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Page 2 Plant-Bacteria Interactions Edited by Iqbal Ahmad, John ...

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15.5 Microbial Enhancement of Metal Ion Removal Capacity of Water Hyacinthj295<br />

therefore, be expected to modify the microbial ecology of rhizosphere [19–23,117].<br />

This can be attained <strong>by</strong> various methods such as addition of organic amendments,<br />

microbial inoculation, manipulation of plant genotypes and exposure to environmental<br />

stresses such as heavy metal contamination.<br />

15.5.2<br />

Mechanisms of Metal Ion Removal <strong>by</strong> <strong>Plant</strong> Roots<br />

Possible mechanisms of toxic metal ion removal <strong>by</strong> plant roots include extracellular<br />

precipitation, cell wall precipitation and adsorption, intracellular uptake followed <strong>by</strong><br />

cytoplasm compartmentalization or vacuolar deposition (Figure 15.1) [130]. The<br />

metal ion uptake and detoxification mechanisms will vary with different plant<br />

species. Most metal ions enter plant cells <strong>by</strong> an energy-dependent, saturable process<br />

through specific or generic metal ion carriers or channels [42]. Toxic metal ions may<br />

employ the same mechanisms that are responsible for the uptake of essential ions.<br />

Paganetto et al. [118] used the patch-clamp techniques to study the transport<br />

properties of vacuolar ion channels from the roots of water hyacinth, Eichhornia<br />

crassipes (Mart. Solms, Pontederiaceae). The vacuolar currents for the transport of<br />

Ni 2+ and Zn 2+ were found to be different from other common ions such as Na + ,Ca 2+<br />

and NH 4 + . However, membrane transport systems such as aqueous pores, ion efflux<br />

pumps, ion selective channels and proton–anion contraports may fail to discriminate<br />

among different metal ions that have similar ionic radii and the same ionic<br />

charge [42,87].<br />

Metabolically important cations from the external solution can accumulate in a<br />

nonmetabolic step (Figure 15.3) [87]. Entry and association of metal ions with plant<br />

cells may occur <strong>by</strong> a number of physical processes, including diffusion, ion exchange,<br />

mass flow and adsorption. The cation-exchange occurs within the free<br />

space of the roots where ions can penetrate without passage through a living<br />

membrane. The exchangeable cations are electrostatically bound to negatively<br />

charged functional groups, probably the free carboxyl groups of pectic cell wall<br />

matrix substances [119–121]. Precipitation and exchangeable sorption remove metal<br />

ions <strong>by</strong> forming insoluble compounds in the free space (Figure 15.3) [87]. Some<br />

natural hyperaccumulators extend and proliferate their roots positively into patches<br />

of high metal ion availability. In contrast, nonaccumulators actively avoid these<br />

areas; this is one of the mechanisms <strong>by</strong> which hyperaccumulators absorb more<br />

metal ions when grown in the same soil [122]. Also, phytochelatins play an important<br />

role in the accumulation and detoxification of excess metal ions such as Cd 2+ in plant<br />

cells [58,87,123–127]. This mechanism further assists phytofiltration technology, as<br />

it increases the specificity of metal ions to binding domains in the plant root. Some<br />

species of plants also have multiple binding capacities for metal ions owing to<br />

induced formation of phytochelatins <strong>by</strong> different metal ions such as Cu 2+ ,Zn 2+ ,<br />

Pb 2+ and Cd 2+ [87]. Indeed, the application of chelates has been shown to induce<br />

substrate accumulation of metals such as Pb 2+ ,U 2+ and Au 3+ in the shoots of<br />

nonhyperaccumulators <strong>by</strong> increasing metal solubility and root-to-shoot translocation<br />

[122].

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