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JXB Advance Access published online on June 22, 2006
Journal of Experimental Botany, doi:10.1093/jxb/erj186
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Published by Oxford University Press [2006] on behalf of the Society for Experimental Biology. This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/2.0/uk/) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.
Received November 4, 2005
Accepted March 13, 2006

RESEARCH PAPER

Stomatal lock-open, a consequence of epidermal cell death, follows transient suppression of stomatal opening in barley attacked by Blumeria graminis

Elena Prats 1 *, Alan P. Gay 1, Luis A. J. Mur 2, Barry J. Thomas 1, and Timothy L. W. Carver 1

1 Institute of Grassland and Environmental Research, Aberystwyth, Ceredigion SY23 3EB, UK
2 University of Wales Aberystwyth, Institute of Biological Sciences, Aberystwyth, Ceredigion SY23 2DA, UK

* To whom correspondence should be addressed.
Elena Prats, E-mail: bb2prpee{at}uco.es


  Abstract

Blumeria graminis f.sp. hordei (Bgh) attack disrupted stomatal behaviour, and hence leaf water conductance (gl), in barley genotypes Pallas and Risø-S (susceptible), P01 (with Mla1 conditioning a hypersensitive response; HR), and P22 and Risø-R (with mlo5 conditioning papilla-based penetration resistance). Inoculation caused some stomatal closure well before the fungus attempted infection. Coinciding with epidermal cell penetration, stomatal opening in light was also impeded, although stomata of susceptible and mlo5 lines remained largely able to close in darkness. Following infection, in susceptible lines stomata closed in darkness but opening in light was persistently impeded. In Risø-R, stomata recovered nearly complete function by ~30 h after inoculation, i.e. after penetration resistance was accomplished. In P01, stomata became locked open and unable to close in darkness shortly after epidermal cells died due to HR. In the P22 background, mlo5 penetration resistance was often followed by consequential death of attacked cells, and here too stomata became locked open, but not until ~24 h after pathogen attack had ceased. The influence of epidermal cell death was localized, and only affected stomata within one or two cells distance. These stomata were unable to close not only in darkness but also after application of abscisic acid and in wilted leaves suffering drought. Thus, resistance to Bgh based on HR or associated with cell death may have previously unsuspected negative consequences for the physiological health of apparently ‘disease-free’ plants. The results are discussed in relation to the control of stomatal aperture in barley by epidermal cells.

Keywords: Barley; Blumeria; hypersensitive response; papilla; powdery mildew; stoma; stomatal conductance; stomatal lock-up.
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