A hyperaccumulator is a plant capable of growing in soils with very high concentrations of metals, absorbing these metals through their roots, and concentrating extremely high levels of metals in their tissues. The metals are concentrated at levels that are toxic to closely related species not adapted to growing on the metalliferous soils. Compared to non-hyperaccumulating species, hyperaccumulator roots extract the metal from the soil at a higher rate, transfer it more quickly to their shoots, and store large amounts in leaves and roots. The ability to hyperaccumulate toxic metals compared to related species has been shown to be due to differential gene expression and regulation of the same genes in both plants. Over 500 species of flowering plants have been identified as having the ability to hyperaccumulate metals in their tissues.
Hyperaccumulating plants hold interest for their ability to extract metals from the soils of contaminated sites (phytoremediation) to return the ecosystem to a less toxic state. The plants also hold potential to be used to mine metals from soils with very high concentrations (phytomining) by growing the plants then harvesting them for the metals in their tissues.
Several gene families are involved in the processes of hyperaccumulation including upregulation of absorption and sequestration of heavy metal metals. These hyperaccumulation genes (HA genes) are found in over 450 plant species, including the model organisms Arabidopsis and Brassicaceae. The expression of such genes is used to determine whether a species is capable of hyperaccumulation1. Expression of HA genes provides the plant with capacity to uptake and sequester metals such as As, Co, Fe, Cu, Cd, Pb, Hg, Se, Mn, Zn, Mo and Ni in 100-1000x the concentration found in sister species or populations.
The capacity for hyperaccumulation is dependent on two major factors: environmental exposure and expression of members of the ZIP gene family. Although experiments have shown that the hyperaccumulation is partially dependent on environmental exposure (i.e. only plants exposed to a metal are observed with high concentrations of that metal), hyperaccumulation is ultimately dependent on the presence and upregulation of genes involved with that process. It has been shown that hyperaccumulation capacities can be inherited in Thlaspi caerulescens (Brassicaceae) and others. As there is wide variety among hyperaccumulating species that span across different plant families, it is likely that HA genes were ecotypically selected for. In most hyperaccumulating plants, the main mechanism for metal transport are the proteins coded by genes in the ZIP family, however other families such as the HMA, MATE, YSL and MTP families have also been observed to be involved. The ZIP gene family is a novel, plant-specific gene family that encodes Cd, Mn, Fe and Zn transporters. The ZIP family plays a role in supplying Zn to metalloproteins.
In one study on Arabidopsis, it was found that the metallophyte A. halleri expressed a member of the ZIP family that was not expressed in a non-metallophytic sister species. This gene was an iron regulated transporter (IRT-protein) that encoded several primary transporters involved with cellular uptake of cations above the concentration gradient. When this gene was transformed into yeast, hyperaccumulation was observed. This suggests that overexpression of ZIP family genes that encode cation transporters is a characteristic genetic feature of hyperaccumulation. Another gene family that has been observed ubiquitously in hyperaccumulators are the ZTP and ZNT families. A study on T. caerulescens identified the ZTP family as a plant specific family with high sequence similarity to other zinc transporter4. Both the ZTP and ZNT families, like the ZIP family, are zinc transporters. It has been observed in hyperaccumulating species, that these genes, specifically ZNT1 and ZNT2 alleles are chronically overexpressed.
While the exact mechanism by which these genes facilitate hyperaccumulation is, as of yet, uncharacterized, expression patterns correlate heavily with individual hyperaccumulation capacity and metal exposure, suggests these gene families play a regulatory role. As the presence and expression zinc transporter gene families is highly prevalent in hyperaccumulators, it is likely that the capacity to accumulate a wide range of heavy metals is likely due to an inability of the zinc transporters to discriminate against certain metal ion. The response of the plants to hyperaccumulation of any metal also supports this theory as it has been observed that AhHMHA3 is expressed in hyperaccumulating individuals. AhHMHA3 has been identified to be expressed in response and aid of Zn detoxification. In another study, using metallophytic and non-metallophytic Arabidopsis populations, back crosses indicated pleiotropy between Cd and Zn tolerances. This response suggests that plants are unable to detect specific metals, and that hyperaccumulation is likely a result of an overexpressed Zn transportation system.
The overall effect of these expression patterns has been hypothesized to assist in plant defense systems. In one hypothesis, “the elemental defense hypothesis,” provided by Poschenrieder, it is suggested that the expression of these genes assist in antiherbivory or pathogen defenses by making tissues toxic to organisms attempting to feed on that plant. Another hypothesis, “the joint hypothesis,” provided by Boyd, suggests that expression of these genes assists in systemic defense
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