Nickel Hyperaccumulator Research Articles

Evaluation of the performance of nickel hyperaccumulator plants as combustion fuel

Agromining is an emerging technology aiming at recovering metals from soils using hyperaccumulator (HA) plants. Plant cultivation and pyro- and hydrometallurgical processes have been designed for nickel (Ni) recovery. In this chain of processes, dry plants are burnt to obtain Ni-rich ashes that are subsequently treated. To ensure the sustainability and the profitability of the process, energy recovered at the combustion stage would add some value. The combustion of three Ni HA plants (Odontarrhena chalcidica, Leptoplax emarginata and Berkheya coddii) was investigated. Namely the higher and lower heating values (HHV and LHV) were measured with a calorimeter bomb and calculated with correlations, thermo-gravimetric analysis was performed, dry plants and ashes were characterized, and gas emissions were monitored during combustion at two temperatures. In a complementary approach, the self-heating behaviour of plants and ashes were compared, and the activation energy of such phenomenon was determined. These results allowed the identification of the step limiting the self-inflammation of the plants, which gives relevant information for their safe storage.

A systematic assessment of the occurrence of trace element hyperaccumulation in the flora of New Caledonia

AbstractNew Caledonia is a global biodiversity hotspot known for its metal hyperaccumulator plants. X-ray fluorescence technology (XRF) has enabled non-destructive and quantitative determination of elemental concentrations in herbarium specimens from the ultramafic flora of the island. Specimens belonging to six major hyperaccumulator families (Cunoniaceae, Phyllanthaceae, Salicaceae, Sapotaceae, Oncothecaceae and Violaceae) and one to four specimens per species of the remaining ultramafic taxa in the herbarium were measured. XRF scanning included a total of c. 11 200 specimens from 35 orders, 96 families, 281 genera and 1484 species (1620 taxa) and covered 88.5% of the ultramafic flora. The study revealed the existence of 99 nickel hyperaccumulator taxa (65 known previously), 74 manganese hyperaccumulator taxa (11 known previously), eight cobalt hyperaccumulator taxa (two known previously) and four zinc hyperaccumulator taxa (none known previously). These results offer new insights into the phylogenetic diversity of hyperaccumulators in New Caledonia. The greatest diversity of nickel hyperaccumulators occur in a few major clades (Malphigiales and Oxalidales) and families (Phyllanthaceae, Salicaceae, Cunoniaceae). In contrast, manganese hyperaccumulation is phylogenetically scattered in the New Caledonian flora.

Synchrotron µXRF imaging of live seedlings of Berkheya coddii and Odontarrhena muralis during germination and seedling growth

Nickel hyperaccumulator plants require highly evolved mechanisms to avoid cellular-level toxicity to cope with the high prevailing concentrations of nickel in their seeds and seedlings. This study aimed to investigate tissue-level distribution and redistribution of Ni and other physiologically relevant elements during the germination of hyperaccumulator plants. Berkheya coddii Roessler (Asteraceae) and Odontarrhena muralis sensu latu (Waldst. & Kit.) Endl. (O. chalcidica), formerly Alyssum murale (Waldst. & Kit.) (Brassicaceae), were germinated for several days at room temperature before using synchrotron micro X-ray Fluorescence Spectroscopy (µXRF) to obtain high-sensitivity and high-resolution elemental images of live/hydrated plants at various stages of seed germination and seedling growth. The results show that fruits and seeds of both species were highly enriched in nickel. In Berkheya coddii, nickel was located in the cotyledons, the micropylar area, the seed coat, and the point of attachment of the pappus with the cypsela body. In Odontarrhena muralis seeds, nickel occurred in the cotyledons and hypocotyl. The emergence of true leaves initiates nickel and calcium redistribution within seedlings in both species. The diversity in physiological responses to nickel in Berkheya coddii and Odontarrhena muralis does not only occur at the mature stage, but is inherent to both species as seed elemental storage and tolerance mechanisms during seedling development differ.

Nickel hyperaccumulation in New Caledonian Hybanthus (Violaceae) and occurrence of nickel-rich phloem in Hybanthus austrocaledonicus.

