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13<br />

The Arabidopsis thaliana<br />

Glutamate-like Receptor Family (AtGLR)<br />

Matthew Gilliham, Malcolm Campbell, Christian Dubos,<br />

Dirk Becker, Romola Davenport<br />

Abstract The 20 genes that encode the Arabidopsis thaliana glutamate-like receptor family<br />

(AtGLR) share significant similarity in amino acid coding sequence and predicted secondary<br />

structure with animal ionotropic glutamate receptor (iGluR) subunits. In animals,<br />

iGluR subunits form glutamate-gated non-selective cation channels (NSCCs) catalysing Na +<br />

and/or Ca 2+ influxintocells;inoneiGluRsubfamilyglycinealsoisrequiredasacoagonist.In<br />

Arabidopsis, both glutamate and glycine have been demonstrated to depolarise the plasma<br />

membrane and increase [Ca 2+ ]cyt, and iGluR antagonists blocked these effects. AtGLRs are<br />

therefore predicted to function in an analogous manner to iGluR. Attempts to functionally<br />

characterise AtGLRs in heterologous expression systems have proved inconclusive with no<br />

ligand-gated activity detected. Research into the glutamate receptor-like family has been<br />

hindered by the lack of phenotypes associated with the AtGLR genes but several phenotypes<br />

associated with AtGLR overexpression and knockout have recently given hints as to their<br />

function. AtGLR have been implicated in light and C:N signalling, hypocotyl detiolation,<br />

root growth, abscisic acid (ABA) metabolism, stress responses, and general ion transport.<br />

This review will concentrate on recent developments in the AtGLR field, including the roles<br />

and effects of glutamate and glycine and related metabolites in plant physiology relative to<br />

potential roles for AtGLRs. It will examine progress made toward defining the functions<br />

of particular AtGLRs and will conclude by recommending potentially fruitful avenues of<br />

future research.<br />

13.1<br />

Introduction<br />

Twenty genes in the Arabidopsis thaliana genome encode subunits of<br />

glutamate-likereceptors(AtGLRs) (Lametal.1998;Lacombe2001a).AtGLR<br />

subunits are so named owing to their similarity in amino acid coding sequence,<br />

and predicted secondary structure, to animal ionotropic glutamate<br />

receptor (iGluR) subunits (Lam et al. 1998; Fig. 13.1). Phylogenetic analysis<br />

suggests that the evolution of the AtGLR subunits predates the divergence<br />

of plants and animals (Chiu et al. 1999, 2002). It is believed that their evolutionary<br />

precursor evolved from the insertion of an inverted K + -selective<br />

ion channel into an amino acid binding protein (Wo and Oswald 1995). The<br />

discovery of a functional glutamate-activated K + -selective channel (GluR0)<br />

in the prokaryotic cyanobacteria Synechocystis provides compelling support<br />

for both hypotheses (Chen et al. 1999). To date, the functional roles of<br />

AtGLR subunits have not been clearly defined.<br />

Communication in Plants<br />

F. Baluška, S. Mancuso, D. Volkmann (Eds.)<br />

© Springer-Verlag Berlin Heidelberg 2006

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