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Hyperaccumulator

A hyperaccumulator is a plant species capable of absorbing and concentrating unusually high levels of heavy metals or metalloids, such as nickel, zinc, cadmium, or arsenic, in its living tissues—particularly the aboveground biomass—often reaching concentrations hundreds or thousands of times greater than those in typical plants, without exhibiting phytotoxic effects.^[1] These plants thrive on metalliferous soils and represent a small fraction of vascular plant species, with over 700 known examples across diverse families as of 2024, enabling them to tolerate and sequester contaminants that would harm most vegetation.^[1]^[2] The concept of hyperaccumulation was first described in 1976 by Jaffré et al., who coined the term for nickel-accumulating species like Pycnandra acuminata (formerly Sebertia acuminata) in New Caledonia, highlighting plants that accumulate over 10% dry weight of nickel in their latex.^[3] Standardized thresholds define hyperaccumulation based on foliar concentrations in natural habitats: greater than 100 μg g⁻¹ for cadmium, thallium, and selenium; 300 μg g⁻¹ for cobalt, copper, and chromium; 1,000 μg g⁻¹ for nickel, arsenic, lead, and rare earth elements; 3,000 μg g⁻¹ for zinc; and 10,000 μg g⁻¹ for manganese, all on a dry weight basis.^[1] These thresholds distinguish hyperaccumulators from regular metal-tolerant plants, emphasizing their role in evolutionary adaptations to serpentine or contaminated soils.^[4] Hyperaccumulators are pivotal in environmental biotechnology, particularly phytoremediation, where they facilitate the extraction of metals from polluted sites through phytoextraction—absorbing contaminants into harvestable biomass for safe disposal or recycling.^[4] Notable examples include Noccaea caerulescens (formerly Thlaspi caerulescens), which hyperaccumulates zinc and cadmium up to 5.4% and 0.3% dry weight, respectively; Pteris vittata, the Chinese brake fern that accumulates arsenic to 2.3%; and Berkheya coddii, a nickel hyperaccumulator reaching 7.6% in leaves.^[1] Beyond cleanup, these plants support phytomining for valuable metals and serve as models for studying metal homeostasis, though many face threats from habitat destruction due to mining activities.^[1]

Definition and Properties

Definition

A hyperaccumulator is defined as a plant capable of accumulating extraordinarily high concentrations of metals or metalloids in its harvestable biomass, often exceeding typical soil levels by orders of magnitude. This accumulation is measured in the dry weight of above-ground tissues, with standard thresholds including greater than 100 μg g⁻¹ (0.01%) for cadmium (Cd), thallium (Tl), and selenium (Se); 300 μg g⁻¹ (0.03%) for cobalt (Co), copper (Cu), and chromium (Cr); 1,000 μg g⁻¹ (0.1%) for nickel (Ni), arsenic (As), lead (Pb), and rare earth elements; 3,000 μg g⁻¹ (0.3%) for zinc (Zn); and 10,000 μg g⁻¹ (1%) for manganese (Mn).^[1] These thresholds ensure that hyperaccumulation represents not just elevated uptake but a biologically significant phenomenon distinct from routine plant metal absorption.^[5] Hyperaccumulators are classified as obligate if restricted to metalliferous soils or facultative if they can grow on normal soils but hyperaccumulate under metal stress.^[1] The term "hyperaccumulator" was first coined in 1976 by Jaffré et al. in their seminal study of nickel accumulation in the New Caledonian tree Sebertia acuminata (now Pycnandra acuminata), where latex nickel concentrations reached up to 25% of dry weight—levels hundreds of times higher than in surrounding soils.^[6] This discovery, building on earlier observations of nickel-tolerant flora in ultramafic soils of New Caledonia, highlighted hyperaccumulation as a rare adaptation in metal-rich environments and spurred global research into such species.^[3] Hyperaccumulation differs from general bioaccumulation in that it is strictly threshold-based, requiring not only high internal metal concentrations but also robust tolerance to prevent toxicity, rather than mere sequestration as a defense mechanism.^[4] Classification criteria emphasize foliar (leaf) concentrations achieved under natural or low-contamination conditions, ensuring the trait reflects inherent physiological capability rather than induced stress responses, as outlined in foundational reviews by Baker et al. and subsequent refinements.^[1]

