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34 ref. doi: 10.1016/j.envpol.2004.10.020; International audience; More knowledge is needed concerning the disturbance of soil organic matter cycling due to heavy metal pollution. The present study deals with the impact of heavy metal pollution on litter breakdown. Our aim was to assess whether heavy metals initially present in the leaves of the metallophyte Arabidopsis halleri: (i) slow down the rate of C mineralization, in relation to metal toxicity towards microflora, and/or (ii) increase the amount of organic C resistant to biodegradation, in relation to an intrinsic resistance of metallophyte residues to biodegradation. We incubated uncontaminated soil samples with either metal-free or metal-rich plant material. Metal-free material was grown in a greenhouse, and metal-rich material was collected in situ. During the 2-month period of incubation, we measured evolved CO2-C and residual plant C in the coarse organic fraction. Our results of CO2-C evolution showed a similar mineralization from the microcosms amended with highly metal-rich leaves of A. halleri and the microcosms amended with the metal-free but otherwise similar plant material. Measuring residual plant C in its input size-fraction gave a more precise insight. Our results suggest that only the large pool of easily decomposable C mineralized similarly from metal-free and from metal-rich plant residues. The pool of less decomposable C seemed on the contrary to be preferentially preserved in the case of metal-rich material. These results support the hypothesis of an annual extra-accumulation in situ of such a slowly decomposable fraction of plant residues which could account to some extent for the observed accumulation of metallophyte litter on the surface of highly metal-polluted soils.

36 ref. doi: 10.1016/j.envpol.2004.07.031; International audience; In order to better understand the fate of metals during the biodegradation of organic matter in soils, an in vitro incubation experiment was conducted with metal-rich and metal-free leaves of Arabidopsis halleri introduced in a non-contaminated soil. During incubation of these microcosms, we followed the partitioning of Zn and Cd between the solution and their solid components, by determining the metal contents of six soil fractions and dissolved metals after granulo-densimetric separations at selected times. Microbial biomass and exchangeable metals in K2SO4 solutions were also determined at the same times, and two main stages were identified. The first one takes place after a fast abiotic transfer of Zn and Cd from readily soluble plant tissues onto fine soil constituents, keeping metals away from the liquid phase: during about 14 days, microbial biomass increased as well as metal contents of some soil fractions, particularly those rich in particulate organic matter. During the second stage, between 14 and 60 days and for the metal-rich microcosms, Zn and Cd contents in solution increased, while microbial biomass decreased instead of staying constant as in control. A change of Zn and Cd speciation is assumed, from non-toxic adsorbed forms to more toxic species in solution. Remaining metal-rich plant residues seem to create a stable organic C compartment in the soil.

Rhizosphere bacterial populations associated with four metallophyte plants in one of major polymetallic (Pb–Zn–Cu) semiarid Moroccan Hercynian province (Draâ Sfar, Marrakech, Morocco) presenting long-term contamination mainly with Zn and Pb were analysed and compared to selected control soils. In the highly Zn-, Cu-, Pb- and Cd- contaminated soils, the total number of culturable heterotrophic bacteria were found in low proportions (< 2.6 × 102 – 1.6 × 104 g−1soil). This bacterial content was slightly similar to that found in moderately polluted and controls soils (6.7 × 104 – 5.8 × 106). However, the bacterial diversity and the rhizosphere/soil ratio, which compares the bacterial content (or bacterial charge) around the metallophyte plants with that in non-rhizosphere soil, were the bacteriological parameters mostly affected by heavy metal contamination. The chronic Zinc-stress results in an increase of tolerance to this metal of both the rhizosphere and non-rhizosphere bacterial communities. However, in general, the rhizosphere bacterial populations exhibited less tolerance to Zn toxicity than the bacterial population of non-rhizosphere soils. This result suggests that toxic effects of Zn decrease in the rhizosphere soils of the metallophyte plants.

