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238 V. Demidchik<br />

Ligand-activated and ROS-activated NSCC can potentially play an important<br />

role in the nutritional K + influx. Demidchik et al. (2004) have demonstrated<br />

that glutamate-activated NSCC in Arabidopsis root epidermis catalyse<br />

significant K + influx currents. High K + permeability of glutamateactivatedNSCCisprobablyresponsibleforaglutamate-induceddepolarisation<br />

of the plasma membrane in Arabidopsis root cells (Dennison and<br />

Spalding 2000). OH•-activated NSCC were shown to mediate large K + and<br />

NH + 4 influx currents (Demidchik et al. 2003). Mechanosensitive NSCC in<br />

Allium cepa bulb epidermis are well permeable to K + (Ding and Pickard<br />

1993) but their role in nutritional K + uptake is difficult to predict owing to<br />

verylimiteddataonthisgroupofchannels.<br />

16.4.2<br />

Calcium and Magnesium<br />

Extracellular Ca 2+ and Mg 2+ concentrations are similar (between 0.1–<br />

2mM)but[Ca 2+ ]cyt (about 100 nM) is dramatically lower than cytosolic<br />

Mg 2+ activity (0.4−2 mM)(Bergmann 1992; Marschner 1995). This results<br />

in very positive ECa (from 87 to 125 mV) and EMg about 0 mV. At negative<br />

membrane potentials, both ions can be passively transported to the cell but<br />

only Ca 2+ can pass a membrane through the channels at positive potentials.<br />

According to Romano et al. (1998) depolarisation-activated K + -permeable<br />

channels in Arabidopsis mesophyll protoplasts can catalyse a significant<br />

Ca 2+ influx. Similar results were found in root protoplasts (Roberts<br />

and Tester 1997; Gilliham et al. 2004). Selectivity series for single Ca 2+ -<br />

permeable outwardly rectifying conductances in the maize root showed that<br />

they were catalysed by NSCC (Roberts and Tester 1997). So, depolarisationactivated<br />

NSCC could catalyse some Ca 2+ influx.<br />

Passive transport of Ca 2+ fornutritionalneedsisakeyquestionofplant<br />

mineral nutrition (Marschner 1995). HACC activate at membrane potentials<br />

more negative than −150 mV (Véry and Davies 2000). When [Ca 2+ ]cyt<br />

increasesHACCactivateatmorepositivevoltagesandthereforeprobably<br />

play a significant role in Ca 2+ uptake in growing tissues (where [Ca 2+ ]cyt is<br />

elevated) (Véry and Davies 2000; Demidchik et al. 2002). DACC have low<br />

amplitude and are only functional in about 35% of protoplasts (Thion et al.<br />

1996). DACC reveal maximal activation at −70 to −80 mV and could be involved<br />

in signalling, having no significant impact on nutritional Ca 2+ influx.<br />

Typically plasma membrane potentials in Arabidopsis root epidermal cells<br />

vary between −101 and −144 mV (Maathuis and Sanders 1993; Demidchik<br />

et al. 2002), thus HACC or DACC cannot be a major Ca 2+ influx pathway<br />

in these conditions. A key role of constitutive Ca 2+ -permeable NSCC<br />

in nutritional Ca 2+ influx has recently been demonstrated in Arabidopsis

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