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The molecular biology and genetics of the Ni-Cd-Zn hyperaccumulator Noccaea caerulescens has been extensively studied, but no information is yet available on Ni and Zn redistribution and mobilization during seed germination. Due to the different physiological functions of these elements, and their associated transporter pathways, we expected differential tissue distribution and different modes of translocation of Ni and Zn during germination. This study used synchrotron X-ray fluorescence tomography techniques as well as planar elemental X-ray imaging to elucidate elemental (re)distribution at various stages of the germination process in contrasting accessions of N. caerulescens. The results show that Ni and Zn are both located primarily in the cotyledons of the emerging seedlings and Ni is highest in the ultramafic accessions (up to 0.15 wt%), whereas Zn is highest in the calamine accession (up to 600 μg g-1). The distribution of Ni and Zn in seeds was very similar, and neither element was translocated during germination. The Fe maps were especially useful to obtain spatial reference within the seeds, as it clearly marked the vasculature. This study shows how a multimodal combination of synchrotron techniques can be used to obtain powerful insights about the metal distribution in physically intact seeds and seedlings., (© The Author(s) 2022. Published by Oxford University Press.)

Nickel (Ni) is used in numerous industrial processes, with large amounts of Ni-rich industrial wastes produced, which are largely sent to landfill. Nickel recovery from waste materials that would otherwise be disposed is of particular interest. Nickel phytomining represents a new technology in which hyperaccumulator plants are cultivated on Ni-rich substrates for commercial metal recovery. The aim of this study was to investigate the possibility of Ni transfer from industrial waste into plant biomass, to support recovery processes from bio-ores. Different industrial galvanic sludges (containing 85-150 g kg -1 Ni) were converted into artificial substrates (i.e. technosols) and the Ni hyperaccumulator Odontarrhena chalcidica (formerly Alyssum murale) was cultivated on these Ni-rich matrices. A greenhouse pot experiment was conducted for three months including an ultramafic soil control and testing fertilized (NPK) and unfertilized replicates. The results showed that fertilization was effective in improving plant biomass for all the substrates and that O. chalcidica was capable of viably growing on technosols, producing a comparable biomass to O. chalcidica on the control (ultramafic soil). On all technosols, O. chalcidica achieved Ni shoot concentrations of more than >1000 mg Ni kg -1 and maximum Ni uptake was obtained from one of the technosols (26.8 g kg -1 Ni, unfertilized; 20.2 g kg -1 Ni, fertilized). Nickel accumulation from three of the technosols resulted to be comparable with the control ultramafic soil. This study demonstrated the feasibility of transferring Ni from toxic waste into the biomass of Odontarrhena chalcidica and that phytomining from galvanic sludge-derived technosols can provide similar Ni yields as from natural ultramafic soils., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Hyperaccumulator plants present the ideal model system for studying the physiological regulation of the essential (and potentially toxic) transition elements nickel and zinc. This study used synchrotron X-ray Fluorescence Microscopy (XFM) elemental imaging and spatially resolved X-ray Absorption Spectroscopy (XAS) to elucidate elemental localization and chemical speciation of nickel and zinc in the hyperaccumulators Noccaea tymphaea and Bornmuellera emarginata (synonym Leptoplax emarginata). The results show that in the leaves of N. tymphaea nickel and zinc have contrasting localization, and whereas nickel is present in vacuoles of epidermal cells, zinc occurs mainly in the mesophyll cells. In the seeds Ni and Zn are similarly localized and strongly enriched in the cotyledons in N. tymphaea. Nickel is strongly enriched in the tip of the radicle of B. emarginata. Noccaea tymphaea has an Fe-rich provascular strand network in the cotyledons of the seed. The chemical speciation of Ni in the seeds of N. tymphaea is unequivocally associated with carboxylic acids, whereas Zn is present as the phytate complex. The spatially resolved spectroscopy did not reveal any spatial variation in chemical speciation of Ni and Zn within the N. tymphaea seed. The dissimilar ecophysiological behaviour of Ni and Zn in N. tymphaea and B. emarginata raises questions about the evolution of hyperaccumulation in these species.

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., (Copyright © 2018. Published by Elsevier B.V.)