Plants that have evolved to survive on metal-rich soils-metallophytes-have key values that must drive research of their unique properties and ultimately their conservation. The ability of metallophytes to tolerate extreme metal concentrations commends them for revegetation of mines and metal-contaminated sites. Metallophytes can also be exploited in environmental technologies, for example, phytostabilization, phytoremediation, and phytomining. Actions towards conserving metallophyte species are imperative, as metallophytes are increasingly under threat of extinction from mining activity. Although many hundreds of papers describe both the biology and applications of metallophytes, few have investigated the urgent need to conserve these unique species. This paper identifies the current state of metallophyte research, and advocates future research needs for the conservation of metallophyte biodiversity and the sustainable uses of metallophyte species in restoration, rehabilitation, contaminated site remediation, and other nascent phytotechnologies. Six fundamental questions are addressed: (1) Is enough known about the global status of metallophytes to ensure their conservation? (2) Are metallophytes threatened by the activities of the minerals industry, and can their potential for the restoration or rehabilitation of mined and disturbed land be realized? (3) What problems exist in gaining prior informed consent to access metallophyte genetic resources and how can the benefits arising from their uses be equitably shared? (4) What potential do metallophytes offer as a resource base for phytotechnologies? (5) Can genetic modification be used to "design" metallophytes to use in the remediation of contaminated land? (6) Does the prospect of using metallophytes in site remediation and restoration raise ethical issues? © 2004 Society for Ecological Restoration International.

In order to assess the importance of colloids in the metal transfer in soils under varying land use (a podzol under a forest, a cultivated luvisol and a luvisol under a metallophyte grassland), gravitational waters were collected in situ by zero-tension lysimeters. Dissolved and colloidal fractions of metals were separated by ultracentrifugation and colloids were studied by transmission electron microscopy (TEM) coupled with energy dispersive X-ray analysis (EDS). The stability of metal complexes in the collected solutions was operationally defined by an ion exchange method. The results show that in the two luvisols about 75% of Pb was always present in colloidal forms and a substantial fraction of Pb in solution was present as stable complexes and labile complexes with slow dissociation kinetics. Zn and Cd were found mostly in dissolved forms and mainly as free ions or labile complexes in solution. In the podzol, all metals were found in dissolved forms and as free ions or labile complexes. The Zn-bearing colloids were organic (biocolloids) or mineral (smectites, Fe-oxyhydroxides, Ca- or Al-rich phosphates) depending on soil type and land use. The results are discussed in terms of metal mobility to explain the different vertical metal distributions observed in the soil profiles.

The recent United Nations RIO+20 Conference on Sustainable Development highlighted the rapid increase in industrial mineral exploration and extraction over the past two decades ([ 1 ][1]). Endemic metallophyte plants, which have evolved where metal accumulations extend to the Earth's surface, have

Using the g-glutamylcysteine synthetase inhibitor, L-buthionine-[S,R]-sulphoximine (BSO), the role for phytochelatins (PCs) was evaluated in Cu, Cd, Zn, As, Ni, and Co tolerance in non-metallicolous and metallicolous, hypertolerant populations of Silene vulgaris (Moench) Garcke, Thlaspi caerulescens J.&C. Presl., Holcus lanatus L., and Agrostis castellana Boiss. et Reuter. Based on plant-internal PCthiol to metal molar ratios, the metals’ tendency to induce PC accumulation decreased in the order As/Cd/Cu > Zn > Ni/Co, and was consistently higher in non-metallicolous plants than in hypertolerant ones, except for the case of As. The sensitivities to Cu, Zn, Ni, and Co were consistently unaffected by BSO treatment, both in non-metallicolous and hypertolerant plants, suggesting that PC-based sequestration is not essential for constitutive tolerance or hypertolerance to these metals. Cd sensitivity was considerably increased by BSO, though exclusively in plants lacking Cd hypertolerance, suggesting that adaptive cadmium hypertolerance is not dependent on PC-mediated sequestration. BSO dramatically increased As sensitivity, both in non-adapted and As-hypertolerant plants, showing that PC-based sequestration is essential for both normal constitutive tolerance and adaptive hypertolerance to this metalloid. The primary function of PC synthase in plants and algae remains elusive.

