A phytogeochemical survey of the flora of ultramafic and adjacent normal soils in North Morocco

Variation in plant elemental composition (Ni, Ca, Mg, Mg/Ca ratio) in relation to soil composition was investigated in a poorly studied ultramafic area in the north of Morocco. A total of 142 leaf samples representing 36 species from 9 sites (5 ultramafic and 4 normal soils from adjacent areas) were analysed. The soil was richer in Mg and Ni and had a higher Mg/Ca ratio in the ultramafic sites than in the control sites, and these differences were qualitatively reflected in the average mineral composition of the plants. However, there were considerable differences in mineral composition among species within serpentinic sites, indicating that species with contrasting mineral nutrition strategies can cope with the mineral element imbalance characteristic of ultramafic soils. Particularly noteworthy was the finding that species with high requirements of Ca are not excluded from serpentinic soils. In view of their high responsiveness to soil nickel and magnesium concentration, Dittrichia viscosa and Lavandula dentata are proposed as bioindicators of these elements in the soil in the Rif area. By contrast, two local serpentine endemics, Halimium atriplicifolium and Notholaena marantae were excluders of nickel and magnesium. This revised version was published online in August 2006 with corrections to the Cover Date.     

Characterisation of arbuscular mycorrhizal fungi colonisation in cluster roots of shape Hakea verrucosa F. Muell (Proteaceae), and its effect on growth and nutrient acquisition in ultramafic soil

Ultramafic soils at Bandalup Hill (Western Australia) are characterised by high concentrations of Ni and low levels of P. Amongst the plant species that can sustain such hostile conditions, Hakea verrucosa F. Muell from a non-mycorrhizal family (Proteaceae) would be expected to rely on cluster roots to access P. However, the acidification of ultramafic soils by cluster roots might increase the dissolution of soil Ni, and therefore its availability to plants. Symbiosis with mycorrhizal fungi, on the other hand, might help to reduce the uptake of Ni by H. verrucosa. Therefore, the aim of this study was to investigate the mycorrhizal status of H. verrucosa, and assess any contribution from mycorrhizal fungi to its growth and nutrient status. Seedlings of H. verrucosa were first grown in undisturbed ultramafic soil cores from Bandalup Hill for 8 weeks to assess the presence of mycorrhizal fungi in their roots. In a second experiment, H. verrucosa seedlings were grown in the same ultramafic soil that was either steamed or left untreated. Seedlings were inoculated with an arbuscular mycorrhizal (AM) fungal consortium from Bandalup Hill. Fungal hyphae, vesicles, as well as intracellular arbuscules and hyphal coils were observed in the cluster roots of H. verrucosa in both experiments. In the first experiment, 57% of the root length was colonized by AM fungi. Seedlings had high (between 1.4 and 1.9) shoot to root ratios and their roots had very few root hairs, despite growing in P-deficient soil. Steaming of the ultramafic soil increased the growth of seedlings and their nutrient uptake. Inoculation with AM fungi reduced the seedling growth in steamed ultramafic soil; however, it increased their shoot P and K concentration and also the shoot K content. The shoot Ni concentration of seedlings was not affected by the presence of AM fungi.     

Natural variation of nickel,zinc and cadmium (hyper)accumulation in facultative serpentinophytes Noccaea kovatsii and N. praecox

Aims

Data on the variability of hyperaccumulation potential of the facultative serpentinophytes Noccaea kovatsii and N. praecox on different geological substrates are scarce. The aim of this study was to assess the accumulation potential of these two species from ultramafic and non-ultramafic substrates, with special emphasis on the hyperaccumulation of Ni, Zn and Cd.

Methods

Samples of plants and corresponding soils were collected from 16 sites covering a wide range of geochemistry. Elemental concentrations were determined in the roots, shoots and the rhizosphere soils. The pH, particle size distribution and content of organic matter were also determined in the soil samples.

Results

All ultramafic accessions of both species hyperaccumulated Ni with high intraspecific variability. Only one accession of N. kovatsii from a schist soil hyperaccumulated Zn, with also a high Cd accumulation. Accumulation and translocation of Ni in both species were much higher in the ultramafic accessions, whereas Zn accumulation and translocation was found in both ultramafic and non-ultramafic accessions.

