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16 Nonselective cation channels 239<br />

root epidermis (Demidchik et al. 2002). These channels were clearly dominant<br />

in Ca 2+ influx currents at resting membrane potentials. 44 Ca 2+ flux<br />

measurements showed that the Ca 2+ uptake rate was not sensitive to verapamil<br />

(HACC blocker) but was inhibited by Gd 3+ (nonspecific blocker<br />

of cation channels). In addition, basal [Ca 2+ ]cyt revealed linear voltage<br />

dependence (NSCC have linear current–voltage relationships) that again<br />

implicates NSCC in Ca 2+ uptake. Constitutive Ca 2+ -permeable NSCC were<br />

permeable to Mg 2+ , and could therefore participate in nutritional Mg 2+<br />

influx into the root.<br />

Glutamate-activated Ca 2+ influx channels have recently been identified in<br />

Arabidopsis root epidermal protoplasts (Demidchik et al. 2004). Glutamateactivated<br />

Ca 2+ conductance appeared as “spiky” currents in about 20%<br />

of protoplasts. Protoplasts isolated from roots of aequorin-transformed<br />

Arabidopsis plants demonstrated steady-state (measured over 2 h) elevation<br />

of [Ca 2+ ]cyt in response to 0.2−2 mM glutamate, suggesting a significant role<br />

of glutamate-activated channels in the regulation of the basal cytosolic Ca 2+<br />

activity. Apoplastic glutamate concentration is between 0.3 and 1.3 mM, as<br />

measured in a range of tissues and species (Lohaus et al. 1995; Lohaus and<br />

Heldt 1997). Therefore, it is possible that apoplastic glutamate, and maybe<br />

glycine too (Dubos et al. 2003), functions as a “permanent” activator of<br />

Ca 2+ influx into the plant cells.<br />

Purine-induced Ca 2+ influx has recently been discovered in Arabidopsis<br />

roots (Demidchik et al. 2003). Application of ATP and its nonhydrolysable<br />

derivativesresultedinamanifoldtransientincreasein[Ca 2+ ]cyt fully recovered<br />

within 10−15 min after the application of purines. This suggests<br />

that, in contrast to glutamate, purines do not cause steady-state changes in<br />

Ca 2+ influx and do not affect nutritional Ca 2+ uptake.<br />

Arabidopsis root and guard cell ROS-activated NSCC catalysed significant<br />

Ca 2+ and Mg 2+ influx that could play a role in nutritional uptake of<br />

these cations (Pei et al. 2000; Demidchik et al. 2003; Foreman et al. 2003).<br />

16.4.3<br />

Microelements and Trace Elements<br />

It is generally accepted that Na + influx is catalysed by root epidermal<br />

NSCC (reviewed by Demidchik et al. 2002; Tester and Davenport 2003).<br />

Constitutive Na + -permeable NSCC have been demonstrated in a range of<br />

tissues and species (Stoeckel and Takeda 1989; Elzenga and van Volkenburgh<br />

1994; White and Lemtiri-Chlieh 1995; Tyerman et al. 1997; Roberts<br />

and Tester 1997; Amtmann et al. 1997; Véry et al. 1998; Demidchik and<br />

Tester 2002). In the majority of preparations these channels were weakly<br />

selective for monovalent cations, voltage-insensitive (or slightly voltage

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