Hybanthus austrocaledonicus (Violaceae) is a nickel (Ni) hyperaccumulator endemic to New Caledonia. One of the specimens stored at the local herbarium had a strip of bark with a remarkably green phloem tissue attached to the sheet containing over 4 wt% Ni. This study aimed to collect field samples from the original H. austrocaledonicus locality to confirm the nature of the green 'nickel-rich phloem' in this taxon and to systematically assess the occurrence of Ni hyperaccumulation in H. austrocaledonicus and Hybanthus caledonicus populations. X-ray fluorescence spectroscopy scanning of all collections of the genus Hybanthus (236 specimens) was undertaken at the Herbarium of New Caledonia to reveal incidences of Ni accumulation in populations of H. austrocaledonicus and H. caledonicus. In parallel, micro-analytical investigations were performed via synchrotron X-ray fluorescence microscopy (XFM) and scanning electron microscopy with X-ray microanalysis (SEM-EDS). The extensive scanning demonstrated that Ni hyperaccumulation is not a characteristic common to all populations in the endemic Hybanthus species. Synchrotron XFM revealed that Ni was exclusively concentrated in the epidermal cells of the leaf blade and petiole, conforming with the majority of (tropical) Ni hyperaccumulator plants studied to date. SEM-EDS of freeze-dried and frozen-hydrated samples revealed the presence of dense solid deposits in the phloem bundles that contained >8 wt% nickel. The occurrence of extremely Ni-rich green phloem tissues appears to be a characteristic feature of tropical Ni hyperaccumulator plants.

Combustion of nickel hyperaccumulator plants investigated by experimental and thermodynamic approaches

This work was undertaken to determine whether thermodynamic calculations can predict the amount and composition of ash produced by combustion of nickel hyperaccumulator (HA) plants in the context of agromining. To this end, three HA plants containing about 1wt% of nickel were submitted to isothermal combustion at 550, 815, 900°C and the solid residues were characterized by complementary techniques (TGA, XRD, ICP-OES, ion chromatography, SEM-EDX), while thermodynamic calculations were performed with the FactSage software using the dry biomass composition as input. We present an in-depth evaluation of inorganics behavior based on the combination and comparison of experimental data and calculations: effect of temperature on ash yield, nature of crystallized phases, ash elemental composition and inorganics volatilization. Our work confirms that equilibrium calculation is a powerful method to evaluate the behavior of inorganic elements during thermal treatment of biomass, despite some lacks in the databases. For the nickel HA plants, an innovative result is that, after combustion, nickel is always in the form of NiO particles, that can mix with MgO to form (Ni,Mg)O solid solutions. These results have direct implications for understanding the combustion process and for better design of the following leaching step aiming at Ni recovery.

Elevated Expression of Vacuolar Nickel Transporter Gene IREG2 Is Associated With Reduced Root-to-Shoot Nickel Translocation in Noccaea japonica.

A number of metal hyperaccumulator plants, including nickel (Ni) hyperaccumulators, have been identified in the genus Noccaea. The ability to accumulate Ni in shoots varies widely among species and ecotypes in this genus; however, little is known about the molecular mechanisms underlying this intra- and inter-specific variation. Here, in hydroponic culture, we compared Ni accumulation patterns between Noccaea japonica, which originated in Ni-enriched serpentine soils in Mt. Yubari (Hokkaido, Japan), and Noccaea caerulescens ecotype Ganges, which originated in zinc/lead-mine soils in Southern France. Both Noccaea species showed extremely high Ni tolerance compared with that of the non-accumulator Arabidopsis thaliana. But, following treatment with 200 μM Ni, N. caerulescens showed leaf chlorosis, whereas N. japonica did not show any stress symptoms. Shoot Ni concentration was higher in N. caerulescens than in N. japonica; this difference was due to higher efficiency of root-to-shoot Ni translocation in N. caerulescens than N. japonica. It is known that the vacuole Ni transporter IREG2 suppresses Ni translocation from roots to shoots by sequestering Ni in the root vacuoles. The expression level of the IREG2 gene in the roots of N. japonica was 10-fold that in the roots of N. caerulescens. Moreover, the copy number of IREG2 per genome was higher in N. japonica than in N. caerulescens, suggesting that IREG2 expression is elevated by gene multiplication in N. japonica. The heterologous expression of IREG2 of N. japonica and N. caerulescens in yeast and A. thaliana confirmed that both IREG2 genes encode functional vacuole Ni transporters. Taking these results together, we hypothesize that the elevation of IREG2 expression by gene multiplication causes the lower root-to-shoot Ni translocation in N. japonica.

