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Aims: Dicranopteris linearis is a rare earth element (REE), aluminium (Al) and silicon (Si) hyperaccumulator plant which occurs in southern China. To date, there have been no studies on the variation in elemental accumulation among populations of Dicranopteris linearis occurring on REE-enriched and non-REE enriched soils. Methods: In total 43 Dicranopteris linearis individuals and the corresponding rhizosphere soils from 7 populations in southern China were sampled and the plant material and soil composition were chemically analysed for REE and other elements. Results: Dicranopteris linearis populations from REE-enriched soils and non-REE enriched soils both have the ability to accumulate high concentrations of REE (> 1000 mg kg− 1), Al (> 1000 mg kg− 1) and Si (> 10 000 mg kg− 1). The ratios of light REEs (LREEs) to heavy REEs (HREEs) in Dicranopteris linearis pinnae are consistently > 1, regardless of their ratios in the corresponding soils. Concentrations of REE in Dicranopteris linearis pinnae vary significantly among populations and have positive correlations with total and extractable soil REE concentrations. The plant Al and Si concentrations in Dicranopteris linearis populations are uncoupled from variations in soil Al and Si concentrations. The plant REE concentrations have no obvious relationships with the prevailing Al concentrations, but are positively correlated with plant Si and Mn, and negatively correlated with soil extractable P concentrations. Conclusions: The hyperaccumulation of REEs (LREE > HREE), Al and Si in non-REE enriched soils is a species-wide trait. The accumulation of REEs is highly variable and associated with soil REE concentrations, nutrients status, and other soil properties.

Aims: The Australian native hemi-aquatic herb C. helmsii (Crassulaceae) is a copper (Cu) accumulator, tolerant to a wide range of climatic conditions and able to concentrate >9000 μg Cu g−1 in its living tissues. These characteristics suggest practical potential for decontamination of Cu-polluted mine waste waters using a phyto-extraction approach. The aim of this study was to investigate Cu uptake in C. helmsii at different Cu2+ concentrations in solution and to test for the effect of pH on the concentration of free Cu2+ and Cu accumulation in this species. Methods: Different solutions were tested in acid (pH 4.0) and mildly acidic (pH 5.8) conditions at five different Cu2+ concentrations (ranging from 0 to 5 mg Cu L−1). The concentration of free Cu2+ was evaluated using Geochem-EZ software. The distribution of Cu in C. helmsii tissues was examined with micro-X-ray Fluorescence (μXRF) elemental mapping and with Scanning Electron Microscopy with Energy Dispersive X-ray Spectroscopy (SEM-EDS). Results: The highest shoot Cu concentration in C. helmsii was 5870 μg Cu g−1 in the 5 mg Cu L−1 treatment. Copper bioaccumulation was positively correlated with the Cu2+ treatment in the different solutions. The μXRF and SEM-EDS analysis revealed that roots accumulated higher Cu concentrations than the shoots. The concentration of free Cu2+ was shown to be dependent upon the type of Fe-chelator (DTPA, EDDHA, HBED) used in the solution due to the displacement of Fe3+ and complexation of Cu2+. Conclusions: Crassula helmsii is highly tolerant to Cu2+ in solution at acidic and mildly acidic pH, and able to accumulate high Cu concentrations in its tissue. This confirms its potential for applications in Cu phyto-extraction of acidic mine effluent waters high in Cu2+.

International audience; Background and Aims: In tropical ultramafic soils, potassium (K) is typically the most growth limiting nutrient. However, tropical nickel (Ni) hyperaccumulator plants, including Phyllanthus rufuschaneyi and Rinorea cf. bengalensis (which are ‘metal crops’ used in agromining) from Malaysia, have unusually high K shoot accumulation compared to other species, despite naturally growing on severely K-impoverished ultramafic soils. This study aimed to establish the response to soil K availability in relation to uptake of K and other elements in the roots and shoots of P. rufuschaneyi and R. cf. bengalensis.Methodology: We undertook an experiment in which soluble K was dosed to ultramafic soil in pots with P. rufuschaneyi and R. cf. bengalensis in Sabah (Malaysia).Results: The results show that root K concentrations increased markedly as the soil K availability increased by 35-fold, whilst the corresponding effect on K accumulation in the shoots of P. rufuschaneyi and R. cf. bengalensis was not significantly different in relation to soil K dosing. Observed divergent responses between root and shoot K accumulation in these species suggests a separate genetic control of K uptake and xylem loading in P. rufuschaneyi and R. cf. bengalensis.Conclusion: The tight control of root-to-shoot K translocation and constrained K accumulation in shoots under a soil K gradient is likely an adaptive mechanism to the evolution of these species to grow in highly nutrient-impoverished ultramafic soils. This study provides information that will be useful for better nutrient management of tropical Ni metal farms that use K-efficient Ni ‘metal crops’.