Key Properties

Hyperaccumulators exhibit exceptional metal tolerance, enabling them to thrive in soils contaminated with high concentrations of heavy metals or metalloids without significant reductions in growth or yield, often through the formation of metal complexes with organic acids such as citrate or histidine that facilitate detoxification and transport.^[7] For instance, species like Noccaea caerulescens can tolerate zinc levels exceeding 10,000 mg/kg in their shoots while maintaining normal physiological functions.^[8] These plants typically demonstrate rapid growth rates and substantial biomass production, which are crucial for their practical applications, with some species achieving yields of 10-20 tons per hectare per year under optimal conditions.^[9] Examples include Berkheya coddii, a nickel hyperaccumulator that produces up to 20 metric tons of biomass per hectare annually on ultramafic soils, enhancing their efficiency in metal extraction processes. A defining trait is their ability to compartmentalize accumulated elements, primarily through vacuolar sequestration in leaf tissues, which isolates metals from sensitive cytoplasmic components and prevents toxicity.^[10] In Arabidopsis halleri, for example, zinc is predominantly stored in leaf vacuoles via transporters like MTP1, allowing the plant to hyperaccumulate without cellular damage.^[11] Ecologically, hyperaccumulators play roles in metalliferous environments, including potential allelopathic effects where leaf litter enriches surrounding soil with metals, inhibiting the growth of competing non-tolerant species.^[12] This trait, observed in selenium hyperaccumulators like Astragalus bisulcatus, may provide a competitive advantage and reflects evolutionary adaptations to naturally metal-rich soils such as serpentine outcrops. These properties are quantified using advanced analytical techniques, such as inductively coupled plasma mass spectrometry (ICP-MS), which provides precise measurements of elemental concentrations in plant tissues to confirm hyperaccumulation thresholds.

Mechanisms of Hyperaccumulation

Physiological Basis

Hyperaccumulators exhibit specialized physiological adaptations that facilitate the efficient uptake of heavy metals from the soil into their roots. Metal ions primarily enter root cells through apoplastic pathways, moving via cell walls and intercellular spaces, or symplastic pathways, crossing plasma membranes into the cytoplasm. These processes are enhanced in hyperaccumulators by modifications in the rhizosphere, such as the secretion of protons that lower local soil pH, thereby increasing metal solubility and bioavailability. Additionally, associations with mycorrhizal fungi, particularly arbuscular mycorrhizae, extend the root system's absorptive surface and can mobilize metals through fungal hyphae, promoting greater uptake without necessarily increasing toxicity to the host plant.^[13]^[14]^[15] Once absorbed, metals are loaded into the xylem for long-distance transport to aboveground tissues, driven primarily by transpiration-induced mass flow. This efficient translocation is supported by low-molecular-weight ligands, such as nicotianamine, which chelate metals like zinc and nickel, maintaining their solubility and preventing precipitation during ascent through the vascular system. In hyperaccumulators, minimal sequestration in root vacuoles ensures that a higher proportion of absorbed metals is directed toward the xylem rather than being retained in the roots, enabling rapid delivery to shoots and leaves.^[13]^[14]^[15] To tolerate high internal metal concentrations, hyperaccumulators employ detoxification strategies that sequester ions away from sensitive cellular compartments. Metals are bound by peptides such as metallothioneins and phytochelatins in the cytosol, which neutralize toxicity and facilitate their transport into vacuoles for long-term storage. These mechanisms also counteract oxidative stress induced by reactive oxygen species (ROS), generated as byproducts of metal-catalyzed reactions, through enhanced antioxidant enzyme activity and glutathione-mediated pathways. Vacuolar compartmentalization is an energy-intensive process, relying on ATP-dependent proton pumps to create electrochemical gradients that drive metal influx via secondary transporters.^[13]^[14]^[15] Experimental evidence from radiotracer studies, such as those using ^{65}Zn in Thlaspi caerulescens, demonstrates the preferential routing of metals to aboveground tissues in hyperaccumulators. In these experiments, the hyperaccumulator showed a 4.5-fold higher influx into root symplasm compared to non-accumulators, with only 5% of absorbed Zn sequestered in root vacuoles versus 12% in controls, resulting in sixfold greater translocation to shoots over 46 hours. Such isotope tracing confirms the physiological efficiency of these pathways, highlighting reduced root retention and enhanced xylem throughput as key to hyperaccumulation.^[16]^[13]