Large ultramafic areas exist in Albania, which could be suitable for phytomining with native Alyssum murale. We undertook a five-year field experiment on an ultramafic Vertisol, aimed at optimizing a low-cost Ni-phytoextraction crop of A. murale which is adapted to the Balkans. The following aspects were studied on 18-m2 plots in natural conditions: the effect of (i) plant phenology and element distribution, (ii) plant nutrition and fertilization, (iii) plant cover and weed control and (iv), planting technique (natural cover vs. sown crop). The optimal harvest time was set at the mid-flowering stage when Ni concentration and biomass yield were highest. The application of N, P, and K fertilizers, and especially a split 100-kg ha(-1) N application, increased the density of A. murale against all other species. It significantly increased shoot yield, without reducing Ni concentration. In natural stands, the control of graminaceous weeds required the use of an anti-monocots herbicide. However, after the optimization of fertilization and harvest time, weed control procured little benefit. Finally, cropping sown A. murale was more efficient than enhancing native stands and gave higher biomass and phytoextraction yields; biomass yields progressively improved from 0.3 to 9.0 t ha(-1) and phytoextracted Ni increased from 1.7 to 105 kg ha(-1).

Until now, there has been little data on the isotope fractionation of nickel (Ni) in higher plants and how this can be affected by plant Ni and zinc (Zn) homeostasis. A hydroponic cultivation was conducted to investigate the isotope fractionation of Ni and Zn during plant uptake and translocation processes. The nonaccumulator Thlaspi arvense, the Ni hyperaccumulator Alyssum murale and the Ni and Zn hyperaccumulator Noccaea caerulescens were grown in low (2 μM) and high (50 μM) Ni and Zn solutions. Results showed that plants were inclined to absorb light Ni isotopes, presumably due to the functioning of low-affinity transport systems across root cell membrane. The Ni isotope fractionation between plant and solution was greater in the hyperaccumulators grown in low Zn treatments (Δ(60)Ni(plant-solution) = -0.90 to -0.63‰) than that in the nonaccumulator T. arvense (Δ(60)Ni(plant-solution) = -0.21‰), thus indicating a greater permeability of the low-affinity transport system in hyperaccumulators. Light isotope enrichment of Zn was observed in most of the plants (Δ(66)Zn(plant-solution) = -0.23 to -0.10‰), but to a lesser extent than for Ni. The rapid uptake of Zn on the root surfaces caused concentration gradients, which induced ion diffusion in the rhizosphere and could result in light Zn isotope enrichment in the hyperaccumulator N. caerulescens. In high Zn treatment, Zn could compete with Ni during the uptake process, which reduced Ni concentration in plants and decreased the extent of Ni isotope fractionation (Δ(60)Ni(plant-solution) = -0.11 to -0.07‰), indicating that plants might take up Ni through a low-affinity transport system of Zn. We propose that isotope composition analysis for transition elements could become an empirical tool to study plant physiological processes.

Ni recovery from serpentine soils by phytomining has proved feasible. Phytomining involves the crop of hyperaccumulating plants with high Ni contents and the valorization of Ni by pyro or hydrometallurgical process. In order to evaluate the Ni content of different plants, we analyzed the organs of 14 hyperaccumulators from three genera: Alyssum, Leptoplax and Bornmuellera. The highest concentration was recorded in the leaves of Leptoplax (34.3 +/- 0.7 mg g(-1)DM). Additionally, we investigated biomass combustion which is the first step of the process we designed to obtain a nickel salt. We showed that temperature and duration were important parameters to ensure a good quality of ashes. At the bench scale, the best conditions were 550 degrees C and 3 h. In this way, we obtained ashes in which Ni could reach 20 wt%. Biomass ashes can be considered as a bio-ore for recovering metal value.

Ni phytoextraction processes need further understanding of the interactions between Ni availability in soils and its absorption by plant roots. The large metal uptake and root exudation by hyperaccumulator species could accelerate the weathering process of Ni-bearing phases in the rhizosphere. The aim of this work was to quantify the weathering of a Ni-bearing mineral phase in the rhizosphere of the Ni-hyperaccumulator Leptoplax emarginata. The studied mineral was chrysotile which was characterized by a low Ni solubility. Column experiments were performed to assess the effect of the Ni-hyperaccumulator L. emarginata and the contribution of rhizobacteria on the dissolution rate of chrysotile. Mineral weathering was monitored by measuring Ni and Mg transferred to leachates or plants throughout the experiment. Results showed that L. emarginata increased chrysotile dissolution by more than 2-fold . The hyperaccumulator L. emarginata accumulated 88% on average of total mobilized Ni. Inoculation with Ni-resistant bacteria in the rhizosphere of L. emarginata had no significant effect on chrysotile dissolution or plant accumulation of Ni in this context. Finally, after 15 weeks of culture, 1.65% of total Ni in the system was mobilized in the planted treatments compared with 0.03% in the unplanted treatments.