Ecophysiology of nickel hyperaccumulating plants from South Africa - from ultramafic soil and mycorrhiza to plants and insects.

An overview of 30 years of studies related to South African nickel hyperaccumulators is presented. Only five species have so far been identified as Ni hyperaccumulator plants among very rich and diversified South African flora. All of them occur on soils derived from ultramafic (serpentine) rocks and belong to the family Asteraceae: Berkheya coddii Roessler, Berkheya zeyheri subsp. rehmannii var. rogersiana, Berkheya nivea, Senecio coronatus, Senecio anomalochrous. Several techniques and methods were used to investigate ecophysiological aspects of the Ni hyperaccumulation phenomenon, from basic field and laboratory studies, to advanced instrumental methods. Analysis of elemental distribution in plant parts showed that in most cases the hyperaccumulated metal was stored in physiologically inactive tissues such as the foliar epidermis. However, an exception is Berkheya coddii, which has a distinctly different pattern of Ni distribution in leaves, with the highest concentration in the mesophyll. Such a distribution suggests that different physiological mechanisms are involved in the Ni transport, storage location and detoxification, compared to other hyperaccumulator species. Berkheya coddii is a plant with high potential for phytoremediation and phytomining due to its large biomass and potentially high Ni yield, that can reach 7.6% of Ni in dry mass of leaves. Senecio coronatus is the only known hyperaccumulator with two genotypes, hyperaccumulating and non-hyperaccumulating, growing on Ni-enriched/metalliferous soil. Detailed ultrastructural studies were undertaken to characterize specialized groups of cells in the root cortex of Ni-hyperaccumulating genotype, that are not known from any other hyperaccumulator. The occurrence of arbuscular mycorrhiza (AM) in Ni-hyperaccumulating plants was found for the first time in South African hyperaccumulator plants, and this type of symbiosis has been proved obligatory in all of them. There is a significant influence of mycorrhiza on the concentration and distribution of several elements. Three highly specialized herbivore insects feeding only on Ni hyperaccumulator plants were identified: Chrysolina clathrata (formerly Chrysolina pardalina), Epilachna nylanderi and Stenoscepa sp. The Ni-elimination strategies of these specialised insects have been established. Microbiological studies have revealed several genera of fungi and bacteria isolated from B. coddii leaves as well as presence of specialised, Ni-resistant yeasts in the C. clathrata gut. Understanding ecophysiological response to harsh environment broadens our knowledge and can have practical applications in cleaning polluted environments through phytomining/agromining. Finally, conservation aspects are also discussed and lines for future research are proposed.

Use and validation of magnetic properties for differentiating nickel hyperaccumulators and non-nickel hyperaccumulators in ultramafic regions

Nickel hyperaccumulators are plants that can absorb a large amount of Ni in their tissues. Nickel hyperaccumulators have a strong potential to be used as bioindicators in mineral explorations. Previous studies show that analysis of magnetic susceptibility has a potential to accelerate the screening of Ni hyperaccumulators. In this study, we investigated the use magnetic characterizations for screening directly Ni hyperaccumulators and non-Ni hyperaccumulators from ultramafic regions. For comparison, the characterization of Ni hyperaccumulators was also performed on Alyssum murale and Alyssum corsicum, which were cultivated on lateritic soils. The metals concentrations in the tissues were determined via inductively coupled plasma optical emission spectroscopy analysis, atomic absorption spectroscopy analysis, and X-ray fluorescence analysis. The magnetic characterizations were conducted by magnetic susceptibility measurement, acquisition of hysteresis curves, and analysis of the thermomagnetic properties. The results showed that the concentration of Ni in A. murale and A. corsicum were correlated to the magnetic susceptibility values. Concurrently, all samples had negative magnetic susceptibility values were non-Ni hyperaccumulators. Two new species of Ni hyperaccumulators were found in Halmahera and Sulawesi and have high potential for use in phytoremediation, phytomining or mineral exploration.