Genetic Basis

The genetic basis of hyperaccumulation in plants involves several key gene families that facilitate the uptake, transport, and sequestration of heavy metals. Prominent among these are the heavy metal ATPases (HMAs), which drive the efflux of metals like zinc (Zn) and cadmium (Cd) into the xylem for root-to-shoot translocation. In the Zn/Cd hyperaccumulator Arabidopsis halleri, the HMA4 gene is triplicated, with cis-regulatory changes enhancing its expression in roots and shoots compared to non-accumulating relatives like A. thaliana, enabling high metal translocation efficiency. Similarly, the ZIP family of transporters, including ZIP4, ZIP6, ZIP9, and ZIP10, is upregulated in hyperaccumulators such as A. halleri and Noccaea caerulescens (formerly Thlaspi caerulescens), promoting influx of Zn and other divalent metals at the plasma membrane of root cells.^[17] These gene families underscore the molecular adaptations that distinguish hyperaccumulators from tolerant but non-accumulating plants. Evolutionary origins of hyperaccumulation traits trace back to gene duplications and polymorphisms selected in metalliferous environments. The triplication of HMA4 in A. halleri arose from segmental duplications, with subsequent mutations in promoter regions driving constitutive high expression, a pattern absent in populations from non-metal-rich soils. In A. halleri, selective sweeps around HMA4 loci indicate local adaptation through standing genetic variation rather than de novo mutations. In N. caerulescens, variation in HMA4 copy number differentiates populations from calamine (Zn/Cd-rich) versus non-calamine soils.^[18] These evolutionary events highlight how hyperaccumulation likely evolved independently multiple times, often via duplication of ancient metal transport genes present across Plantae.^[19] Quantitative trait loci (QTL) mapping studies reveal that hyperaccumulation is under polygenic control, involving multiple genomic regions rather than single major genes. In A. halleri, QTL analysis of Zn accumulation and tolerance in interspecific crosses with A. lyrata identified three major QTLs for Zn hypertolerance, one of which colocalizes with HMA4, with the QTLs together explaining up to 36% of phenotypic variance, while others contribute to root/shoot partitioning. Similar mapping in N. caerulescens populations confirms polygenic inheritance, with QTLs for Cd and Zn uptake distributed across chromosomes, interacting with environmental factors like soil metal levels to modulate trait expression.^[20] This polygenic architecture allows fine-tuned responses, enhancing fitness on heterogeneous metalliferous soils. Epigenetic modifications, particularly DNA methylation, play a crucial role in regulating gene expression during metal exposure in hyperaccumulators. In N. caerulescens, hypermethylation of cytosine residues in promoter regions of stress-responsive genes occurs upon nickel (Ni) exposure, preserving genomic stability by silencing transposable elements and preventing mutagenesis from reactive oxygen species induced by metals.^[21] Similarly, Cd exposure in N. caerulescens leads to increased global DNA methylation levels, correlating with upregulated expression of metal transporters like HMA4 through demethylation of specific loci, thus facilitating adaptive hyperaccumulation without permanent genetic changes.^[22] These reversible modifications enable rapid, heritable responses to fluctuating metal stresses across generations. Comparative genomics across hyperaccumulator species reveals conserved pathways despite phylogenetic distance, emphasizing shared evolutionary pressures. Transcriptome comparisons between N. caerulescens and its non-accumulating relative Arabidopsis thaliana show over 2,000 differentially expressed genes in N. caerulescens, with enriched pathways for metal homeostasis including HMA and ZIP families, achieving 88% nucleotide identity in coding regions. In unrelated hyperaccumulators like A. halleri and N. caerulescens, both from the Brassicaceae family, orthologous genes in Zn/Cd transport pathways exhibit similar upregulation patterns, suggesting convergence on core regulatory networks for xylem loading and chelation, independent of speciation events.^[23] This conservation facilitates cross-species insights into engineering tolerance traits.