A relatively small yet diverse group of plants are capable of sequestering metals in their shoot tissues at remarkably high concentrations that would be toxic to most organisms. This process, known as metal hyperaccumulation, is of interest for several reasons, including its relevance to the phytoremediation of metalpolluted soils. Most research on hyperaccumulators has focused on the physiological mechanisms of metal uptake, transport, and sequestration, but relatively little is known regarding the genetic basis of hyperaccumulation. There are no known cases of major genetic polymorphisms in which some members of a species are capable of hyperaccumulation and others are not. This is in contrast to the related phenomenon of metal tolerance, in which most species that possess any metal tolerance are polymorphic, evolving tolerance only in local populations on metalliferous soil. However, although some degree of hyperaccumulation occurs in all members of the species that can hyperaccumulate, there is evidence of quantitative genetic variation in ability to hyperaccumulate, both between and within populations. Such variation does not appear to correlate positively with either the metal concentration in the soil or the degree of metal tolerance in the plant. Studies using controlled crosses, interspecific hybrids, and molecular markers are beginning to shed light on the genetic control of this variation. As molecular physiology provides greater insights into the specific genes that control metal accumulation, we may learn more about the genetic and regulatory factors that influence variable expression of the hyperaccumulation phenotype.

This work was undertaken to measure the in situ phytoextraction of zinc using a former zinc-smelter site where metallophyte plants have been growing for 30 years. The site exhibited a gradient in the total metal concentration in the upper horizon (from 3230 to 8530 mg Zn kg −1 ). Soils were sampled from four different sectors (I–IV), and plant shoots were harvested, identified, their biomass weighed and analysed for zinc. The results showed that three plant species were dominant on the site, including Arabidopsis halleri (cress), Armeria maritima (seathrift), and Arrhenatherum elatius (fromental). A. maritima was the predominant species according to the biomass production on the most polluted sector I. As the concentration of metals in soils decreased, A. maritima disappeared and A. halleri increased. The biomass of A. elatius was the highest on the less polluted soils. Concentrations in zinc in the aerial parts of plants varied from 73 (sector IV) to 6269 mg kg −1 DM (sector I). The concentration of Zn in A. halleri decreased with the decrease in concentration of zinc in soil. Phytoextraction was calculated from the biomass and its concentration of metal. It was at a maximum in sector III with a high contribution of A. halleri and A. elatius and reached 10 kg Zn ha −1 , a promising amount for phytoextraction considering the absence of any agricultural practices. In sector I, phytoextraction was four times lower despite a 2.6 times higher concentration of Zn in the upper horizon. In conclusion, phytoextraction was strongly dependent on the concentration of the available metal in soils which may limit the growth of plants, and favour tolerant but low biomass plant species such as A. maritima .

Some higher plant species have developed heavy metal tolerance strategies which enable them to survive and reproduce in highly metal-contaminated soils. We have investigated such heavy metal uptake and accumulation strategies of two absolute metallophyte species (Armeria maritima ssp. halleri and Cardaminopsis halleri) and one pseudometallophyte (Agrostis tenuis) growing near a former metal smelter. Samples of plant parts and soil were analysed for Zn, Cd, Pb, and Cu. In soil, there were two dominant types of metal concentration gradients with depth. Under the absolute metallophytes, extremely high metal contents were measured in the surficial Ah horizon, followed by a strong decrease in the underlying soil horizons (L(11) and L(12)). Under the pseudometallophyte, metal concentrations in the Ah horizon were much lower and fewer differences were observed in metal concentrations among the Ah, L(11), and L(12) horizons. The concentrations of Zn, Cd, Pb, and Cu in Agrostis tenuis roots were greater than concentrations in leaves, indicating significant metal immobilisation by the roots. For C. halleri, Zn and Cd concentrations in leaves were20,000 and100 mg kg(-1), respectively, indicating hyperaccumulation of these elements. Armeria maritima ssp. halleri exhibited root concentrations of Pb and Cu that were 20 and 88 times greater, respectively, than those in green leaves, suggesting an exclusion strategy by metal immobilisation in roots. However, Zn, Cd, Pb, and Cu concentrations in brown leaves of Armeria maritima ssp. halleri were 3-8 times greater than in green leaves, suggesting a second strategy, i.e. detoxification mechanism by leaf fall.