Bacterial community diversity in the rhizosphere of nickel hyperaccumulator plant species from Borneo Island (Malaysia).

The Island of Borneo is a major biodiversity hotspot, and in the Malaysian state of Sabah, ultramafic soils are extensive and home to more than 31 endemic nickel hyperaccumulator plants. The aim of this study was to characterize the structure and the diversity of the rhizosphere bacterial communities of several of these nickel hyperaccumulator plants and factors that affect these bacterial communities in Sabah. The most abundant phyla were Proteobacteria, Acidobacteria and Actinobacteria. At family level, Burkholderiaceae and Xanthobacteraceae (Proteobacteria phylum) were the most abundant families in the hyperaccumulator rhizospheres. Redundancy analysis based on soil chemical analyses and relative abundances of the major bacterial phyla showed that abiotic factors of the studied sites drove the bacterial diversity. For all R. aff. bengalensis rhizosphere soil samples, irrespective of studied site, the bacterial diversity was similar. Moreover, the Saprospiraceae family showed a high representativeness in the R. aff. bengalensis rhizosphere soils and was linked with the nickel availability in soils. The ability of R. aff. bengalensis to concentrate nickel in its rhizosphere appears to be the major factor driving the rhizobacterial community diversity unlike for other hyperaccumulator species.

X-ray fluorescence elemental mapping of roots, stems and leaves of the nickel hyperaccumulators Rinorea cf. bengalensis and Rinorea cf. javanica (Violaceae) from Sabah (Malaysia), Borneo

There are major knowledge gaps in understanding the translocation leading from nickel uptake in the root to accumulation in other tissues in tropical nickel hyperaccumulator plant species. This study focuses on two species, Rinorea cf. bengalensis and Rinorea cf. javanica and aims to elucidate the similarities and differences in the distribution of nickel and physiologically relevant elements (potassium, calcium, manganese and zinc) in various organs and tissues. High-resolution X-ray fluorescence microscopy (XFM) of frozen-hydrated and fresh-hydrated tissue samples and nuclear microprobe (micro-PIXE) analysis of freeze-dried samples were used to provide insights into the in situ elemental distribution in these plant species. This study has shown that the distribution pattern of nickel hyperaccumulation is typified by very high levels of accumulation in the phloem bundles of roots and stems. In the leaves, nickel is preferentially located in epidermal cell region, whereas manganese is located mainly in the lower epidermis and zinc in the upper epidermis and palisade mesophyll. The abundant formation of calcium-oxalate crystals, lining the collenchyma, is a prominent feature of both Rinorea cf. bengalensis and Rinorea cf. javanica. Future investigations on Rinorea cf. bengalensis and Rinorea cf. javanica should focus on unravelling the mechanisms of nickel uptake in the root, specifically targeting the identification of nickelspecific membrane transporters.

A preliminary survey of nickel, manganese and zinc (hyper)accumulation in the flora of Papua New Guinea from herbarium X-ray fluorescence scanning

The flora of Papua New Guinea is amongst the richest in the world with an estimated 25,000 plant species. The extreme levels of biodiversity, climatic ranges and soil types suggest a high possibility of metal hyperaccumulator plants existing in Papua New Guinea. However, no hyperaccumulator plants have been reported from this region yet. The use of handheld X-ray fluorescence instruments is a non-destructive and effective method for the systematic quantitative assessment of hyperaccumulation in vast numbers of herbarium specimens. X-ray fluorescence scanning was undertaken at the Queensland Herbarium (Australia) on all Papua New Guinea specimens from seven major families (Celastraceae, Cunoniaceae, Phyllanthaceae, Proteaceae, Rubiaceae, Salicaceae and Violaceae), covering 3164 plant specimens. This preliminary survey revealed the existence of ten zinc hyperaccumulator species (> 3000 µg g−1 Zn), eight manganese accumulator species (> 5000 µg g−1 Mn) and one nickel hyperaccumulator species (> 1000 µg g−1 Ni). These results highlight the potential for discovery of numerous new metal hyperaccumulator plants from the flora of Papua New Guinea if larger-scale systematic screening efforts were undertaken.