Molecular Transporters

In hyperaccumulating plants, plasma membrane transporters from the ZIP (ZRT/IRT-like protein) family play a crucial role in the influx of essential and non-essential metals from the soil into root cells. These transporters facilitate the uptake of divalent cations such as zinc (Zn²⁺), iron (Fe²⁺), manganese (Mn²⁺), and cadmium (Cd²⁺) by mimicking nutrient ions, with the ZIP subfamily member IRT1 (iron-regulated transporter 1) particularly noted for its adaptation to transport Cd²⁺ alongside Fe²⁺ and Mn²⁺ in hyperaccumulators like Thlaspi caerulescens (now Noccaea caerulescens).^[24]^[25] Overexpression of IRT1 in plants leads to elevated accumulation of Cd²⁺ and Zn²⁺, underscoring its efficiency in metal entry under high soil concentrations typical of contaminated environments.^[24] Other ZIP members, such as ZIP2 and ZIP3, similarly contribute to Cd²⁺ influx, enabling hyperaccumulators to achieve rapid root uptake without immediate toxicity.^[26] Tonoplast transporters, including the metal tolerance proteins (MTPs) from the cation diffusion facilitator (CDF) family, are essential for vacuolar sequestration of excess metals within cells, thereby preventing cytosolic overload and oxidative damage. In Zn/Cd hyperaccumulators like Arabidopsis halleri and N. caerulescens, MTP1 localizes to the tonoplast and drives the accumulation of Zn²⁺ and Cd²⁺ into vacuoles, enhancing metal tolerance by compartmentalizing up to 50% of cellular metal content.^[27]^[28] This sequestration mechanism is particularly pronounced in leaf mesophyll cells, where MTP3 and MTP8 isoforms further support vacuolar storage of Mn²⁺ and Zn²⁺, maintaining homeostasis during hyperaccumulation.^[29] By isolating metals in the vacuole, MTPs allow hyperaccumulators to tolerate concentrations exceeding 1000 µg/g dry weight without disrupting enzymatic functions.^[30] Efflux pumps from the P1B-ATPase family, such as heavy metal ATPases (HMAs), actively drive metal export across membranes, facilitating long-distance transport from roots to shoots in hyperaccumulators. While HMA3 primarily sequesters Cd²⁺, Zn²⁺, and Pb²⁺ in root vacuoles to modulate availability, in species like Sedum plumbizincicola, its shoot-localized activity supports hypertolerance by preventing free cytosolic Cd²⁺ buildup during translocation.^[31]^[32] Related members like HMA4, however, are key for xylem loading and root-to-shoot export of Zn²⁺ and Cd²⁺, with upregulated expression in hyperaccumulators such as N. caerulescens enabling shoot accumulation levels over 10,000 µg/g.^[33] These ATP-driven pumps use energy from ATP hydrolysis to extrude metals against concentration gradients, ensuring efficient aerial hyperaccumulation.^[34] The activity of these transporters is tightly regulated by transcription factors responsive to metal signaling, including bZIP and WRKY families, which coordinate gene expression under stress. bZIP factors like bZIP19 and bZIP23 bind to zinc deficiency-responsive elements, upregulating ZIP transporters in hyperaccumulators to enhance metal influx during exposure.^[35] WRKY transcription factors, such as those in A. halleri, activate HMA and MTP genes via MAPK cascades, promoting sequestration and transport in response to Cd²⁺ or Zn²⁺ signals, thereby fine-tuning hyperaccumulation without toxicity.^[36]^[37] This regulatory network ensures adaptive expression, with bZIP-WRKY interactions amplifying transporter levels by 5- to 10-fold under metal excess.^[38] Structural studies of these transporters reveal conserved metal-binding motifs that underpin their specificity and function, often elucidated through homology modeling and mutagenesis due to limited plant-specific crystallography. ZIP proteins feature variable N-terminal regions with histidine-rich motifs that coordinate Zn²⁺ and Cd²⁺, facilitating selective influx.^[39] P1B-ATPases like HMA3/4 contain signature CPx (cysteine-proline-cysteine) motifs in their transmembrane domains, which form intramembrane metal-binding sites essential for Cd²⁺ and Zn²⁺ transport, as confirmed by site-directed mutagenesis showing reduced activity upon mutation.^[33] MTPs exhibit six transmembrane helices with aspartate- and histidine-coordinated cation efflux domains, enabling vacuolar pumping while avoiding non-specific binding.^[40] These motifs highlight evolutionary adaptations in hyperaccumulators for high-affinity metal handling.

Applications

Phytoremediation

Phytoremediation employs hyperaccumulator plants to decontaminate soils polluted with heavy metals by exploiting their exceptional capacity to uptake and sequester contaminants into harvestable biomass. The process begins with selecting and planting suitable hyperaccumulators on contaminated sites, where they grow over one or more seasons, absorbing metals through their roots and translocating them to aboveground tissues. Once mature, the plants are harvested, and the metal-laden biomass is removed and disposed of through methods such as incineration, smelting, or safe landfilling, thereby reducing soil metal concentrations over repeated cycles. This in situ approach minimizes ecosystem disruption while progressively restoring site usability.^[41] Compared to conventional remediation techniques like soil excavation and disposal, phytoremediation offers significant advantages in cost-effectiveness and environmental compatibility. Traditional excavation can cost $80–150 per cubic meter due to equipment, labor, and off-site disposal requirements, whereas phytoremediation typically ranges from $0.05–1 per cubic meter treated, primarily involving seed or seedling costs, minimal maintenance, and harvesting.^[42]^[43] Additionally, it preserves soil structure, prevents erosion, and enhances biodiversity without generating large waste volumes or requiring heavy machinery. Field applications, such as the use of Alyssum bertolonii for nickel extraction from serpentine soils in Italy since the mid-1990s, have demonstrated practical success, with trials showing up to 30% reduction in soil nickel levels after multiple harvests on ultramafic sites.^[44] Despite these benefits, phytoremediation faces limitations that can hinder its efficacy, including the inherently slow growth rates of many hyperaccumulators, which prolong remediation timelines—especially in colder climates where reduced metabolic activity further delays biomass production. Improper management of harvested biomass also poses risks, such as metal leaching into groundwater if disposal is inadequate, potentially exacerbating contamination. To address bioavailability constraints and accelerate metal uptake, assisted phytoremediation techniques are often integrated, such as applying chelators like EDTA to soil, which complexes metals and enhances their solubility for root absorption; studies have shown EDTA can increase cadmium uptake by up to several-fold in plants like Brassica juncea, though its persistence raises secondary environmental concerns.^[45]^[46]^[47]

