Life history variation in the heavy metal tolerant plant Thlaspi caerulescens growing in a network of contaminated and noncontaminated sites in southern France: role of gene flow, selection and phenotypic plasticity

* Here we explore life history differences in a set of neighbouring metallicolous and nonmetallicolous populations of the heavy metal tolerant plant Thlaspi caerulescens. * We contrasted data from field observations and from a common garden experiment, in which soil zinc (Zn) concentration and light availability were manipulated, and data on microsatellite molecular variation. * The two ecotypes showed few differences in life history in the field, but large differences in their response to Zn concentration in the common garden. Soil toxicity affected most characters in nonmetallicolous plants, while it had no effect on metallicolous plants. The two ecotypes responded similarly to light. Genetic differentiation for quantitative characters between ecotypes contrasted with the absence of differentiation for microsatellites. Conversely, populations of the same ecotype showed similar responses to Zn, despite their high differentiation for molecular markers. * We conclude that divergent selection related to soil toxicity has had a predominant role in shaping life history differences between ecotypes, gene flow weakly opposing local adaptation despite geographical proximity.     

Seasonal fluctuations of selenium and sulfur accumulation in selenium hyperaccumulators and related nonaccumulators

Some plants hyperaccumulate selenium (Se) up to 1% of dry weight. This study was performed to obtain insight into whole-plant Se fluxes in hyperaccumulators. Selenium hyperaccumulators Astragalus bisulcatus and Stanleya pinnata were monitored over two growing seasons for seasonal fluctuations in concentrations of Se and the chemically similar element sulfur (S). The related nonhyperaccumulators Astragalus sericoleucus, Oxytropis sericea and Thlaspi montanum were included for comparison. In both hyperaccumulators leaf Se decreased from April to October, coinciding with Se hyperaccumulation in flowers and seeds. Root Se levels were lowest in summer. Selenium concentration decreased with leaf age in both hyperaccumulators. Leaf S levels peaked in summer in all plant species, as did Se levels in nonhyperaccumulators. Selenium and S levels tended to be negatively correlated in hyperaccumulators, and positively correlated in nonhyperaccumulators. These results suggest a specific flow of Se in hyperaccumulator plants over the growing season, from root to young leaves in spring, followed by remobilization from aging leaves to reproductive tissues in summer, and back to roots in the autumn.     

Plant Cd2+ and Zn2+ status effects on root and shoot heavy metal accumulation in Thlaspi caerulescens

* In this study we address the impact of changes in plant heavy metal, (i.e. zinc (Zn) and cadmium (Cd)) status on metal accumulation in the Zn/Cd hyperaccumulator, Thlaspi caerulescens. * Thlaspi caerulescens plants were grown hydroponically on both high and low Zn and Cd regimes and whole-shoot and -root metal accumulation, and root (109)Cd(2+) influx were determined. * High-Zn-grown (500 microm Zn) plants were found to be more Cd-tolerant than plants grown in standard Zn conditions (1 microm Zn). Furthermore, shoot Cd accumulation was significantly greater in the high-Zn-grown plants. A positive correlation was also found between shoot Zn accumulation and increased plant Cd status. Radiotracer (109)Cd root flux experiments demonstrated that high-Zn-grown plants maintained significantly higher root Cd(2+) influx than plants grown on 1 microm Zn. It was also found that both nickel (Ni) and copper (Cu) shoot accumulation were stimulated by high plant Zn status, while manganese (Mn) accumulation was not affected. * A speculative model is presented to explain these findings, suggesting that xylem loading may be one of the key sites responsible for the hyperaccumulation of Zn and Cd accumulation in Thlaspi caerulescens.     

