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Zinc (Zn) is an essential micronutrient for plants and animals. Unfortunately, deficiency of Zn in humans has increased on a global scale. The main reason of this micronutrient deficiency is dietary intakes of food with low Zn levels. Adoption of biofortification approaches would result in Zn enrichment of target tissue to a considerable extent. However, there is a basic need to understand Zn absorption mechanisms in plants prior to exploitation of such practical approaches. Zn absorption is a complex physiological trait which is mainly governed by Zn transporters and metal chelators of plant system. Plant growth stage, edaphic factors, season etc. also influence Zn efficiency of particular species. Molecular studies in Zn hyperaccumulators have already demonstrated the participation of specific Zn transporters, vacuolar sequestration and detoxification mechanisms in maintenance of Zn homeostasis. These have been described in detail in present review and provide opportunities for utilization in biofortification programmes. However, issues such as lesser bioavailability of Zn in target organ, uptake of toxic divalent cations (Cd, Ni, Pb, As etc.) along with Zn, sink activity and dilution in Zn concentration in response to sink number etc. in biofortified crops need further investigation. In order to design novel strategy in biofortification programmes, future researches should focus on physiological performance and yield penalties in concerned crop, metabolic load in term of organic acid production and crosstalk of Zn with other mineral nutrients under low and high Zn conditions.

Background Plants that accumulate metal and metalloid trace elements to extraordinarily high concentrations in their living biomass have inspired much research worldwide during the last decades. Hyperaccumulators have been recorded and experimentally confirmed for elements such as nickel, zinc, cadmium, manganese, arsenic and selenium. However, to date, hyperaccumulation of lead, copper, cobalt, chromium and thallium remain largely unconfirmed. Recent uses of the term in relation to rare-earth elements require critical evaluation. Scope Since the mid-1970s the term 'hyperaccumulator' has been used millions of times by thousands of people, with varying degrees of precision, aptness and understanding that have not always corresponded with the views of the originators of the terminology and of the present authors. There is therefore a need to clarify the circumstances in which the term 'hyperaccumulator' is appropriate and to set out the conditions that should be met when the terms are used. We outline here the main considerations for establishing metal or metalloid hyperaccumulation status of plants, (re) defme some of the terminology and note potential pitfalls. Conclusions Unambiguous communication will require the international scientific community to adopt standard terminology and methods for confirming the reliability of analytical data in relation to metal and metalloid hyperaccumulators.

Cadmium (Cd) is highly toxic to most organisms, but some rare plant species can hyperaccumulate Cd in aboveground tissues without suffering from toxicity. The mechanism underlying Cd detoxification by hyperaccumulators is interesting but unclear. Here, the heavy metal ATPase 3 (SpHMA3) gene responsible for Cd detoxification was isolated from the Cd/zinc (Zn) hyperaccumulator Sedum plumbizincicola. RNA interference (RNAi)-mediated silencing and overexpression of SpHMA3 were induced to investigate its physiological functions in S. plumbizincicola and a nonhyperaccumulating ecotype of Sedum alfredii. Heterologous expression of SpHMA3 in Saccharomyces cerevisiae showed Cd-specific transport activity. SpHMA3 was highly expressed in the shoots and the protein was localized to the tonoplast. The SpHMA3-RNAi lines were hypersensitive to Cd but not to Zn, with the growth of shoots and young leaves being severely inhibited by Cd. Overexpressing SpHMA3 in the nonhyperaccumulating ecotype of S. alfredii greatly increased its tolerance to and accumulation of Cd, but not Zn. These results indicate that elevated expression of the tonoplast-localized SpHMA3 in the shoots plays an essential role in Cd detoxification, which contributes to the maintenance of the normal growth of young leaves of S. plumbizincicola in Cd-contaminated soils.