Uptake, translocation and accumulation of nickel and cobalt in Berkheya coddii, a 'metal crop' from South Africa.

Hyperaccumulator plants have the ability to efficiently concentrate metallic elements, e.g. nickel, from low-grade sources into their living biomass. Although the majority of nickel hyperaccumulator plant species restrict cobalt uptake, some species are able to co-accumulate cobalt when growing in ultramafic soils. The asteraceous perennial herb Berkheya coddii from South Africa is one of the most promising agromining crops known globally. It may accumulate nickel in excess of 30 000 μg g-1 in dry leaves, while co-accumulating up to 600 μg g-1 cobalt. This study aimed to elucidate the interactions between nickel and cobalt for uptake by and translocation into B. coddii through a pot experiment including various cobalt/nickel treatment combinations in soil, after which uptake and localisation were recorded. Cobalt in the substrate limits nickel uptake by B. coddii plants and is mainly retained in the basal leaves in contrast to Ni that is rapidly transferred to the top of the plant. B. coddii was more tolerant to high Ni concentration, whether in the substrate or internally but remains a promising crop which could be used, with suitable agronomic measures and practices, for cobalt agromining in areas with high soil cobalt but low soil nickel. A yield of 77 kg ha-1 nickel and 16.5 kg ha-1 cobalt may be attainable under optimum conditions.

Phylogenetic and geographic distribution of nickel hyperaccumulation in neotropical Psychotria

Hyperaccumulation of heavy metals in plants has never been documented from Central America or Mexico. Psychotria grandis, P. costivenia, and P. glomerata (Rubiaceae) have been reported to hyperaccumulate nickel in the Greater Antilles, but they also occur widely across the neotropics. The goals of this research were to investigate the geographic distribution of hyperaccumulation in these species and explore the phylogenetic distribution of hyperaccumulation in this clade by testing related species. Portable x-ray fluorescence (XRF) spectroscopy was used to analyze 565 specimens representing eight species of Psychotria from the Missouri Botanical Garden herbarium. Nickelhyperaccumulation was found in specimens of Psychotria costivenia ranging from Mexico to Costa Rica and in specimens of P. grandis from Guatemala to Ecuador and Venezuela. Among related species, nickel hyperaccumulation is reported for the first time in P. lorenciana and P. papantlensis, but no evidence of hyperaccumulation was found in P. clivorum, P. flava, or P. pleuropoda. Previous reports of hyperaccumulation in P. glomerata appear to be erroneous, resulting from taxonomic synonymy and specimen misidentification. Hyperaccumulation of nickel by Psychotria is now known to occur widely from southern Mexico through Central America to northwestern South America, including some areas not known to have ultramafic soils. Novel aspects of this research include the successful prediction of new hyperaccumulator species based on molecular phylogeny, use of XRF technology to nondestructively obtain elemental data from herbarium specimens, and documentation of previously unknown areas of ultramafic or nickel-rich soil based on such data.

Localization of nickel in the hyperaccumulator plant Breynia cernua (Poir.) Mull.Arg. discovered in the nickeliferous laterites of Zambales, the Philippines

A newly discovered nickel (Ni) hyperaccumulator plant, Breynia cernua (Phyllanthaceae) from the nickeliferous laterites of Acoje, Zambales, has elevated Ni concentrations. Its computed bioaccumulation factor is 1.7 and the translocation factor is 4.6. Leaf tissues which contain the highest Ni concentrations were analyzed using optical microscopy and Scanning Electron Microscopy with Energy-Dispersive X-ray (SEM/EDX). The results indicated that most of the Ni are stored in the epidermal cells of the leaves, followed by the mesophyll cells then the vascular bundles. The accumulation of Ni found in the subcellular locations, probably in the vacuoles, may prevent phytotoxicity that could affect the normal function of other cellular structures. A possible resistance mechanism through strain avoidance towards Ni accumulation makes B. cernua capable of tolerating elevated Ni concentrations in its tissues. The tolerance of B. cernua to Ni phytotoxicity suggests possible applications of this hyperaccumulator plant in phytoextraction technology.