Conclusions

Ni accumulation and translocation was restricted to ultramafic populations of both species, whereas it is a species-wide trait for Zn. This study provides new and comprehensive information on the natural variation of hyperaccumulation of Ni, Zn and Cd in N. kovatsii and N. praecox.

     

Nickel bioavailability in an ultramafic toposequence in the Vosges Mountains (France)

A serpentinised harzburgite outcrop located in the Vosges Mountains hosts a population of the Ni-hyperaccumulator Thlaspi caerulescens J. & C. Presl. A complete study was undertaken to relate the variability of Ni availability along the ultramafic toposequence to pedogenesis, soil mineralogy and functioning with X-Ray Diffraction, Transmission Electron Microscope observations coupled with Isotopic Exchange Kinetics and diethylenetriamine pentaacetic acid extraction of Ni. The soil profiles ranging from Dystric Cambisol to Hypermagnesic Hypereutric Cambisol were distributed unevenly along the toposequence probably due to geochemical variability of the bedrock and also complex quaternary erosion features. The richest soils were characterised by slight mineral weathering leading to Ni, Cr and Fe accumulation in the B horizons whereas the lowest saturated soils had very low-metal contents. Most soil minerals were inherited from the parent materials and there were only few traces of formation of secondary minerals. Primary minerals (e.g. serpentine, chlorite) contained low Ni concentrations (0.2%) whereas neoformed goethite, mainly in the B horizons of the richest soils, contained up to 4.3% Ni. Ni was probably sorbed onto amorphous Fe oxy−hydroxide particles (oxalate extraction) rather than incorporated within the crystal lattice of goethite. Ni availability in the B horizon of Hypereutric Cambisols was extremely high and so was the oxalate extractable Fe. At the toposequence level, there was a high level of Ni availability in the least weathered soils and a very low-availability level in the more intensively weathered soils (strongly acidic pH). Ni availability was unexpectedly positively correlated to pH and was controlled by soil mineralogy and Ni-bearing mineral phases. Ni hyperaccumulation (above 1,000 mg kg⁻¹) by native T. caerulescens was only reached in the Ni-rich soils as a consequence of the local edaphic factors. Ni uptake by T. caerulescens is strongly regulated by Ni availability in soils and therefore related to pedogenesis.     

Does soil determine the boundaries of monodominant rain forest with adjacent mixed rain forest and maquis on ultramafic soils in New Caledonia?

Aim To determine the soil characteristics of Nothofagus‐dominated rain forests in an ultramafic region (i.e. soils having high concentrations of metals including Mg, Fe and Ni), and whether soil characteristics may explain the location of monodominant rain forest in relation to adjacent mixed rain forest and maquis (shrub‐dominated vegetation). Location New Caledonia. Methods Soil characteristics were compared among six Nothofagus‐dominated rain forests from a range of altitudes and topographic positions. At four of these sites, comparisons were made with soils of adjacent mixed rain forest and maquis. Results Soil characteristics varied among the monodominant Nothofagus forests, largely due to differences between ultramafic soils and soils influenced by non‐ultramafic intrusions. The soils of all vegetation types had low concentrations of nutrients, particularly P, K and Ca (both total and extractable/exchangeable), and high total concentrations of Ni, Fe, Cr and Mn. There were significant differences between the rain forests and adjacent maquis in soil concentrations of several elements (N, P, Ca, Mg and Mn), more so in surface soils than at depth, but much of this pattern may be caused by effects of vegetation on the soil, rather than of soil on the vegetation. However, there were no significant differences in soil concentrations of any mineral elements between Nothofagus forest and adjacent mixed rain forest. Main conclusions We found no evidence for soil mediation of boundaries of Nothofagus rain forest with mixed rain forest, and little evidence for the boundaries of either forest type with maquis. We suggest that the local abrupt boundaries of these monodominant Nothofagus forests are directly related to temporal factors, such as time since the last wildfire and frequency of wildfire, and that disturbance is therefore a major causal factor in the occurrence of these forests.     