Hyperaccumulator plants are unusual plants that accumulate particular metals or metalloids, such as nickel, zinc, cadmium and arsenic, in their living tissues to concentrations that are hundreds to thousands of times greater than what is normal for most plants. The hyperaccumulation phenomenon is rare (exhibited by less than 0.2% of all angiosperms), with most of the similar to 500 hyperaccumulator species known globally for nickel.This review highlights the contemporary understanding of nickel hyperaccumulation processes, which include root uptake and sequestration, xylem loading and transport, leaf compartmentation and phloem translocation processes.Hyperaccumulator plants have evolved highly efficient physiological mechanisms for taking up nickel in their roots followed by rapid translocation and sequestration into the aerial shoots. The uptake of nickel is mainly involved with low affinity transport systems, presumably from the ZIP family. The presence of high concentrations of histidine prevents nickel sequestration in roots. Nickel is efficiently loaded into the xylem, where it mainly presents as Ni2+. The leaf is the main storage organ, which sequestrates nickel in non-active sites, e.g. vacuoles and apoplast. Recent studies show that phloem translocates high levels of nickel, which has a strong impact on nickel accumulation in young growing tissues.

Chromium (Cr) is the second most common metal contaminant in ground water, soil, and sediments due to its wide industrial application, hence posing a serious environmental concern. Among various valence states, Cr(III) and Cr(VI) are the most stable forms. Cr(VI) is the most persistent in the soil and is highly toxic for biota. Since Cr is a non-essential element for plants, there is no uptake mechanism; Cr is taken up along essential elements such as sulfate through sulfate transporters. Cr accumulation in plants causes high toxicity in terms of reduction in growth and biomass accumulation, and Cr induces structural alterations. Cr interferes with photosynthetic and respiration processes, and water and minerals uptake mechanism. Various enzymatic activities related to starch and nitrogen metabolism are decreased by Cr toxicity either by direct interference with the enzymes or through the production of reactive oxygen species. Cr causes oxidative damage by destruction of membrane lipids and DNA damage. Cr may even cause the death of plant species. Few plant species are able to accumulate high amount of Cr without being damaged. Such Cr-tolerant, hyperaccumulator plants are exploited for their bioremediation property. The present review discusses Cr availability in the environment, Cr transfer to biota, toxicity issues, effect on germination and plant growth, morphological and ultrastructural aberrations, biochemical and physiological alterations, effect on metabolic processes, Cr-induced alterations at the molecular level, Cr hyperaccumulation and Cr detoxification mechanism, and the role of arbuscular mycorrhizae in Cr toxicity, in plants.

Heavy metals (HMs) normally occur in nature and are rapidly released into ecosystems by anthropogenic activities, leading to a series of threats to plant productivity as well as human health. Phytoremediation is a clean, eco-friendly, and cost-effective method for reducing soil toxicity, particularly in weedy plants (invasive plant species (IPS)). This method provides a favorable tool for HM hyperaccumulation using invasive plants. Improving the phytoremediation strategy requires a profound knowledge of HM uptake and translocation as well as the development of resistance or tolerance to HMs. This review describes a comprehensive mechanism of uptake and translocation of HMs and their subsequent detoxification with the IPS via phytoremediation. Additionally, the improvement of phytoremediation through advanced biotechnological strategies, including genetic engineering, nanoparticles, microorganisms, CRISPR-Cas9, and protein basis, is discussed. In summary, this appraisal will provide a new platform for the uptake, translocation, and detoxification of HMs via the phytoremediation process of the IPS.

Background When studying metallophytes and hyperaccumulator plants, it is often desired to assess the level of tolerance of a specific trace metal/metalloid in a putative tolerant species, to determine root and shoot accumulation of the trace metal/metalloid of interest, or to establish whether a trace metal/metalloid has an essential function. The use of hydroponics has proven to be a powerful tool in answering such questions in relation to the physiological regulation of metal/metalloids in plants. Carefully designing experiments requires considering nutrient solution formulation, dose rate regime, and environmental conditions, but this is often overlooked. Aims This review aims to bring together key information for hydroponics studies in physiological, evolutionary, and genetics/molecular biological research of trace metal/metalloid tolerance and accumulation in plants, focussing on metallophytes and hyperaccumulator plants. Conclusions It is not possible to define a ‘universal’ nutrient solution that is both sufficient and non-toxic for all plants, although it is often possible, dependent on plant species under study and the research question to be addressed, to ‘adapt’ commonly used ‘standard formulations’. Well-designed and executed hydroponics experiments can yield powerful insights in the regulation of essential and toxic metal/metalloid trace elements, and this extends far beyond hyperaccumulator plants.