This chapter presents knowledge of the mechanisms and genetics of heavy metal tolerance in hyperaccumulating and nonhyperaccumulatitig metallophytes. Strongly heavy metal-enriched substrates are hostile to plant growth. The colonization of metalliferous substrates involves evolutionary adaptation. Plants of facultative metallophyte populations from metalliferous soil almost invariably exhibit higher levels of metal tolerance than those of normal soil populations. Metalliferous soils are often enriched in combinations of different heavy metals and, therefore, local metallophyte populations often exhibit combined tolerance to different metals. The physiological and biochemical mechanisms of metal tolerance are poorly understood. In general, tolerance in unadapted and metal-adapted plants might depend on qualitatively different mechanisms. Hyperaccumulators, just like nonhyperaccumulating metallophytes, may show relatively high levels of tolerance to metals that are not present at toxic concentrations in soil at the sites of population origin. Hyperaccumulators seem to exhibit a high degree of metal specificity with regard to foliar accumulation.

Metallophytes, plant species that grow only on soils rich in metals, are used for bioremediation of polluted soils. However, little is known about the functioning of such soil–plant systems. We have investigated soil organic matter (SOM) dynamics and its effects on the fate of heavy metals under a metallophyte grassland highly polluted by industrial dust fallout. Both litter and soil horizons were sampled. Particle-size and density fractionations were carried out to separate particulate organic matter (POM), i.e. the light fraction >50 μm. All samples were analysed for C, N, Zn, Pb, and Cd. Bulk densities were determined for all horizons and stocks of elements per unit area were calculated. Compared to broad uncontaminated temperate grasslands, SOM displayed similar quantities but differed significantly as to its quality and dynamics. The main differences were a lack of incorporation of plant returns in the soil profile and an imbalance of SOM composition towards more POM and less fine humified material. Zn, Pb, and, to a lesser extent, Cd were located mainly in the organic-rich, superficial soil layer. Heavy metal concentrations of POM of different sizes were similar within each horizon. Heavy metal concentrations of total POM increased strongly according to a depth–time scale. Our results suggest a selective decomposition of portions of metallophyte-derived debris with initially low heavy metal concentrations and resistance to biodegradation of those portions with initially high heavy metal concentrations. Such a mechanism may constitute a process of mutual protection, in this soil–plant system, of plant debris towards biodegradation and of heavy metals towards mobility.

The tolerance of the metallophyte Silene vulgaris, a plant suitable for the phytostabilisation of metal(loid)-contaminated soils, to arsenic (As), mercury (Hg) and cadmium (Cd) was evaluated in a semi-hydroponic culture system under controlled environmental conditions. The appearance of oxidative stress, alteration of photochemical processes and modification of biothiol content were studied as physiological parameters of metal(loid) stress in plants treated with 0, 6 and 30 μM (As, Hg or Cd) for 7 days. In spite of the metal(loid) excluder behaviour of S. vulgaris, Cd was translocated to the aerial part of the plant at a higher rate than Hg or As. The major toxic effects were observed in roots, where lipid peroxidation was increased in a dose-dependent manner. Redox enzymes such as glutathione reductase (GR) were severely inhibited by Hg, whereas GR was overexpressed. The accumulation of Cd produced a remarkable production of phytochelatins (PCs) in roots, whereas Hg and As led to modest PCs synthesis. There was a severe loss of chlorophyll content in Cd-treated plants, accompanied with a significant decrease in photosystem II efficiency (ΦPSII) and photochemical quenching (qP). Similar negative effects were observed in Hg- and As-exposed plants, but to a lesser degree. The exposure to the highest dose of each toxic element (30 μM) caused depletion of the light harvesting complex b1 protein. In conclusion, specific stress signatures to each metal(loid) were observed, with As being the least toxic element, suggesting that different mechanisms of tolerance were exerted. These results could be applied in future experiments to select tolerant ecotypes to optimize the phytostabilisation of metal(loid) multipolluted soils.