Arbuscular mycorrhizal fungi from New Caledonian ultramafic soils improve tolerance to nickel of endemic plant species

In order to improve knowledge about the role of arbuscular mycorrhizal fungi (AMF) in the tolerance to heavy metals in ultramafic soils, the present study investigated the influence of two Glomus etunicatum isolates from New Caledonian ultramafic maquis (shrubland), on nickel tolerance of a model plant species Sorghum vulgare, and of two ultramafic endemic plant species, Alphitonia neocaledonica and Cloezia artensis. In a first step, plants were grown in a greenhouse, on sand with defined concentrations of Ni, to appreciate the effects of the two isolates on the alleviation of Ni toxicity in controlled conditions. In a second step, the influence of the AMF on A. neocaledonica and C. artensis plants grown in a New Caledonian ultramafic soil rich in extractable nickel was investigated. Ni reduced mycorrhizal colonization and sporulation of the fungal isolates, but the symbionts increased plant growth and adaptation of endemic plant species to ultramafic conditions. One of the two G. etunicatum isolates showed a stronger positive effect on plant biomass and phosphorus uptake, and a greater reduction in toxicity symptoms and Ni concentration in roots and shoots. The symbionts seemed to act as a barrier to the absorption of Ni by the plant and reduced root-to-shoot Ni translocation. Results indicate the potential of selected native AMF isolates from ultramafic areas for ecological restoration of such degraded ecosystems.     

Tolerance and induction of tolerance to Ni of arbuscular mycorrhizal fungi from New Caledonian ultramafic soils

The influence of Ni on arbuscular mycorrhizal fungi (AMF) has not been studied yet. We tested the tolerance to Ni of five AMF isolates from New Caledonian ultramafic soils. Spore germination indicated that these isolates were clearly more tolerant to Ni than three other isolates from non-ultramafic soils. They were able to germinate at 30 μg g⁻¹ Ni, whereas spores of the non-ultramafic isolates were totally inhibited at 15 μg g⁻¹ Ni. Among the ultramafic isolates, two were obtained from roots of Ni-hyperaccumulating plants. Their tolerance to Ni was clearly higher than all the other isolates. The proportion of germinated spores of the different isolates in contact with ultramafic soils showed the same tendencies as those observed with Ni solutions. Tolerance to Ni increased when spores were produced from mycorrhiza on plants grown on sand containing 20 μg g⁻¹ Ni, in comparison with those produced on sand without Ni. These results indicate that the tolerance to Ni of AMF spores can be induced by the presence of this metal in the substrate.

Localization of Nickel in Tissues of Streptanthus polygaloides Gray (Cruciferae) and Endemic Nickel Hyperaccumulators from California

The genus Streptanthus Nutt. is one of the most important indicators of ultramafic floras in western North America. This genus contains taxa that are endemic or tolerant of ultramafic soils. Streptanthus polygaloides is an annual nickel hyperaccumulator strictly confined to ultramafic soils throughout the Californian Sierra Nevada foothills. Nickel concentration in S. polygaloides populations was evaluated by elemental microanalysis using inductively coupled plasma mass spectrometry (ICP-MS). Representative samples of S. polygaloides roots, stems, leaves, flowers, and fruits were studied by scanning electron microscopy (SEM) coupled to an energy-dispersive X-ray probe (SEM-EDX). Results show Ni accumulation values between 0.09 and 1.18 %, and a distribution pattern similar to that observed in other Ni hyperaccumulator taxa, with the leaf epidermis accumulating the largest concentrations.     

Nickel distribution in the hyperaccumulator Alyssum serpyllifolium Desf. spp. from the Iberian Peninsula

Abstract

The relative concentration and distribution of nickel (Ni) in vegetative tissues (leaves, stems and trichomes) and reproductive organs (seeds) was studied using energy-dispersive X-ray microanalysis (EDXS) and scanning electron microscopy (SEM) in two previously studied Ni-hyperaccumulator subspecies of Alyssum serpyllifolium Desf. growing naturally in ultramafic soils of the Iberian Peninsula: A. serpyllifolium ssp. lusitanicum Dudley & P. Silva and A. serpyllifolium ssp. malacitanum Rivas Goday ex G. López. Both taxa showed that Ni accumulates preferentially in the leaves, exhibiting a homogeneous distribution on both epidermis surfaces. The highest Ni concentrations were found inside the epidermal cells and at the base of trichome stalks. Ni accumulation in seeds was lower than in the vegetative organs. The location of Ni in these plants allows us to postulate that its accumulation is a protection mechanism against external stress.     