AimsMetal hyperaccumulation is a rare phenomenon described for an increasing number of plant taxa. In this study we investigated the root growth responses of the well-known nickel, zinc, cadmium hyperaccumulatorNoccaea caerulescensand of the metal tolerant (non-accumulator)Stellaria media, in order to observe root foraging vs avoidance responses to nickel. MethodsTo allow for observations of root growth and foraging preferences, two accessions ofNoccaea caerulescensand two accessions ofStellaria mediaorginating from high nickel and low nickel habitats were grown in rhizotrons with localized nickel enrichment.ResultsThe root density in the control and nickel-enriched soil areas in the rhizotrons with differentN. caerulescensaccessions had distinct responses: moderate nickel avoidance was recorded for the non-nickel accession, while a clear foraging response was observed inN. caerulescensfrom the nickel accession. In contrast, nickel rooting avoidance was observed for bothS. mediaaccessions and was more pronounced in the non-nickel accession.ConclusionsThis study shows thatN. caerulescensoriginating from different accessions responded differently to soil nickel enrichment, with the nickel accession ofN. caerulescensactively foraging for nickel, suggesting a physiological adaptation and demand for this metal. In contrast, a clear nickel avoidance response by a metal tolerant species, S. media, was observed in this study, a phenomenon which has not been previously described; this suggests that root avoidance responses might play a role in the adaptation of metal tolerant species to Ni-rich soils.

Background: Phytoextraction is an in situ technique that can be applied to minerals and mining wastes using hyperaccumulator plants to purposely bio-concentrate high levels of metals or metalloids into their shoots in order to remove them from the substrate, while achieving monetary gain. Phytoextraction can be applied to a limited number of elements depending on the existence of hyperaccumulator plants with suitable characteristics. Although phytoextraction has been trialled in experimental settings, it requires testing at field scale to assess commercial broad-scale potential. Scope: The novelty and purported environmental benefits of phytoextraction have attracted substantial scientific inquiry. The main limitation of phytoextraction with hyperaccumulators is the number of suitable plants with a high accumulation capacity for a target element. We outline the main considerations for applying phytoextraction using selected elemental case studies in which key characteristics of the element, hyperaccumulation and economic considerations are evaluated. Conclusions: The metals cobalt, cadmium, thallium and rhenium and the metalloids arsenic and selenium are present in many types of minerals wastes, especially base metal mining tailings, at concentrations amenable for economic phytoextraction. Phytoextraction should focus on the most toxic elements (arsenic, cadmium, and thallium) or especially valuable elements (selenium, cobalt, and rhenium). The value proposition is in the clean-up of contaminated land in the case of toxic elements, whereas it is in the ‘bio-ore’ generated by the process in the case of valuable elements.

Aims: 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. Methods : 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. Results: 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. Conclusions: 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.

International audience; Background and aims: Pioneer plants may improve the ecological restoration of degraded ultramafic areas by plant-soil interaction processes. In this study, we assess the effect of the pioneer actinorhizal tree C. terminale (Casuarinaceae) on the recovery of plant communities and soil functions on degraded tropical ultramafic sites.Methods: Soil and plant samples were collected along a tree-age gradient in two degraded ultramafic sites in Sabah (Northern Borneo, Malaysia): a Technosol and a Leptosol. Chemical composition of plants and soils, and biological activity of soils were assessed at both sites. Plant colonisation was assessed by plot vegetation surveys.Results: An improvement in soil fertility parameters (pH reduction from 8.5 to 6.8, an increase in the concentrations of several nutrients and enhanced soil enzyme activities) was observed along the C. terminale age gradient. However, plant cover and diversity was only improved around mature trees at the site that was not impacted by mining.Conclusion: C. terminale promotes the recovery of several soil functions, mainly related to the storage and recycling of N, P, K, S. Besides plant-soil feedback, other environmental factors (i.e. exposition to sunlight, drought) may play an important role on revegetation of ultramafic soils.