Stackhousia tryonii Bailey is a rare, serpentine-endemic herb, with potential for use in phytoremediation and/or phytomining. This study evaluates the use of herbaceous heel cuttings to propagate S. tryonii on three rooting media [sand, Medium I; a commercial nursery mix, Medium II; and sand : peat moss : perlite (2 : 2 : 1; v/v), Medium III] following the application of plant growth hormones, viz. indole-3-butyric acid and naphthalene acetic acid (both as pure and commercial formulations, viz. Clonex Gel-green, Gel-purple and Gel-red), and honey. Cutting survival, rooting percentage, the number of primary and secondary roots produced, the length of the longest root and the total root length were evaluated after 10 weeks. Results show that there was a positive (P < 0.001) influence of plant growth regulator treatments on cuttings' survival. Medium III failed to support survival of cuttings. Percentage rooting (as a proportion of original number of cuttings) was 10–30% higher in Medium I than in Medium II. Cuttings treated with Clonex Gel-red, IBA at 1000 ppm and NAA at 250 ppm had 60–70% higher percentage rooting in Medium�I than in Medium II. Total number of primary roots per rooted cuttings was significantly (P�

Metalliferous substrates are important in the distribution of various plants such asArmeria maritima, Minuartia verna, Thlaspi alpestre, Silene maritima andCochlearia officinalis, in parts of the United Kingdom. Metalliferous substrates are similarly important for a slightly different assemblage of metallophytes in the Aachen-Liege orefield in Belgium and Germany. The significance of metalliferous substrates in the distribution of the metallophytes varies from orefield to orefield and this is investigated. Certain sites, by virtue of their environmental and species diversity, are outstanding as habitats for the metallophytes and some of these are described. Observations on the importance and problems of the conservation of the metallophyte habitats are included.

The plant-communities from habitats of the metallophyte species Minuartia verna and Thlaspi alpestre (T. caerulescens) at sites disturbed and undisturbed by mining are described. Four communities were delineated by cluster and principal component analysis. Group 1 comprised species-poor communities on disturbed non-calcareous soils; group 2, relatively species-rich communities on disturbed calcareous soils; group 3, species-rich communities in the main on undisturbed calcareous soils. Group 4 consisted of species-rich communities with an alpine element, in damp habitats on base-rich soils derived from igneous rocks. Total and exchangeable elements As, Ba, Ca, Cd, Co, Cr, Cu, Fe, Mg, Mn, Ni, Pb and Zn were determined for the soils of these sites. Levels of soil Ca, Cd, Pb and Zn accounted for most of the variation along the first axis of the PCA and soil nutrient levels were probably the main predictor along the second.

Experiments were carried out on the tolerance to and uptake of copper and cobalt by three members of a phylogenetic series of taxa within the genusSilene from Zaire which were reputed to represent a progression of increasing adaptation to metalliferous soils. Plants studied were the widespread and presumably non-tolerantSilene burchelli var.angustifolia (B), a more tolerant variant of this taxon (E) designated an ecotype, and the metallophyteS. cobalticola (C). Studies on the effects of both metals, singly and in combination, on seed germination, seedling and plant performance and yield, and metal uptake from soil cultures, confirmed in general the sequence B

Plants colonizing metalliferous soils have evolved physiological mechanisms which enable them to tolerate metal toxicity. These mechanisms do not generally suppress metal uptake but result in internal detoxification. Two basic strategies of plant response are suggested, accumulators and excluders. In the former, metals can be concentrated in plant parts from low or high background levels. By contrast, differential uptake and transport between root and shoot in excluders, lead to more or less constant low shoot levels over a wide range of external concentration. ‘Indicators’ are seen as a further mode of response where proportional relationships exist between metal levels in the soil, uptake and accumulation in plant parts. The physiological properties of accumulator and excluder species are considered in relation to metal tolerance mechanisms.

Publisher Summary This chapter illustrates the literature on those plants and micro-organisms which can combat excessive quantities of heavy metal ions. Heavy metals include those metals which have density greater than five. Their common feature in regard to biological life is that in excessive quantities they are poisonous and can cause death of most living organisms. However, certain organisms possess an ability to survive under conditions of metal contamination which can prove toxic to other living things. Toxic levels of heavy metals can occur under several circumstances. The chapter focuses on the type of contamination in which the soil itself contains large quantities of these metals. The contamination results from the presence of undisturbed metal ore near the soil surface causing anomalies or from the actual mining of ore bodies. The vegetation in such areas is influenced by one overriding factor, namely, metal concentration. The habitats are usually spatially distinct and clear cut. It is observed that the sequence and pattern of genetic change responsible for colonization of metal-contaminated areas serves as a unique record of natural selection in action.