Nickel Hyperaccumulation in the Serpentine Flora of Cuba

Extraordinary uptake (hyperaccumulation) of nickel (Ni), reachingconcentrations of 0.1–5.0%, about 1000-times greater thanthose usually found in flowering plants, has been reported overthe period 1948–1996 in about 190 species that grow onNi-rich serpentine soils derived from ultramafic rocks in variousparts of the world. A recent study of the families Buxaceaeand Euphorbiaceae identified a further 80 Ni hyperaccumulatorsfrom the very large ultramafic flora of Cuba, the largest numberfound to date in any one country. A much wider investigationof the elemental content of plants from the Cuban ultramaficflora, reported here with representative analyses of the correspondingsoils, has revealed Ni hyperaccumulation in an additional 50taxa (in 16 genera and eight families). The number of hyperaccumulatorsis greatest on the oldest serpentine soils, which are believedto have been available for colonization for the last 10–30million years. Both Ni hyperaccumulators, and serpentine endemicspecies generally, are much more frequent on these old soils,occurring in the eastern and western extremities of Cuba, thanon those developed within the last million years in the centralpart of the country. Hyperaccumulating plants of the familiesAcanthaceae, Asteraceae, Clusiaceae, Myrtaceae, Ochnaceae, Oleaceae,Rubiaceae and Tiliaceae are discussed.Copyright 1999 Annalsof Botany Company. Nickel, hyperaccumulation, serpentine flora, Cuba.     

Aspects of reproduction in Cerastium alpinum on calcic and ultramafic soils in Central Norway

Cerastium alpinum is one of several species of Caryophyllaceae able to grow on soils of deviating chemical composition, including ultramafic soils. To compare mating systems between plants growing on calcic and ultramafic soils, pollination experiments were performed in one ultramafic and four calcic sites in Central Norway. Two populations on calcic soils were classified as ssp. lanatum and two as ssp. alpinum . The results suggest that C. alpinum is a highly self compatible mixed mater with a large proportion of facilitated selfing. Plants on different soils showed no significant difference in mating system. However, relative seed set and number of initial ovules differed. The population on ultramafic soil produced a significantly lower number of initial ovules than those on calcic soils. Relative seed set was also lower on ultramafic soil. The difference in number of initiated ovules, along with known differences in various morphological traits, i.e. plants on ultramafic soils display a more dwarfed stature, suggest that plants on ultramafic soils allocate more resources to survival than to reproduction and growth compared to plants on calcic soils. In addition, on calcic soils a higher relative seed set was found in ssp. lanatum than in ssp. alpinum , probably caused by habitat differences between the investigated populations rather than by genetic differences between the two taxa. The obvious success of C. alpinum on ultramafic soils may partly be ascribed to easy establishment in this scarcely vegetated habitat.     

Bacterially Induced Weathering of Ultramafic Rock and Its Implications for Phytoextraction

The bioavailability of metals in soil is often cited as a limiting factor of phytoextraction (or phytomining). Bacterial metabolites, such as organic acids, siderophores, or biosurfactants, have been shown to mobilize metals, and their use to improve metal extraction has been proposed. In this study, the weathering capacities of, and Ni mobilization by, bacterial strains were evaluated. Minimal medium containing ground ultramafic rock was inoculated with either of two Arthrobacter strains: LA44 (indole acetic acid [IAA] producer) or SBA82 (siderophore producer, PO₄ solubilizer, and IAA producer). Trace elements and organic compounds were determined in aliquots taken at different time intervals after inoculation. Trace metal fractionation was carried out on the remaining rock at the end of the experiment. The results suggest that the strains act upon different mineral phases. LA44 is a more efficient Ni mobilizer, apparently solubilizing Ni associated with Mn oxides, and this appeared to be related to oxalate production. SBA82 also leads to release of Ni and Mn, albeit to a much lower extent. In this case, the concurrent mobilization of Fe and Si indicates preferential weathering of Fe oxides and serpentine minerals, possibly related to the siderophore production capacity of the strain. The same bacterial strains were tested in a soil-plant system: the Ni hyperaccumulator Alyssum serpyllifolium subsp. malacitanum was grown in ultramafic soil in a rhizobox system and inoculated with each bacterial strain. At harvest, biomass production and shoot Ni concentrations were higher in plants from inoculated pots than from noninoculated pots. Ni yield was significantly enhanced in plants inoculated with LA44. These results suggest that Ni-mobilizing inoculants could be useful for improving Ni uptake by hyperaccumulator plants.     

