approved February 25, 2010 (received for review September 29, 2009)

Abstract
Upon release from the anther, pollen grains of angiosperm flowers are exposed to a dry environment and dehydrate. To survive this process, pollen grains possess a variety of physiological and structural adaptations. Perhaps the most striking of these adaptations is the ability of the pollen wall to fold onto itself to prevent further desiccation. Roger P. Wodehouse coined the term harmomegathy for this folding process in recognition of the critical role it plays in the survival of the pollen grain. There is still, however, no quantitative theory that explains how the structure of the pollen wall contributes to harmomegathy. Here we demonstrate that simple geometrical and mechanical principles explain how wall structure guides pollen grains toward distinct folding pathways. We found that the presence of axially elongated apertures of high compliance is critical for achieving a predictable and reversible folding pattern. Moreover, the intricate sculpturing of the wall assists pollen closure by preventing mirror buckling of the surface. These results constitute quantitative structure-function relationships for pollen harmomegathy and provide a framework to elucidate the functional significance of the very diverse pollen morphologies observed in angiosperms.
Footnotes
- 1To whom correspondence should be addressed. E-mail: ekatifori{at}mail.rockefeller.edu.
Author contributions: E.K., S.A., E.C., D.R.N., and J.D. designed research; E.K., S.A., E.C., and J.D. performed research; E.K., E.C., D.R.N., and J.D. contributed new reagents/analytic tools; E.K., S.A., E.C., and J.D. analyzed data; and E.K., D.R.N., and J.D. wrote the paper.
The authors declare no conflict of interest.
This article is a PNAS Direct Submission.
This article contains supporting information online at www.pnas.org/cgi/content/full/0911223107/DCSupplemental.
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