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11 AA transport in plant versus neutrotransmission 165<br />

(Grallath et al. 2005; Rentsch et al. 1996). AtProT1isexpressedinthe<br />

phloem and phloem parenchyma cells, whereas AtProT2 expression is<br />

restricted to the epidermis and cortex cells of roots. AtProT3 finally is only<br />

expressed in the aerial parts of the plant, in the epidermal cells of the leaves.<br />

Induction of the expression of the AtProT upon salt stress implies a role<br />

in stress adaptation or a function in organs of plants that desiccate like<br />

pollen and seeds (Rentsch and Frommer 1996; Schwacke et al. 1999). In<br />

mangroves, where salt concentrations are permanently high, the transport<br />

of compatible solutes is thought to be an important factor for adaptation<br />

to the environment (Waditee et al. 2002).<br />

In plants nothing is known about the compartimentation of AA or the<br />

transport steps into the vacuole. Some plant ATF1 members might fulfil<br />

a function in import or export of solutes in vesicles and vacuoles. A similar<br />

mechanism like the concentration of GABA in synaptic vesicles via VGAT<br />

(Fig. 11.1) is possible for the related uncharacterized plant proteins in the<br />

ATF family.<br />

The plant members of the ATF family investigated so far all function as<br />

H + -coupled systems, probably playing a role in accumulating AA within<br />

the plant cell. Subcellular localization studies could help to identify the<br />

transporters involved in the compartimentation of AA into the vacuole.<br />

11.4<br />

Conclusions and Future Prospects<br />

Taken together, plant AA transporters can be grouped in three superfamilies<br />

with homologs from the animal kingdom (Lalonde et al. 2004; Wipf<br />

et al. 2002): (1) the ATF1 superfamily (SLC32, SLC36 and SLC38 and Arabidopsis<br />

ATF1 members) (Fig. 11.3); (2) the APC superfamily (SLC7, AtCATs<br />

and AtLATs) (Fig. 11.2); (3) AA transporters within the major facilitator<br />

superfamily (MFS) (SLC17, homologs of unknown function in Arabidopsis<br />

(Wipf et al. 2002).<br />

As described, several export steps are required for AA distribution in<br />

plants (Fig. 11.3): phloem-to-xylem transfer, release of AA into the apoplast<br />

of the leaf mesophyll as the first step for phloem loading, unloading in sink<br />

organs, supply of symplasmic isolated cells, e.g., growing embryo, guard<br />

cells and pollen. Multiple possibilities exist, including vesicular transport<br />

or carrier-mediated transport, but until now no exporter has been identified<br />

in plants. Vesicular export is well established for AA and analogs in<br />

mammalian nerve cells. Carriers for the active uptake of AA into secretory<br />

vesicles have been identified (Fig. 11.1) and homologous plant genes exist<br />

(Fig. 11.3). Furthermore, certain mammalian AA transporters are implicatedincellularexportattheplasmamembrane,especiallyfromintestinal

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