Biofortification and Agriculture

Hyperaccumulators have been explored for biofortification of staple crops by transferring traits that enhance uptake and accumulation of essential micronutrients such as zinc (Zn) and selenium (Se). Plants like Arabidopsis halleri and Noccaea caerulescens hyperaccumulate Zn through elevated expression of metal transporter genes, including ZIP family members and HMA4, which facilitate root-to-shoot translocation.^[48] Genetic introgression of such traits into crops like wheat has been achieved by incorporating the Gpc-B1 locus from wild relatives, resulting in up to 10-15% higher grain Zn concentrations without yield penalties.^[48] For Se, hyperaccumulators such as Astragalus bisulcatus and Stanleya pinnata inspire transgenic approaches, where overexpression of genes like selenocysteine methyltransferase in model plants increases Se accumulation in edible tissues by 2-5 fold, improving bioavailability for human consumption.^[49] These strategies aim to address micronutrient deficiencies in populations reliant on staples like rice and wheat, with field trials showing enhanced nutritional profiles in biofortified varieties.^[48] In agricultural settings, hyperaccumulators contribute to revegetation of mine tailings by stabilizing degraded soils and mitigating erosion risks. Tolerant species, including some hyperaccumulators like Atriplex spp., establish root systems that bind loose tailings, reducing wind and water erosion in arid environments through rhizosphere precipitation of metals and improved soil aggregation.^[50] This approach supports land rehabilitation for future farming by preventing contaminant dispersal and enhancing soil structure, with greenhouse studies demonstrating effective vegetation cover when combined with organic amendments.^[50] Such phytostabilization is particularly valuable in semiarid regions, where it lowers reclamation costs to $0.40-26 per cubic meter compared to conventional methods.^[50] Despite these benefits, the use of hyperaccumulator biomass in agriculture poses risks due to elevated metal concentrations, necessitating strict regulations. Biomass from plants like Reynoutria japonica grown on contaminated sites can accumulate cadmium (Cd) levels approaching or exceeding EU maximum limits for animal feed, such as 1 mg/kg for complete feed.^[51] EU Regulation (EC) No 1881/2006 and Directive 2002/32/EC set thresholds for heavy metals like lead (3 mg/kg), mercury (0.1 mg/kg), and arsenic (2 mg/kg) in feed materials to protect animal health and prevent bioaccumulation in the food chain.^[52] ^[53] Improper use of such biomass as fodder could lead to toxicity, prompting guidelines that prohibit its direct application without processing to remove contaminants.^[52] Research advances since 2015 have leveraged CRISPR-Cas9 to incorporate hyperaccumulation-like traits for iron (Fe) enrichment in rice, enhancing crop nutrition. Editing of the OsNRAMP7 gene, a metal transporter analogous to those in hyperaccumulators, has increased Fe accumulation in polished rice grains by 20-50% in edited lines of varieties like TBR225, without affecting yield or introducing toxicity.^[54] These modifications target root uptake and phloem loading mechanisms, drawing from hyperaccumulator physiology to improve Fe bioavailability in Fe-deficient diets.^[55] Similar edits to OsNAS2 promoters in IR64 rice have boosted Fe translocation, achieving 1.5-2 times higher concentrations in endosperm since initial demonstrations in 2016.^[56] Economic assessments highlight the potential market value of biofortified grains from such applications in nutrient-deficient regions. The global biofortification market, driven by micronutrient-enriched staples, was valued at $110.2 million in 2022 and is projected to reach $221.5 million by 2031, with high demand in Asia and Africa where 2 billion people face hidden hunger.^[57] Programs like those from HarvestPlus estimate that scaling Zn- and Fe-biofortified crops could yield $17 in health benefits per $1 invested, particularly in low-income areas with staple-dependent diets.^[58] These projections underscore the role of hyperaccumulator-derived traits in sustainable agriculture, though commercialization requires addressing regulatory hurdles for genetically edited varieties.^[57] As of 2025, ongoing field trials in Asia have integrated microbial-assisted phytoremediation using hyperaccumulators, enhancing metal uptake efficiency.^[59]

Examples

Notable Plant Species

Noccaea caerulescens (formerly Thlaspi caerulescens), commonly known as alpine penny-cress, is a prominent zinc hyperaccumulator native to metalliferous calamine soils across Europe, where it thrives in environments enriched with zinc and lead. This perennial herb from the Brassicaceae family can accumulate up to 54,000 μg/g (5.4% dry weight) of zinc in its leaves, enabling it to tolerate and extract high concentrations from contaminated substrates without exhibiting toxicity symptoms.^[60]^[61] Noccaea goesingensis, a metallophyte endemic to Austrian metalliferous regions such as serpentine soils, exemplifies cadmium hyperaccumulation, reaching levels up to 1,000 μg/g in its tissues. This Brassicaceae species is adapted to heavy metal-rich habitats in central Europe, where it selectively uptakes cadmium alongside other metals like nickel.^[62]^[63] Pteris vittata, or Chinese brake fern, serves as a key arsenic hyperaccumulator, with fronds capable of concentrating up to 22,600 μg/g of arsenic, far exceeding typical plant tolerances. Native to and widespread in subtropical and tropical regions, including parts of Asia, this Pteridaceae fern grows in diverse environments from old mine sites to naturally arsenic-enriched soils.^[64]^[65] Hyperaccumulator species exhibit broad distribution patterns, with 721 known taxa (as documented in the Global Hyperaccumulator Database as of 2017) spanning approximately 50 families, predominantly in metalliferous ecosystems like ultramafic, calamine, and serpentine soils across temperate, tropical, and subtropical zones. Many of these plants face conservation challenges due to their restricted ranges and vulnerability to habitat disruption from mining activities, underscoring the need for protected status in biodiversity hotspots.^[1]^[66] Effective cultivation of hyperaccumulators generally requires well-drained soils to prevent waterlogging, which can inhibit root function and metal uptake; coarse-textured substrates, such as sandy loams, optimize growth and accumulation performance across species like those in Brassicaceae and Pteridaceae.^[67]^[68]