Rhizosphere Microbial Densities and Trace Metal Tolerance of the Nickel Hyperaccumulator Alyssum Serpyllifolium Subsp. Lusitanicum

In this study we determine culturable microbial densities (total heterotrophs, ammonifiers, amylolytics and cellulolytics) and bacterial resistance to Co, Cr, and Ni in bulk and rhizosphere soils of three populations of the Ni-hyperaccumulator Alyssum serpyllifolium subsp. lusitanicum and the excluder Dactylis glomerata from ultramafic sites (two populations in Northeast (NE) Portugal (Samil (S), Morais (M)) and one population in Northwest (NW) Spain (Melide (L)). The relationship between bioavailable metal concentrations (H₂O-soluble) and microbial densities were analysed. Significant differences in microbial densities and metal-resistance were observed between the two species and their three populations. The hyperaccumulator showed higher microbial densities (except cellulolytics) and a greater rhizosphere effect, but this was only observed in S and M populations. These populations of A. serpyllifolium also showed selective enrichment of Ni-tolerant bacteria at the rhizosphere where Ni solubility was enhanced (densities of Ni-resistant bacteria were positively correlated with H₂O-soluble Ni). These rhizobacteria could solubilise Ni in the soil and potentially improve phytoextraction strategies.     

In Vitro Health Risk Assessment of Ingesting Metal-Enriched Soils and Dusts in a Chinese Mining City

To evaluate the potential health hazards caused by extensive vanadium–titanium magnetite mining, bioavailability and bioaccessibility of metals were assessed in the pluralistic mining–agriculture–residential city of Panzhihua, China. Intensive mining and related heavy traffic may have contributed to Ni, Cr, and Zn contamination and Mn and V accumulation in soils and road dusts. The metal bioavailability estimated by water extraction was significantly lower than metal bioaccessibility on the basis of the simple bioaccessibility extraction test. The bioaccessible metal concentrations were significantly and positively correlated with soil/dust total metal concentrations (p < .05). However, bioavailable metal concentrations exhibited no relationship with total metal concentrations except for V. The bioaccessibility of metals significantly varied and exhibited the following order: Pb > Zn > Mn > Ni > V > Cr. Health risk assessment indicated that the carcinogenic and non-carcinogenic risks posed by these metals were at an acceptable level, but Cr in soils of the mining area and V in surface soils along the Jinsha River were close to the safe level for children. Therefore, the potential health risk attributed to the exposure of children to metals in surface soils and road dusts in such areas should not be overlooked.     

Arbuscular mycorrhiza of Berkheya coddii and other Ni-hyperaccumulating members of Asteraceae from ultramafic soils in South Africa

The occurrence of arbuscular mycorrhiza (AM) in nickel-(Ni)-hyperaccumulating plants of the Asteraceae family growing on Ni-enriched ultramafic soils in South Africa was surveyed. All plants were found to be consistently colonised by AM fungi, with the abundant formation of arbuscules. Berkheya coddii, which is an important species for phytomining, formed well-developed mycorrhiza under greenhouse conditions. Plants cultivated under greenhouse conditions and inoculated with native fungi had a higher shoot biomass and Ni content than noninoculated plants. Inoculation of B. coddii with Glomus intraradices (BEG) was successful, but only plants with abundantly developed arbuscules showed increased yield. In other cases, shoot biomass was similar to noninoculated plants. Dense depositions localised on top of the arbuscules were often observed in roots collected from the field and from pot cultures.     

Release of Ni and Co by microbial activity in New Caledonian ultramafic soils

A high positive correlation was found between extractable Ni and Co contents and microbial activity of 40 ultramafic soil samples from New Caledonia, suggesting a possible role of microorganisms in the release of these two metals. A saprolite (ultramafic subsoil) and a hypermagnesian brown soil were incubated 9 months in different conditions. Ni and Co release, measured by their extractability, occurred without reduction of soil pH but did not occur when the native microflora was eliminated by heat treatment. However, when autoclaved soil was re-inoculated with a pinch of the same nonheated soil, the release of metals was recorded. The concentrations of extractable Ni and Co were much higher in soils amended with organic compounds in which microbial activity was enhanced, but only if the soil was not heat treated. The presence of Grevillea exul, a metal-tolerant plant, stimulated the metal release process, but the stimulation was less effective than it was in the compost-amended soil without plant. The influence of the plant in extractable Ni and Co contents in this amended soil was not significant. The release of the two metals therefore seemed to be induced by the activity of specialized organotrophic microorganisms.     