Biogeochemistry of the flora of Weda Bay, Halmahera Island (Indonesia) focusing on nickel hyperaccumulation

Indonesia has one of the largest surface expressions of ultramafic rocks on Earth and in parallel hosts one of the most species-rich floras. Despite the extensive knowledge of the botanical diversity and the chemistry of these substrates, until recently the records for nickel hyperaccumulator plant species in the region have been scant. Identification of native local hyperaccumulator species is the critical initial step for developing local phytomining as these species naturally match ambient bioclimatic, geochemical and physiological conditions. Prior to this research just 11 nickel hyperaccumulator plant species were known from the whole of Indonesia. This field-based investigation at Weda Bay on the Island of Halmahera in the North Maluku Province revealed the existence of 13 nickel, two cobalt, one manganese and 10 aluminium hyperaccumulator plant species. Phylogenetic affinity for nickel hyperaccumulation is diverse and spans several orders, but was most frequent in the Malpighiales, as in other ultramafic regions of Southeast Asia. In contrast to global patterns, hyperaccumulation was infrequent in the Phyllanthaceae.

Herbarium X-ray fluorescence screening for nickel, cobalt and manganese hyperaccumulator plants in the flora of Sabah (Malaysia, Borneo Island)

Sabah (Malaysia) on the Island of Borneo has a high plant diversity (>8000 species) occurring on a wide range of soils, including ultramafic soils which are known to host hyperaccumulator plants. In this study a new approach (“Herbarium X-ray Fluorescence Ionomics”) was used to obtain elemental data from herbarium specimens using non-destructive X-ray Fluorescence spectroscopy analysis. In total ~7300 specimens were thus analysed for nickel, cobalt, and manganese concentrations at the Herbarium of the Forest Research Centre (FRC) in Sepilok, Sabah. The measurements led to recording a total of 759 specimens (originating from 17 families in 30 genera and 74 species) as trace element hyperaccumulators, including 28 nickel hyperaccumulator species (in 10 families, 17 genera), 12 cobalt hyperaccumulator species (in 3 families, 7 genera), and 51 manganese hyperaccumulator species (in 12 families, 24 genera). The outcomes of this research demonstrate that X-ray Fluorescence scanning is highly useful approach for hyperaccumulator plant discovery in herbarium collections that has the potential to add vast numbers of hyperaccumulating taxa to the global inventory.

Rhizosphere chemistry and above-ground elemental fractionation of nickel hyperaccumulator species from Weda Bay (Indonesia)

The identification and use of hyperaccumulator plants in mining projects has been recognized as an important component of mine planning at several sites around the world. The objective of this research was to provide information on relevant plant tissue chemistry and an indicative assessment of the potential for phytomining at Weda Bay Nickel (WBN), Halmahera. The first stage was the identification of native nickel hyperaccumulator plants. In total, 280 plant tissue samples from 10 nickel accumulator species and 46 matching rhizosphere soil samples were collected. Chemical analyses of plant tissue samples were performed and physico-chemical parameters of the rhizosphere soils were also measured. A total of three species were considered as metal crops: Rinorea aff. bengalensis (up to 22,200 mg kg−1 dry weight at 2 m above ground level), Ficus trachypison (1060 mg kg−1) and Trichospermum morotaiense (5180 mg kg−1), but only R. aff. bengalensis has sufficiently high Ni concentrations in biomass to warrant field trials. Utilising a successional planting strategy, F. trachypison and T. morotaiense could be used to facilitate site conditions, followed by the metal crop R. aff. bengalensis. Using this design, a nickel yield of 330 kg per hectare would be possible every 4 years. In addition to allowing the recovery of nickel, this approach could be an integrated mine site rehabilitation strategy to mitigate environmental impacts, improve soil quality and facilitate transition to other land-uses such as native forest.