Solute active transport or exclusion by plants can be identified by the values of the Transpiration Stream Concentration Factor (TSCF=xylem:solution solute concentration ratio). The aim of this study was to estimate this parameter for Ni uptake by the Ni-hyperaccumulator Leptoplax emarginata or the Ni-excluder Triticum aestivum cultivar 'Fidel'. The Intact Plant TSCF for nickel (IPTSCF(Ni)) was calculated as the ratio between the nickel mass accumulation in the leaves and the nickel concentration in solution per volume of water transpired. Predominantly, Ni active transport occurred for L. emarginata, with IPTSCF(Ni) values of 4.7-7.2 and convective component proportions of the root Ni uptake flow of only 15-20% for a range of Ni concentrations in solutions of 2-16 µmol Ni l(-1), regardless of the growth period and the time of Ni uptake. Hyperaccumulator roots were permeable to both water and nickel (mean reflection coefficient for Ni, σ(Ni), of 0.06), which was mainly attributed to an absence of exodermis. Results provide a new view of the mechanisms of Ni hyperaccumulation. By contrast, the wheat excluder was characterized by an extremely low mean IPTSCF(Ni) value of 0.006, characterizing a predominantly Ni sequestration in roots. From a methodological viewpoint, the 'microscopic' TSCF(Ni), measured directly on excised plants was 2.4 times larger than its recommended 'macroscopic' IPTSCF(Ni) counterpart. Overall, IPTSCF and σ determined on intact transpiring plants appeared to be very useful biophysical parameters in the study of the mechanisms involved in metal uptake and accumulation by plants, and in their modelling.

Stable Zn isotope signatures offer a potential tool for tracing Zn uptake and transfer mechanisms within plant-soil systems. Zinc isotopic compositions were determined in the Zn hyperaccumulator Noccaea caerulescens collected at a Zn-contaminated site (Viviez), a serpentine site (Vosges), and a noncontaminated site (Sainte Eulalie) in France. Meanwhile, a Zn-tolerant plant ( Silene vulgaris ) was also collected at Viviez for comparison. While δ(66)Zn was substantially differentiated among N. caerulescens from the three localities, they all exhibited an enrichment in heavy Zn isotopes of 0.40-0.72‰ from soil to root, followed by a depletion in heavy Zn from root to shoot (-0.10 to -0.50‰). The enrichment of heavy Zn in roots is ascribed to the transport systems responsible for Zn absorption into root symplast and root-to-shoot translocation, while the depletion in heavy Zn in shoots is likely to be mediated by a diffusive process and an efficient translocation driven by energy-required transporters (e.g., NcHMA4). The mass balance yielded a bulk Zn isotopic composition between plant and soil (Δ(66)Zn(plant-soil)) of -0.01‰ to 0.63‰ in N. caerulescens , indicative of high- and/or low-affinity transport systems operating in the three ecotypes. In S. vulgaris , however, there was no significant isotope fractionation between whole plant and rhizosphere soil and between root and shoot, suggesting that this species appears to have a particular Zn homeostasis. We confirm that quantifying stable Zn isotopes is useful for understanding Zn accumulation mechanisms in plants.

A pot experiment was conducted to investigate the effect of nickel concentration on physiological characteristics of Alyssum murale when grown in a soil mixed with sewage sludge (at the rate of 2.8%). Two types of sludge were used: agricultural sewage sludge (S1) and industrial sewage sludge with an increasing nickel concentration (S2, S3, and S4). Results showed that Ni in shoots was higher than Ni in roots. A. murale is able to concentrate up to 12730 mg/kg Ni in leaves. The highest dry matter yield was observed with plants grown with agricultural sewage sludge. An addition of S2 and S3 increased shoot biomass. However, application of S4 reduced 40% shoot dry weight as compared to the control Ni treatment did not affect all chlorophyll fluorescence parameters. The F(v)/F(m) ratio was stable between Ni treatments. Photosynthesis rate (A) increased with agricultural sewage sludge, but remained stable with variable Ni rates from the industrial sludge. The chlorophyll content increased with S1, S2 and S3 but it remains constant with S4 when compared to the control Therefore, high nickel concentration did not affect the function of the photosynthetic machine of A. murale.