Foliar chemistry and leaf herbivory on mount Bloomfield,Palawan, Philippines

Summary

Mount Bloomfield has a range of vegetation types on non-ultramafic and ultramafic soils. We report here foliar chemical analyses for N, P, K, Na, Ca, Mg, AI, B, Co, Cr, Cu, Fe, Mo, Mn, Ni, Si, Zn made on a large number of tree species. The most important results were: the notably high calcium in the ultramafic forest leaves, the occurrence of an aluminium accumulating species and two nickel accumulators on the ultramafic soils. Measurements of herbivory by the leaf-hole method showed that the percentage leaf area consumed by herbivores was similar in all the plots but the actual leaf area consumed was greater in the non-ultramafic forest because this had the largest leaves. There was no relationship between herbivory, and any of the leaf mineral elements investigated. Even the accumulating species were attacked by herbivores, of which the gall-formers and leaf-miners must be tolerant of high element concentrations, since the whole of their juvenile existence is in the leaf tissue.     

Vegetation and soil and plant chemistry on ultramafic rocks in the tropical Far East

The tropical Far East has many outcrops of ultramafic rock including very large areas in Sulawesi (c. 8000 km²) and New Caledonia (c. 5500 km²). The outcrops occur under several different climates, and give rise to a range of soils, the characteristics of which are reviewed. The vegetation on them is very varied. Under the same climate one can find grassland, scrub, and both short and tall rain forests. The variation in species richness on the ultramafics is difficult to explain. The degree of endemism varies too; it is probably less dependent on soil characteristics than on historical factors. The causes of the various unusual types of vegetation on ultramafic outcrops are discussed. It is possible that the somewhat dwarfed forests result from a shortage of one or more major nutrients or from very high soil Mg/Ca quotients or high Ni concentrations. The distinct ‘maquis’ vegetation of New Caledonia, and probably ultramafic scrub elsewhere, has evolved in relation to not only the soil chemical factors just listed but also periodic fire and varying degrees of drought. Fires are certainly more important than was once thought and the adverse soil factors may have a role in delaying recolonisation. The plant chemistry is notable for the presence of species which hyperaccumulate certain elements, notably Ni. This phenomenon is discussed in relation to its ecological importance, which may be protection of the hyperaccumulators against herbivores. The need for a conservation policy for the ultramafic areas is stressed, and mention is made of the restoration work on sites damaged by nickel mining in New Caledonia.     

镍污染土壤植物采矿技术关键过程及其研究进展

植物采矿是利用超积累植物高量吸收土壤中的重金属,并从中提取、冶炼金属产品,在修复污染土壤的同时实现金属的资源化。全世界广泛分布着自然风化的镍污染土壤,植物采矿因其重要的环境、生态及资源价值,被作为一种环境友好且具备经济效益的土壤修复技术,在此类地区具有广阔的应用前景。该植物采矿技术关键过程主要包括超积累植物镍高选择性根际环境响应、植物镍高效吸收转运以及生物质中镍高附加值资源化等过程。近30年,污染土壤中镍的植物采矿已经在美国、阿尔巴尼亚、马来西亚等多个国家进行了野外实践,取得了良好效果。然而,相关技术在我国的研究与应用仍然处于起步阶段。文中通过综述植物采矿技术的关键过程的研究进展,发现其中的瓶颈,为接下来植物采矿的科学研究和技术在全世界推广提供理论基础和技术指导。     

Bacterial community diversity and functional roles in the rhizosphere of Rinorea cf. bengalensis and Phyllanthus rufuschaneyi under a nickel concentration gradient

Plant and Soil - The tropical nickel (Ni) hyperaccumulator plants, Rinorea cf. bengalensis and Phyllanthus rufuschaneyi, are locally common on ultramafic soils in the Malaysian state of Sabah on...