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232 S.J. Roux, C. Song, C. Jeter<br />

animal cells, although several works have used luciferin–luciferase-based<br />

reporters localized to the ECM to obtain initial data on this critical measurement<br />

(Joseph et al. 2003), and plant scientists are testing this approach also<br />

in plants (Gary Stacey, University of Missouri, personal communication).<br />

Ifandwhenapurinoceptorisidentifiedinplantsandmethodsfor<br />

measuring changes in eATP concentration are perfected, the field of nucleotide<br />

signaling will rapidly expand in significance, as will the field of<br />

ectoapyrase studies. Since virtually all growth-signaling pathways are interlinked,<br />

future breakthroughs on the impact of extracellular nucleotides<br />

on plant growth and development will also inform diverse other fields that<br />

focus on growth control (e.g., hormone physiology, photomorphogenesis,<br />

gravitropism/phototropism, plant reproduction). The field of extracellular<br />

nucleotidesignalinginplantsisinitsinfancy,andweexpectthemost<br />

exciting insights are yet to come.<br />

Acknowledgements. The authors thank Wenqiang Tang, Timothy Butterfield,<br />

X. Wang, and Stephen Stout for data and valuable discussion. Research<br />

of the authors reviewed here was supported by grants from the National<br />

Science Foundation (IBN 0344221) and the National Aeronautics and Space<br />

Administration (NAG2-1586 and NAG10-295).<br />

<strong>References</strong><br />

Arteca, J, Arteca N (1999) A multi-responsive gene encoding 1-aminocyclopropane-1carboxylate<br />

synthase (ACS6) in mature Arabidopsis leaves. Plant Mol Biol 39:209–219<br />

Burnstock G (1972) Purinergic nerves. Pharmacol Rev 24:509–581<br />

Burnstock G (1996) Purinoceptors: ontogeny and phylogeny. Drug Dev Res 39:204–242<br />

Burnstock G, Knight GE (2004) Cellular distribution and functions of P2 receptor subtypes<br />

in different systems. Int Rev Cytol 240:31–51<br />

Clark G, Lee D, Dauwalder M, Roux SJ (2005) Immunolocalization and histochemical evidence<br />

for the association of two different Arabidopsis annexins with secretion during<br />

early seedling growth and development. Planta 220:621–631<br />

Coco S, Calegari F, Pravettoni E, Pozzi D, Taverna E, Rosa P, Matteoli M, Verderio C (2003)<br />

Storage and release of ATP from astrocytes in culture. J Biol Chem 278:1354–1362<br />

CohnJR,UhmT,RamuS,NamY-W,KimD-J,PenmetsaV,WoodTC,DennyRL,YoungND,<br />

Cook, DR, Stacey G (2001) Differential regulation of a family of apyrase genes from<br />

Medicago truncatula. Plant Physiol 125:2104–2119<br />

Demidchik V, Nichols C, Oliynyk M, Dark A, Glover, BJ, Davies JM (2003) Is ATP a signaling<br />

agent in plants? Plant Physiol 133:456–461<br />

Gout E, Bligny R, Douce R (1992) Regulation of intracellular pH values in higher plant<br />

cells. Carbon-13 and phosphorus-31 nuclear magnetic resonance studies. J Biol Chem<br />

267:13903–13909<br />

Jabs T, Dietrich RA, Dangl JL (1996) Initiation of runaway cell death in an Arabidopsis<br />

mutant by extracellular superoxide. Science 273:1853–1856<br />

Jaffe MJ (1973) The role of ATP in mechanically stimulated rapid closure of the Venus’sflytrap.<br />

Plant Physiol 51:17–18

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