Element-Specific Accumulators

Hyperaccumulators exhibit remarkable specificity in targeting particular elements, often tied to the geochemical properties of their native soils. Nickel (Ni) hyperaccumulators, defined by thresholds exceeding 1,000 μg Ni g⁻¹ dry weight in aboveground tissues, are particularly prevalent on ultramafic soils rich in serpentine-derived substrates.^[69] A prominent example is Berkheya coddii, an Asteraceae species endemic to South African ultramafic outcrops, which can accumulate up to 76,000 μg Ni g⁻¹ dry weight (7.6%) in leaves, enabling it to thrive in Ni-contaminated environments while excluding other metals.^[70] These plants dominate in regions like the Barberton Greenstone Belt, where ultramafic weathering contributes to elevated soil Ni levels, fostering specialized floras adapted to metalliferous stress.^[71] Selenium (Se) hyperaccumulators, typically those exceeding 1,000 μg Se g⁻¹ dry weight, are adapted to seleniferous soils where Se toxicity poses risks to non-tolerant species. Astragalus bisulcatus, a Fabaceae native to western U.S. regions such as Wyoming and Colorado, exemplifies this by accumulating up to 14,000 μg Se g⁻¹ dry weight in shoots, primarily in young leaves, which confers herbivore deterrence but heightens toxicity risks for grazing livestock in these arid, alkaline soils.^[72] Ecological implications include altered arthropod communities, with fewer herbivores on high-Se plants, underscoring Se's role in plant defense within seleniferous hotspots like the Great Plains.^[73] For rare earth elements (REEs), hyperaccumulation is characterized by concentrations surpassing 1,000 μg g⁻¹ total REEs in dry biomass, often linked to ion-adsorption clay deposits. The fern Dicranopteris linearis (Gleicheniaceae), found on southern Chinese mine tailings, hyperaccumulates lanthanum (La) up to 2,000 μg g⁻¹ dry weight in fronds, alongside other light REEs like cerium, facilitating tolerance to acidic, REE-enriched soils derived from weathered granites. This species' ecology is tied to subtropical forests overlying ion-adsorption deposits in Jiangxi Province, where it aids in stabilizing eroded, REE-contaminated landscapes.^[74] Some hyperaccumulators demonstrate multi-element capabilities, accumulating multiple metals simultaneously due to overlapping soil geochemistry. Haumaniastrum robertii (Lamiaceae), an obligate metallophyte from the Democratic Republic of Congo's Copperbelt, hyperaccumulates both cobalt (Co) up to 10,000 μg g⁻¹ and copper (Cu) exceeding 5,000 μg g⁻¹ dry weight in leaves, thriving on Cu-Co rich katangan supergene ores.^[75] This dual tolerance highlights adaptations to polymetallic substrates, where Co and Cu co-occur at high levels, influencing local biodiversity in tropical savannas.^[76] Global hotspots for element-specific hyperaccumulator diversity are concentrated in geologically ancient, metalliferous terrains. Cuba stands out for nickel endemics, hosting over 128 Ni hyperaccumulator species—more than any other region—primarily in the serpentine floras of eastern and western ultramafic massifs, such as the Sierra del Cristal, where endemics like those in Buxaceae and Euphorbiaceae evolved in isolation on Ni-rich ophiolites.^[1] These areas, alongside New Caledonia and the Congo Basin, represent evolutionary cradles for metal-specific adaptations, with Cuban species contributing significantly to ultramafic endemism.^[77]