We have investigated the accumulation of nickel in a hyperaccumulating plant from the Brassicacae family Leptoplax emarginata (Boiss.) O.E. Schulz. Two supplementary hyperaccumulating plants, which have been the subject of a high number of publications, Alyssum murale Waldst. & Kit and Thlaspi caerulescens J.&C. Presl, and a nonaccumulating species Aurinia saxatilis were also studied for reference. The plants were grown during 4 months in specific rhizoboxes with Ni-bearing minerals as a source of nickel. Nickel speciation was analyzed through X-ray absorption spectroscopy at Ni K-edge (X-ray absorption near edge spectroscopy and extended X-ray absorption fine structure spectroscopy) in the different parts of the plants (leaves, stems and roots) and compared with aqueous solutions containing different organo-Ni(II) complexes. Carboxylic acids (citrate, malate) appeared as the main ligands responsible of nickel transfer within those plants. Citrate was found as the predominant ligand for Ni in stems of Leptoplax and Alyssum, whereas in leaves of the three plants, malate appeared as the chelating organic acid of accumulated metal. Histidine could not be detected either in leaves, stems nor roots of any studied plant sample.

Leptoplax emarginata and Bornmuellera tymphaea are nickel hyperaccumulators of the Brassicaceae family endemic to serpentine soils in Greece. The aims of this work were to compare the growth and uptake behavior of these plants with the Ni hyperaccumulator species Thlaspi caerulescens and Alyssum murale, and to evaluate their effect on soil Ni availability. Plants were grown for 3 mo on three soils that differ in Ni availability. Ni availability in soils was measuredby isotopic exchange kinetics and DTPA-TEA extractions. Results showed that L. emarginata produced significantly more biomass than other plants. On the serpentine soil, B. tymphaea showed the highest Ni concentration in shoots. However, Niphytoextraction on the three soils was maximal with L. emarginata. The high initial Ni availability of soil Serp (470.5 mg kg(-1)) was the main explanation for the high Ni concentrations measured in plant shoots grown on this soil, compared to those grown on soils Calc and Silt A. murale was the least efficient in reducing Ni availability on the serpentine soil L. emarginata appeared as the most efficient species for Ni phytoextraction and decrease of the Ni available pool.

The nickel hyperaccumulator Bornmuellera tymphaea has been evaluated for use in agromining in Greece with very promising results. This study examines propagation of this species based on cuttings, which provides rapid multiplication and clonal propagation of genotypes with useful genetic traits and offers obvious advantages over propagation by seed. Apical cuttings, 10 cm long, obtained from plants in the field, were propagated on mist benches under greenhouse conditions. Four treatments based on the rooting medium were evaluated: (a) perlite, (b) perlite + peat 1:1 (v/v), (c) ultramafic soil, and (d) ultramafic soil + perlite 1:1 (v/v). Completely randomized blocks were used in a 4 × 2 factorial design, using 4 different propagation rooting media and 2 levels of auxins (with or without the use of indole‐3‐butyric acid [IBA]), with 3 replicates of 28 cuttings. The response to perlite medium was best in terms of rooting: 75% without IBA and 71.33% with IBA, and the dry weight of roots was 0.18 g and of shoots 0.34 g. Perlite + peat 1:1 (v/v) medium also resulted in rooting in more than 55% of cuttings whereas cuttings grown in ultramafic soil showed very low rooting (<27%). Based on the findings, B. tymphaea has an aptitude for vegetative propagation by cuttings with survival values and rooting greater than 70%. Perlite combined with the application of auxin (ΙΒΑ) is recommended. [ABSTRACT FROM AUTHOR]

The Greek endemic Bornmuellera emarginata, known for its Ni hyperaccumulation capacity, is a promising candidate species for use in agromining. At the same time, the installation of photovoltaic systems on agricultural land has recently been increasing. Field observations that populations of this species usually occupy shaded places under shrubs or trees led us to study the effect of different shading levels on the growth of B. emarginata, with a view to combine the cultivation of the species with photovoltaic panels in the same area. The experiment was conducted under greenhouse conditions and lasted 210 days after sowing (DAS). Seeds were sown directly into 2‐L pots filled with ultramafic soil collected from the field. Three plant samplings (81, 140, and 210 DAS) were conducted during the experiment to determine a set of morphological and physiological parameters of the species (fresh and dry plant biomass, plant height, number of lateral shoots, total number of leaves, proline and chlorophyll concentrations) alongside with its nickel yield at different levels of shading (0%, 20%, 35%, 60%, and 90%). Growth of B. emarginata was only significantly affected at the highest shading level. Nickel yield of the species increased over time and was maximized at the end of the experiment. The highest nickel yield per plant was observed at the intermediate shading level (35%). Bornmuellera emarginata can be considered as a candidate nickel hyperaccumulator for cultivation in ultramafic or nickel‐polluted areas under photovoltaic panels with corresponding shading, providing important environmental benefits. [ABSTRACT FROM AUTHOR]