Pollution by heavy metals (HM) represents a serious threat for both the environment and human health. Due to their elemental character, HM cannot be chemically degraded, and their detoxification in the environment mostly resides either in stabilization or in their removal from the matrix, e.g., soil. For this purpose, phytoremediation, i.e., the application of plants for the restoration of a polluted environment, has been proposed as a promising green alternative to traditional physical and chemical methods. Among the phytoremediation techniques, phytoextraction refers to the removal of HM from the matrix through their uptake by a plant. It possesses considerable advantages over traditional techniques, especially due to its cost effectiveness, potential treatment of multiple HM simultaneously, no need for the excavation of contaminated soil, good acceptance by the public, the possibility of follow-up processing of the biomass produced, etc. In this review, we focused on three basic HM phytoextraction strategies that differ in the type of plant species being employed: natural hyperaccumulators, fast-growing plant species with high-biomass production and, potentially, plants genetically engineered toward a phenotype that favors efficient HM uptake and boosted HM tolerance. Considerable knowledge on the applicability of plants for HM phytoextraction has been gathered to date from both lab-scale studies performed under controlled model conditions and field trials using real environmental conditions. Based on this knowledge, many specific applications of plants for the remediation of HM-polluted soils have been proposed. Such studies often also include suggestions for the further processing of HM-contaminated biomass, therefore providing an added economical value. Based on the examples presented here, we recommend that intensive research be performed on the selection of appropriate plant taxa for various sets of conditions, environmental risk assessment, the fate of HM-enriched biomass, economical aspects of the process, etc.

Toxic metals/metalloids such as arsenic are environmental pollutants that are damaging to living organisms. Commonly found metals pose a serious threat to human health unless they are controlled. In this study, the accumulation of As metalloid, which has entered the food chain for specific reasons, in plant organs was determined, and it was examined whether plants are hyperaccumulators against As concentration. As a result, Pinus pinaster Aiton., Cupressus arizonica Greene., Picea orientalis (L.) Peterm., Cedrus atlantica (Endl.), and Pseudotsuga menziesii (Mirb.) species accumulated more than 1 mg/kg As concentration. The highest average As concentration (7.91 mg/kg) was found in the northern bark of P. menziesii. The highest As concentrations in wood were found in the eastern direction for P. pinaster (5.21 mg/kg), C. arizonica (4.99 mg/kg), and P. orientalis (4.02 mg/kg), and in the western direction for C. atlantica (3.56 mg/kg) and P. menziesii (3.88 mg/kg). Additionally, it was determined that As concentration varied depending on location, direction, species, and year.

Globally, ultramafic outcrops are renowned for hosting floras with high levels of endemism, including plants with specialised adaptations such as nickel or manganese hyperaccumulation. Soils derived from ultramafic regoliths are generally nutrient-deficient, have major cation imbalances, and have concomitant high concentrations of potentially phytotoxic trace elements, especially nickel. The South and Southeast Asian region has the largest surface occurrences of ultramafic regoliths in the world, but the geoecology of these outcrops is still poorly studied despite severe conservation threats. Due to the paucity of systematic plant collections in many areas and the lack of georeferenced herbarium records and databased information, it is not possible to determine the distribution of species, levels of endemism, and the species most threatened. However, site-specific studies provide insights to the ultramafic geoecology of several locations in South and Southeast Asia. The geoecology of tropical ultramafic regions differs substantially from those in temperate regions in that the vegetation at lower elevations is generally tall forest with relatively low levels of endemism. On ultramafic mountaintops, where the combined forces of edaphic and climatic factors intersect, obligate ultramafic species and hyperendemics often occur. Forest clearing, agricultural development, mining, and climate change-related stressors have contributed to rapid and unprecedented loss of ultramafic-associated habitats in the region. The geoecology of the large ultramafic outcrops of Indonesia’s Sulawesi, Obi and Halmahera, and many other smaller outcrops in South and Southeast Asia, remains largely unexplored, and should be prioritised for study and conservation.