Towards an Understanding of the Molecular Basis of Nickel Hyperaccumulation in Plants.

Metal hyperaccumulation is a rare and fascinating phenomenon, whereby plants actively accumulate high concentrations of metal ions in their above-ground tissues. Enhanced uptake and root-to-shoot translocation of specific metal ions coupled with an increased capacity for detoxification and sequestration of these ions are thought to constitute the physiological basis of the hyperaccumulation phenotype. Nickel hyperaccumulators were the first to be discovered and are the most numerous, accounting for some seventy-five percent of all known hyperaccumulators. However, our understanding of the molecular basis of the physiological processes underpinning Ni hyperaccumulation has lagged behind that of Zn and Cd hyperaccumulation, in large part due to a lack of genomic resources for Ni hyperaccumulators. The advent of RNA-Seq technology, which allows both transcriptome assembly and profiling of global gene expression without the need for a reference genome, has offered a new route for the analysis of Ni hyperaccumulators, and several such studies have recently been reported. Here we review the current state of our understanding of the molecular basis of Ni hyperaccumulation in plants, with an emphasis on insights gained from recent RNA-Seq experiments, highlight commonalities and differences between Ni hyperaccumulators, and suggest potential future avenues of research in this field.

Effects of the interactions between nickel and other trace metals on their accumulation in the hyperaccumulator Noccaea caerulescens

Nickel hyperaccumulators are a group of particular plant species which possess high efficient Ni uptake transport systems in their roots, the mechanisms of which however are far from being understood. Study on the interactions between nickel and other trace metals during root absorption process can provide insights into the Ni uptake patterns. Therefore, this work used model hyperaccumulator species Noccaea caerulescens to comprehensively assess the interaction between Ni and other trace metals (Zn, Fe, Mn, Cu and Co) during the root uptake process. Plants were grown in nutrient solutions containing different concentrations of Ni, Zn, Fe and Co, and the metal concentrations in roots and shoots were determined. Furthermore, the gene expression levels of Zn and Fe transporters were determined by quantitative real-time reverse transcriptase PCR. Results showed that Ni uptake could be severely suppressed by the presence of Zn or Co, while greatly stimulated by Zn or Fe deficiency. Iron uptake could be also stimulated by Ni exposure. In addition, the expression levels of the Zn transporter ZIP10 and the Fe transporter IRT1 were increased after Ni exposure. It could be concluded that the great competition between Ni and the non-essential element Co indicates that Ni uptake is mainly controlled by non-specific transporters. Nickel may be mainly taken up via Zn transporters, while Fe transporters also play a role for Ni absorption.

Community diversity and potential functions of rhizosphere-associated bacteria of nickel hyperaccumulators found in Albania

Ultramafic (i.e. serpentine) soils are widespread in the Balkans and particularly in Albania. They account for a large part of plant endemism in that region and host several hyperaccumulator species, which are characterized by leaf nickel concentrations frequently above 1%. This rich nickel hyperaccumulating flora could serve as candidate to be used in phytoextraction and agromining. Despite recent interest in metal hyperaccumulating plants and agromining, very few studies have investigated the bacterial diversity and the influence of environmental factors on microbial gene profiles in the rhizosphere of hyperaccumulator plants growing on ultramafic soils. Because rhizospheric bacteria could be crucial to the success of phytoremediation, we studied a total of 48 nickel-hyperaccumulating plants which were sampled from four species that are widespread in Albania: Noccaea ochroleuca, Odontarrhena smolikana, O. rigida and O. chalcidica. All samples were taken from the ultramafic regions of Librazhd and Pogradec in eastern Albania in October 2015. Our study shows that Proteobacteria, Actinobacteria and Acidobacteria dominated the soil bacterial communities. Of these three phyla, only Proteobacteria was relatively abundant. This study underlines the influence of soil Cation Exchange Capacity on the bacterial community's diversity and structure. Based on the predicted metagenomes, the genes belonging to amino acid, lipid and carbohydrate metabolisms were identified as major gene families. Our study sheds some light on our understanding of how bacterial communities are structured within and affect the rhizosphere of hyperaccumulator plants from ultramafic soils in Albania.