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Jun 2, 2017 · HMA3, a heavy metal transporter, plays an important role in heavy metal detoxification in plants by sequestering toxic or excess heavy metals into the vacuoles.
[32]
Protein Biochemistry and Expression Regulation of Cadmium/Zinc ...
May 22, 2017 · Since hyperaccumulators are well-known for having an enhanced root to shoot translocation, the enhanced expression of HMA transporters ...
[33]
ZmHMA3, a Member of the Heavy-Metal-Transporting ATPase ... - NIH
P1B-ATPases ... These genes play a primary and important role in metal translocation from the root to the shoot and subsequent metal accumulation within the shoot ...
[34]
The molecular mechanism of zinc and cadmium stress response in ...
In this review, we will compare the different molecular mechanisms of nutritional (Zn) and non-nutritional (Cd) metal homeostasis between metal-sensitive and ...Metal Chelation · Metal Sequestration And... · Ros And Stress Response
[35]
MAPK Cascades and Transcriptional Factors: Regulation of Heavy ...
Apr 18, 2022 · In addition to bZIP, MYB, WRKY and HSF, other transcription factor families also regulate the heavy metal response. In Aegilops markgrafii, ...
[36]
Genome-wide analysis of WRKY gene family and the dynamic ...
Oct 10, 2024 · The WRKY transcription factors comprise one of the most extensive gene families and serve as pivotal regulators of plant responses to heavy ...
[37]
Traversing the Links between Heavy Metal Stress and Plant Signaling
Feb 5, 2018 · ... WRKY transcription factors can work coordinately with MAPK cascade during heavy metal stress tolerance. Plant bZIP transcription factors are ...
[38]
Genome-wide analysis of plant metal transporters, with an emphasis ...
In angiosperms, the high number of ZIP transporters would facilitate the absorption of metal ions from the soil and the distribution throughout the plant. ...
[39]
Vacuolar Transporters – Companions on a Longtime Journey - PMC
By storing and releasing a multitude of compounds, vacuoles play a multifaceted role in the plant development and response to environment Al changes.
[40]
Phytoremediation of heavy metals in soil and water: An eco-friendly ...
Phytoremediation is a realistic and promising strategy for heavy metal removal from polluted areas, based on the employment of hyper-accumulator plant species.Phytoremediation Of Heavy... · 4. Phytoremediation Of Hms... · 6. Factors Affecting...<|control11|><|separator|>
[41]
[PDF] Phytoremediation of Contaminated Soil and Ground Water at ...
For phytostabilization, cropping system costs have been estimated at $200 to $10,000 per hectare, equivalent to $0.02 to $1.00 per cubic meter of soil, assuming ...
[42]
Comparative Cost Analysis of Asbestos Remediation Methods
Nov 1, 2024 · Cost Range: $80 - $150 per cubic meter, affected by site location, depth of contamination, and transportation distance to licensed facilities.
[43]
The nickel hyperaccumulator plant Alyssum bertolonii as a potential ...
Experiments were carried out in Italy on the potential use of the hyperaccumulator Alyssum bertolonii in phytomining of ultramafic soils for Ni.
[44]
Phytoremediation: Mechanisms, plant selection and enhancement ...
Phytoremediation is a cost-effective and environmentally friendly technique that utilizes plants to immobilize, uptake, reduce toxicity, stabilize, or degrade ...
[45]
Comparing the risk of metal leaching in phytoremediation using ...
Metals that cannot be accumulated are defined as non-hyperaccumulated metals and can increase the leaching risk in phytoremediation.
[46]
EDTA-assisted Pb phytoextraction - PubMed
Phytoextraction has been proposed as an inexpensive, sustainable, in situ plant-based technology that makes use of natural hyperaccumulators as well as high ...
[47]
Biofortification of crops with seven mineral elements often lacking in ...
Jan 15, 2009 · This article reviews aspects of soil science, plant physiology and genetics underpinning crop biofortification strategies
[48]
Biofortification and phytoremediation of selenium in China - Frontiers
Biofortiﬁcation is an agricultural process that increases the uptake and accumulation of specific nutrients (Rouached, 2013), e.g., selenium (Se), in ...
[49]
Phytostabilization of Mine Tailings in Arid and Semiarid ... - NIH
Phytostabilization, the use of plants for in situ stabilization of tailings and metal contaminants, is a feasible alternative to costly remediation practices.
[50]
Accumulation of metallic trace elements in Reynoutria japonica
May 6, 2022 · When grown on the POL soil, R. japonica stem and leaf Cd content was close to the EU maximum regulatory limit for organic amendments or animal ...Soil Material And Analyses · Results · Plant Growth Parameters
[51]
Undesirable substances - Food Safety - European Commission
The most important prohibits the dilution of contaminated feed materials. It also includes maximum limits for heavy metals such as arsenic, lead, mercury and ...
[52]
Commission Regulation (EU) No 1275/2013 - Legislation.gov.uk
Higher maximum levels for these heavy metals in long-term supply formulations do not, however, entail a risk for animal or public health or the environment as ...
[53]
CRISPR-enhanced iron accumulation in TBR225 rice via <i ...