Do metal-rich plants deter herbivores? A field test of the defence hypothesis

Some plant species growing on metalliferous soils are able to accumulate heavy metals in their shoots up to very high concentrations, but the selective advantage of this behaviour is still unknown. The most popular hypothesis, that metals protect plants against herbivores, has been tested several times in laboratory conditions, with contradictory results. We carried out the first large-scale test of the defence hypothesis in eight natural populations of the model Zn hyperaccumulator Thlaspi caerulescens J. and C. Presl (Brassicaceae). In two climatic regions (temperate, Belgium–Luxembourg, and Mediterranean, southern France), we worked in metalliferous and in normal, uncontaminated environments, with plants spanning a wide range of Zn concentrations. We also examined the importance of glucosinolates (main secondary metabolites of Brassicaceae) as antiherbivore defences. When exposed to natural herbivore populations, T. caerulescens suffered lower herbivory pressures in metal-enriched soils than in normal soils, both in Belgium–Luxembourg and in southern France. The trapping of gastropods shows an overall lower population density in metalliferous compared to normal environments, which suggests that herbivory pressure from gastropods is lower on metalliferous soils. In addition, foliar concentration of glucosinolates was constitutively lower in all populations from metal-enriched soils, suggesting that these have evolved towards lower investment in organic defences in response to lower herbivory pressure. The Zn concentration of plants had a protective role only for Belgian metallicolous plants when transplanted in normal soils of Luxembourg. These results do not support the hypothesis that Zn plays a key role in the protection of T. caerulescens against enemies. In contrast, glucosinolates appear to be directly involved in the defence of this hyperaccumulator against herbivores.     

The potential of Thlaspi caerulescens for phytoremediation of contaminated soils

Uptake of Cd, Zn, Pb and Mn by the hyperaccumulator Thlaspi caerulescens was studied by pot trials in plant growth units and in populations of wild plants growing over Pb/Zn base-metal mine wastes at Les Malines in the south of France. The pot trials utilised metal-contaminated soils from Auby in the Lille area. Zinc and Cd concentrations in wild plants averaged 1.16% and 0.16% (dry weight) respectively. The unfertilised biomass of the plants was 2.6 t/ha. A single fertilised crop with the above metal content could remove 60 kg of Zn and 8.4 kg Cd per hectare. Experiments with pot-grown and wild plants showed that metal concentrations (dry weight basis) were up to 1% Zn (4% Zn in the soil) and just over 0.1% Cd (0.02% Cd in the soil). The metal content of the plants was correlated strongly with the plant-available fraction in the soils as measured by extraction with ammonium acetate and was inversely correlated with pH. Bioaccumulation coefficients (plant/soil metal concentration quotients) were in general higher for Cd than for Zn except at low metal concentrations in the soil. There was a tendency for these coefficients to increase with decreasing metal concentrations in the soil. It is proposed that phytoremediation using Thlaspi caerulescens would be entirely feasible for low levels of Cd where only a single crop would be needed to halve a Cd content of 10 g/g in the soil. It will never be possible to remediate elevated Zn concentrations within an economic time frame (<10 yr) because of the lower bioaccumulation coefficient for this element coupled with the much higher Zn content of the soils.     

Distribution and Metal-Accumulating Behavior of Thlaspi caerulescens and Associated Metallophytes in France

The heavy metal hyperaccumulator Thlaspi caerulescens is widespread in France on many kinds of sites and substrates, including Zn/Pb/Cd mine and smelter wastes, Ni-rich serpentine outcrops and a variety of nonmetalliferous soils. Thlaspi caerulescens is remarkable among the metallophytes of France because it accumulates Zn to high concentrations (almost always >0.1%, and often >1% in the dry matter) regardless of the total Zn concentration of the substrate. The extraordinary uptake of Zn from soils of normal Zn concentration draws attention to the need for studies of the mechanisms by which such mobilization and uptake can occur. Different populations of Thlaspi caerulescens in France show considerable variation in their ability to accumulate Cd; individuals in some populations contain as much as 0.1 to 0.4% Cd, the highest levels recorded in vascular plants. The hyperaccumulation of Ni (sometimes exceeding 1%) from serpentine soils in France is also noteworthy. Despite the generally low biomass, some very large individuals occur, giving good potential for selective breeding to improve the value of Thlaspi caerulescens for phytoremediation, especially of Cd. The high Zn uptake from all kinds of soils is a property shared by the related T. brachypetalum, and T. alpinum shows dual Zn- and Ni uptake, depending on the substrate. The extent to which other species of Thlaspi occurring in France exhibit metal accumulation is also discussed.