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172 A. Fait, A. Yellin, H. Fromm<br />

Fig.12.1. Overview of the γ-aminobutyric acid (GABA) shunt and relevant metabolic<br />

branches in plants and other organisms. GABA is metabolized via a short pathway known<br />

as the GABA shunt, which bypasses two steps of the tricarboxylic acid cycle (TCA), and<br />

is composed of three enzymes: the cytosolic-localized Ca 2+ -calmodulin (CaM)-regulated<br />

glutamate decarboxylase (GAD), and the mitochondrial-localized GABA trans-aminase<br />

(GABA-T) and succinic semialdehyde dehydrogenase (SSADH). The SSADH substrate<br />

succinic semialdehyde (SSA) isalsocatabolizedtoγ-hydroxybutyrate (GHB) byGHBdehydrogenase<br />

(GHBDH), which is cytosolic in mammals. GHB catabolism occurs in mammals<br />

either in the reverse direction by GHBDH/semialdehyde reductase (see text) or by<br />

the mitochondrial-localized L-glucuronate reductase (GR; EC 1.1.1.19). In plants, GHB<br />

catabolism has not been characterized. Chemical structures are given for GABA-shunt<br />

substrates, intermediates, and products. Unknown functions and transport systems are<br />

depicted as dotted arrows and question marks. Glu glutamate, α-KG α-ketoglutarate, GABA-<br />

TK/P α-ketoglutaric acid/pyruvate-dependent GABA-T<br />

in plants. We also address the occurrence, in plants, of γ-hydroxybutyrate<br />

(GHB), a by-product of the GABA shunt (Fig. 12.1) and a neurotransmitter,<br />

which was only recently discovered in plants (Allan et al. 2003; Breitkreuz et<br />

al. 2003; Fait et al. 2005). For further details on the different roles of GABA<br />

in plants the readers are referred to two recent reviews (Bouché et al. 2003;<br />

Bouché and Fromm 2004) and to earlier reviews by Shelp et al. (1999) and<br />

Kinnersley and Turano (2000).

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