X-ray Fluorescence Spectroscopy (XRF) is a non-destructive analytical technique that has recently been developed as a tool for high throughput screening for the discovery of trace element hyperaccumulator species in herbarium collections. Herbarium XRF analysis was undertaken of representatives of the Brassicaceae family in the two herbaria in Belgrade (Serbia). This was coupled to field collection and chemical analysis of new samples of Cardamine waldsteinii. The data revealed a potentially new zinc hyperaccumulator species, C. waldsteinii, with 3050 mg kg−1 Zn in its leaves. This compares to 2210 mg kg−1 in the leaves of this species collected from the field. These values are below the notional hyperaccumulation threshold for zinc but are nevertheless unusual. The remarkably high accumulation factor for zinc are suggestive of the exceptional potential of C. waldsteinii for zinc uptake, even in non-metalliferous, zinc-deficient soils. We recommend following up this study with experimental investigations on C. waldsteinii to elucidate its ecophysiology and its zinc tolerance and accumulation characteristics. [ABSTRACT FROM AUTHOR]

Aims: Based on a recent taxonomic revision of the genus Odontarrhena in Albania, our work aimed at comparing the taxa and populations of this genus in terms of nickel accumulation and levels of other metals in relation to their concentrations in the soil at distinct Albanian ultramafic outcrops. Methods: Several populations of different Odontarrhena species were sampled from 20 different outcrops across the entire Albanian territory. Concentrations of Ni, Co, Cr, Mg, Ca, K, Fe and Mn were determined in soil, roots and shoots of all specimens sampled. Results: Ultramafic soils across Albania showed variable metal composition, with significant differences between sites. Large differences in mineral element concentrations were also found between the plants which were not related to species identity or geographical distribution of their populations. Conclusions: Shoot Ni concentrations in Albanian Odontarrhena taxa depend on soil Ni concentrations but not on species identity. For O. chalcidica, the more widely distributed species, the environmental fingerprint for shoot metal composition was found not only for Ni, but also for Ca and Mg, the latter showing a positive linear relationship between soil and plant concentrations. [ABSTRACT FROM AUTHOR]

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 μM Ni or Zn, and then used for phloem sap extraction by means of the EDTA-stimulated exudation method. In the second experiment, Ni 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 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. [ABSTRACT FROM AUTHOR]

Albanian taxa and populations of the genus Odontarrhena are most promising candidates for research on metal tolerance and Ni-agromining, but their genetic structure remains unknown. We investigated phylogenetic relationships and genetic differentiation in relation to distribution and ploidy of the taxa, anthropic site disturbance, elevation, soil type, and trace metals at each population site. After performing DNA sequencing of selected accessions, we applied DNA-fingerprinting to analyze the genetic structure of 32 populations from ultramafic and non-ultramafic outcrops across Albania. Low sequence divergence resulted in poorly resolved phylograms, but supported affinity between the two diploid serpentine endemics O. moravensis and O. rigida. Analysis of molecular variance (AMOVA) revealed significant population differentiation, but no isolation by distance. Among-population variation was higher in polyploids than in diploids, in which genetic distances were lower. Genetic admixing at population and individual level occurred especially in the polyploids O. chalcidica, O. decipiens, and O. smolikana. Admixing increased with site disturbance. Outlier loci were higher in serpentine populations but decreased along altitude with lower drought and heat stress. Genetic variability gained by gene flow and hybridization at contact zones with "resident" species of primary ultramafic habitats promoted expansion of the tetraploid O. chalcidica across anthropogenic sites. [ABSTRACT FROM AUTHOR]

Abstract: To characterize the complete Ni root sink over the short term, the contributions of apoplast and symplast to the metal root uptake were measured for both maize and the Ni hyperaccumulator Leptoplax emarginata, using a physico-chemical fractionation of root Ni after exposure. The apoplastic uptake accounted for 81.3–88.0% and for 90.6–95.5% of the total root uptake of maize and leptoplax, respectively. This could be modelled using exchange isotherms, such as the Langmuir or Freundlich isotherms. Root cell walls of both plant species showed similar affinity to Ni, although the hyperaccumulator had a higher apoplastic uptake. For maize as for leptoplax, the symplastic influx fitted a Michaelis–Menten function with similar K m and I max values, although the two plant species have contrasting abilities to accumulate Ni. This suggests that symplastic uptake is not the main factor of hyperaccumulation and that the Ni transport system might be similar in these two plants species. More research is needed to understand how the high level of Ni adsorption on root apoplast might affect the overall Ni uptake. [Copyright &y& Elsevier]