Mar 26, 2025 · This study builds upon previous research that used CRISPR/Cas9 technology to overexpress a key iron transporter gene (OsNRAMP7) in TBR225.Missing: hyperaccumulation | Show results with:hyperaccumulation
[54]
Iron-biofortified Rice Developed to Help Combat Iron Deficiency ...
Apr 2, 2025 · Vietnam National University researchers reported the development of iron-rich rice using CRISPR-Cas9 gene editing. The results are published ...Missing: enrichment 2015
[55]
CRISPR-mediated promoter editing of a cis-regulatory element of ...
Jan 23, 2024 · This research studied the OsNAS2 promoter sequence modification introduced using the genome-editing tool CRISPR in the elite rice variety IR64.Missing: hyperaccumulation | Show results with:hyperaccumulation
[56]
Biofortifications Market Size, Share, Industry, Forecast and Outlook ...
Oct 30, 2024 · Global Biofortifications Market reached US$ 110.2 million in 2022 and is expected to reach US$ 221.5 million by 2031.
[57]
[PDF] Biofortification Evidence | HarvestPlus
Biofortification increases the amount of micronutrients in staple food crops through conventional plant breeding, agronomic practices,.
[58]
Identification of Thlaspi caerulescens Genes That May Be Involved ...
Certain ecotypes of T. caerulescens can accumulate Zn and Cd to extremely high levels in the shoot, with Zn reaching levels as high as 30,000 μg g−1 (Brown ...
[59]
Thlaspi caerulescens, an attractive model species to study heavy ...
Aug 9, 2025 · ... Hyperaccumulator plants can grow and reproduce on natural metalliferous soils such as calamine soils (enriched in Zn and Pb) and serpentine ...
[60]
Harnessing hyperaccumulator plants to recover technology‐critical ...
Mar 11, 2025 · The widely accepted definition of trace element hyperaccumulators refers to plants that, when grown in their natural habitats (as opposed to ...<|control11|><|separator|>
[61]
A nickel phytomining field trial using Odontarrhena chalcidica and ...
Jul 15, 2019 · Research article. A nickel phytomining field trial using Odontarrhena chalcidica and Noccaea goesingensis on an Austrian serpentine soil.
[62]
Comparison of root absorption, translocation and tolerance of ...
Nov 12, 2004 · The first As hyperaccumulator described recently was the fern Pteris vittata, which accumulated up to 7500 mg As kg−1 in its fronds at an As- ...
[63]
New evidence of the arsenic uptake and translocation in As ...
Pteris vittata (Chinese brake fern) is a tropical-subtropical plant and was the first identified arsenic hyperaccumulator [5]. As accumulation and ...
[64]
Global Distribution and Ecology of Hyperaccumulator Plants
Around 400 species of plant from 45 families ... Conservation issues are important because many hyperaccumulators are rare and restricted in their distribution.
[65]
Soil Phytomining: Recent Developments—A Review - MDPI
Hyperaccumulators have the potential of absorbing metals from soil via their root system and translocating them into their aerial parts [4]. In this way, a ...
[66]
Soil texture and climate limit cultivation of the arsenic ...
Aug 15, 2022 · We show that Pteris vittata, an arsenic hyperaccumulating fern used to phytoextract arsenic, grows better in coarse- compared to fine-textured soils.
[67]
Uptake and distribution of nickel and other metals in the ...
Apr 11, 2003 · Berkheya coddii is a fast-growing, high biomass nickel (Ni) hyperaccumulator plant that has recently attracted attention for its possible use in the ...
[68]
Uptake, translocation and accumulation of nickel and cobalt in ...
Jun 19, 2020 · The hyperaccumulator plant Berkheya coddii from South Africa is one of the most promising 'agromining crops' known globally. This article ...
[69]
Ecophysiology of nickel hyperaccumulating plants from South Africa
May 27, 2020 · Berkheya coddii is a plant with high potential for phytoremediation and phytomining due to its large biomass and potentially high Ni yield, that ...
[70]
Selenium hyperaccumulation reduces plant arthropod loads in the ...
Nov 19, 2007 · The Se hyperaccumulating plant species, which contained 1000–14 000 µg Se g−1 DW, harbored significantly fewer arthropods (c. twofold) and fewer ...
[71]
Spatial Imaging, Speciation, and Quantification of Selenium in the ...
Astragalus bisulcatus and Stanleya pinnata hyperaccumulate selenium (Se) up to 1% of plant dry weight. In the field, Se was mostly present in the young leaves ...
[72]
Rare earth elements in naturally grown fern Dicranopteris linearis in ...
The situations mentioned above are mainly because of the very low concentrations of REEs in soil and plant (10−4–10−2 ug/g; Laul et al., 1979, Sun et al., 1994 ...
[73]
Copper and cobalt accumulation in plants: a critical assessment of ...
Sep 14, 2016 · This review synthesizes contemporary understanding of copper–cobalt (Cu–Co) tolerance and accumulation in plants.
[74]
[PDF] Copper and cobalt accumulation in plants - HAL
In the Copper Flora, Haumaniastrum robertii, with leaf Co. 364 concentration can >4000 μg g-1 (Morrison et al., 1981), might be a good candidate for Co. 365.
[75]
Nickel Hyperaccumulation in the Serpentine Flora of Cuba
A recent study of the families Buxaceae and Euphorbiaceae identified a further 80 Ni hyperaccumulators from the very large ultramafic flora of Cuba.