Aims: Dicranopteris linearis is a fern that accumulates unusually high concentrations (up to 0.3% dry weight) of rare earth elements (REEs) in China. Previously, we reported that D. linearis accumulates high concentrations of aluminium (Al) and silicon (Si) in the fronds, but the interactions between these elements and REEs were unknown. Methods: In this study, a range of analytical techniques, including chemical extractions, Scanning-Electron Microscopy with Energy-Dispersive Spectroscopy (SEM-EDS) and Laser Ablation Inductively Coupled Plasma Mass Spectrometry (LA-ICP-MS) were used to study the association of REEs with Al and Si. Results: The results show that D. linearis accumulates high concentrations of REEs (up to 3830 mg kg−1), Al (up to 9660 mg kg−1) and Si (up to 20,300 mg kg−1), with concentrations increasing with age and frond order of pinna. The extraction patterns suggest the existence of REEs-Si and Al-Si complexes. The SEM-EDS analysis confirmed the existence of phytoliths (Al) deposits in the protoplast and apoplast of the pinna cells. The upper epidermis of the pinnule and the pericycle of the midvein are more concentrated in phytoliths (Al) particles. The LA-ICP-MS analysis revealed that REEs and Al are preferentially compartmentalized within bio-inactive tissues of the pinnules, e.g. necrotic tissues (REEs) and in the margins (Al). Conclusions: Co-deposition of Si with REEs and Al may be a mechanism for dealing with the high concentrations of REEs and Al in D. linearis fronds.

International audience; Aim 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. Methods 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. Results 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. Conclusions 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.

Background: Nickel (Ni) phytomining operations cultivate hyperaccumulator plants (‘metal crops’) on Ni-rich (ultramafic) soils, followed by harvesting and incineration of the biomass to produce a high-grade ‘bio-ore’ from which Ni metal or pure Ni salts are recovered. Scope: This review examines the current status, progress and challenges in the development of Ni phytomining agronomy since the first field trial over two decades ago. To date, the agronomy of less than 10 species has been tested, while most research focussed on Alyssum murale and A. corsicum. Nickel phytomining trials have so far been undertaken in Albania, Canada, France, Italy, New Zealand, Spain and USA using ultramafic or Ni-contaminated soils with 0.05–1 % total Ni. Conclusions: N, P and K fertilisation significantly increases the biomass of Ni hyperaccumulator plants, and causes negligible dilution in shoot Ni concentration. Organic matter additions have pronounced positive effects on the biomass of Ni hyperaccumulator plants, but may reduce shoot Ni concentration. Soil pH adjustments, S additions, N fertilisation, and bacterial inoculation generally increase Ni phytoavailability, and consequently, Ni yield in ‘metal crops’. Calcium soil amendments are necessary because substantial amounts of Ca are removed through the harvesting of ‘bio-ore’. Organic amendments generally improve the physical properties of ultramafic soil, and soil moisture has a pronounced positive effect on Ni yield. Repeated ‘metal crop’ harvesting depletes soil phytoavailable Ni, but also promotes transfer of non-labile soil Ni to phytoavailable forms. Traditional chemical soil extractants used to estimate phytoavailability of trace elements are of limited use to predict Ni phytoavailability to ‘metal crop’ species and hence Ni uptake.

Background and aims: Kinabalu Park is the world’s most species-rich hotspot with over 5000 plant species recorded for an area 1200 km2. The aim of this study was to characterise the vegetation on ultramafic edaphic ‘islands’ in relation to soil chemistry and elevation. Methods: In total 87 non-permanent vegetation plots were established covering 12 ultramafic edaphic ‘islands’ from 474 to 2950 m asl in which 2854 plant species in 742 genera and 188 families were recorded from 14 662 collections. Results: The results show that plant diversity decreases with elevation, but a mid-elevation (circum 1500 m asl) ‘hump’ occurs for some plant groups (orchids, pteridophytes) as a result of the presence of cloud forests. Six main vegetation classes with associated soil types were discerned: (i) Sub-Alpine Scrub; and (ii) Graminoid Scrub, both associated with Hypermagnesic Cambisols (‘hypermagnesian soils’); (iii) Montane Cloud Forest, associated with Cambisols often with accumulation of humus; (iv) Mixed Dipterocarp Forest, associated with deep Ferralsols (‘laterites’); (v) Pioneer Casuarina Scrub; (vi) Mature Mixed Casuarina Forest, both associated with Hypermagnesic Leptosols. Conclusions: We hypothesised that ‘adverse’ soil chemistry would exacerbate vegetation stunting, and the results confirmed that stunted vegetation and elevational floristic compression occurs on chemically adverse soils (mainly hypermagnesian soils). However, no clear correlation with plant diversity was found, as some of the most ‘adverse’ soils on the summit of Mount Tambuyukon had up to 132 species per 250 m2.

International audience; Aims This study evaluated the effect of phloem translocation on Ni accumulation in the hyperaccumulator Noccaea caerulescens. Methods The first experiment assessed the metal and organic compound concentrations in phloem sap. Leaves were cut from plants grown in nutrient solutions added with 100 mu M Ni or Zn, and then used for phloem sap extraction by means of the EDTA-stimulated exudation method. In the second experiment, Ni-61(2+) was applied to old leaves as a foliar spray to assess bidirectional movement of Ni in phloem. Results In the first experiment, enrichment of Ni or Zn was found in phloem exudates, indicating high Ni and Zn phloem loading and translocation capacity in N. caerulescens. Amino acids, e.g. nicotianamine and histidine, were present at low concentrations in exudates, which are insufficient for Ni and Zn chelation. On the contrary, concentrations of organic acids, especially malate, were high in all treatments, which may be involved in Ni and Zn chelation in phloem. The second experiment showed that 89 % of Ni-61 exported from old leaves was translocated upward to young leaves, whereas only 11% moved downward to roots. Conclusions Phloem sap of N. caerulescens is enriched in Ni and malate, the majority of which moves upward to young tissues. Phloem translocation may play an important role for Ni accumulation in young leaves of N. caerulescens.

International audience; Background and aims Noccaea caerulescens is a model plant for the understanding of trace metal accumulation and a source of cultivars for phytoextraction. The aim of this study was to investigate natural variation for trace metal accumulation, major nutrient uptake and growth parameters in 22 populations. The correlations among these traits were particularly examined to better understand the eco-physiology and the phytoextraction potential of the species. Methods Populations from three edaphic groups, i.e. calamine (CAL), serpentine (SERP) and non metalliferous (NMET) sites were grown in hydroponics for seven weeks at moderate trace metal exposure. Growth indicators, element contents and correlations between these variables were compared. Results All the phenotypic characteristics showed a wide variability among groups and populations. The SERP populations showed a smaller plant size, higher cation contents and strong correlations between all element concentrations. NMET populations did not differ in plant size from the CAL ones, but had higher Zn and Ni contents. The CAL populations showed higher Cd and Mn accumulations and lower Ca contents. The trade-off between biomass production and Cd, Ni and Zn accumulation was high in SERP populations and low in the CAL and NMET ones. Conclusions N. caerulescens is a genetically diverse species, showing specific features depending on the group and the population. These features may reflect the wide adaptive capacities of the species, and also reveal promising potential for phytoextraction of Cd, Ni and Zn.

Aims The primary aim of this study was to investigate the impact of active nickel and cadmium transport, transpiration and shoot biomass production on Ni and Cd accumulation in the leaves of the Ni-hyperaccumulator Leptoplax emarginata. A secondary objective was to observe the effects of various concentrations of nickel and cadmium in solutions on the plant growth and ecophysiological characteristics of these plants. Finally, the study sought to identify possible nickel and cadmium concentration gradients in solution as a function of the root distance. Methods The Intact Plant Transpiration Stream Concentration Factor (TSCF=xylem/solution solute concentration ratio) was determined for both Ni and Cd and for the selected intact transpiring Ni-hyperaccumulator Leptoplax emarginata, cultivated on two contrasting fertilized and Ni-Cd-contaminated sandy porous media (rhizotrons with central root compartments, linked to Mariotte tubes operated at -1 kPa). IPTSCF(Ni) and IPTSCF(Cd) were calculated as the ratios between the hyperaccumulator plant's nickel or cadmium mass in the leaves and the nickel or cadmium concentration in solution by the volume of water transpired during the period of culture. Plant growth characteristics and gas exchanges were also recorded. Results IPTSCF values were much greater than 1 (IPTSCF(Ni) = 5.2 +/- 0.9 and IPTSCF(Cd) = 4.4 +/- 0.6) whatever the amount of available Ni and Cd. This characterized a predominantly active plant metal uptake. Moreover, biological regulation was reported: plant growth and transpiration were significantly lower for hyperaccumulator plants cultivated in sand which was rich in available Ni and Cd, than for hyperaccumulator plants cultivated in topsoil, poor in available Ni and Cd. In the soil rhizosphere, capillary flow was related to transpiration and a depletion pattern was developed for Ni and sometimes for Cd. Conclusions Overall, the Intact Plant Transpiration Stream Concentration Factor appeared to be a relevant metal bioconcentration factor taking into account the predominant type of metal transport from roots to leaves, plant growth and transpiration coupling and metal availability. IPTSCF(Ni) and IPTSCF(Cd) values were much greater than 1 and similar whatever the amount of available Ni and Cd. This characterized a predominantly active plant combining Ni and Cd uptake and biological regulations dependent of the Ni and Cd concentrations in solution.

A glasshouse experiment was set up to compare processes and organisms in two soils planted with genetically modified (GM) herbicide tolerant (HT) maize treated with appropriate herbicides. This was part of a wider project (ECOGEN) looking at the consequences of GM cropping systems on soil biology using a tiered approach at laboratory, glasshouse and field scales. Soil for the experiment was taken from field sites where the same maize cultivars were grown to allow comparison between results under glasshouse and field conditions. The maize cultivars T25 (GM HT glufosinate-ammonium tolerant), Orient (non HT near isogenic control for T25) and Monumental (a conventional, non HT variety) were grown in contrasting sandy loam and clay loam soils, half were sprayed with the appropriate herbicide as used in the field and soil samples were taken at the five-leaf and flowering plant growth stage. The main effects on all measured parameters were those of soil type and plant growth stage, with four categories of subsequent interaction: (1) there were no effects of herbicide on plant growth or soil microarthropods: (2) the maize cultivar (but not the GM HT trait) had effects on the decomposition of cotton strips and the nematode community; (3) herbicide application in general altered the community level physiological profile of the microbial community and reduced both soil basal respiration and the abundance of protozoa; and (4) the specific application of glufosinate-ammonium to T25 maize altered soil microbial community structure measured by ester linked fatty acids. The results from this glasshouse experiment support the findings from the field that there are effects of herbicide application on the soil microbial and meso-faunal community but that, compared to other standard agricultural practices, the differences are relatively small.

Our purpose was to quantify the effect of rhizosphere processes on the availability of soil cadmium and zinc to various plant species. E values for Cd and Zn were measured on two contaminated soils differing mainly in their pH (6.2 and 8.1). L values were measured for six plant species with contrasting metal uptakes. The difference between E and L values quantifies the mobilization of the element which was non-phyto-available prior to cultivation. For Zn, L values were 1.2 to 5.6 times greater than the E values, depending on the plant species. The increase in Zn availability was greater in the basic soil. For Cd, L values in the basic soil were 1.1 to 2 times greater than the E value. In the slightly acid soil, plants did not enhance the cadmium availability. The mobilization of non-labile metal could be due to exudates from roots or microflora, phytosiderophores being responsible for the highest mobilization. The lower availability of Fe or Zn could explain the greater mobilization in the basic soil.