Local Adaptation of Metallicolous and Non‐Metallicolous Anthyllis vulneraria Populations: Their Utilization in Soil Restoration

Restoration of metalliferous mine soils requires using plant species tolerant to high metal concentrations and adapted to nutrient‐poor soil. Legumes can increase plant productivity through N₂‐fixation, but they are often scarce in metalliferous sites. We examined survival, growth, and tolerance of four populations of a legume, Anthyllis vulneraria, from two metalliferous (MET) Zn‐Pb mine sites, Avinières (AV) ([Zn‐EDTA] = 26,000 mg/kg) and Eylie (EY) ([Zn‐EDTA] = 4,632 mg/kg), and two non‐metalliferous (NMET) sites located in the south of France with the aim to select the most appropriate populations for restoration of mined soils. In a common garden experiment, plants from each population were reciprocally grown in soil from the provenance of each population. The two NMET populations exhibited high mortality and low growth rates in soil from the mined sites. The AV MET exhibited a high growth rate in metalliferous soils, but showed high mortality in non‐metalliferous soils. The growth of the EY MET was very low in the AV‐contaminated soil, but was the highest of all populations in moderately and non‐metalliferous soils. Plants from the AV MET population showed a high growth and survival in metalliferous soil and would be appropriate in the restoration of metal‐contaminated sites (>30,000 mg Zn kg^?1). The EY MET population would be adapted to the restoration of moderate metal‐contaminated soils (<30,000 mg Zn kg^?1). Taking into account the broad distribution of A. vulneraria, these two populations could be suitable for the restoration of derelict mine sites in mediterranean and temperate regions of Europe and North America.     

The Genetic Basis of Metal Hyperaccumulation in Plants

Referee: Professor Alan J.M. Baker, School of Botany, The University of Melbourne, VIC 3010, Australia 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.     

Morphological and structural plasticity of grassland species in response to a gradient in saline‐sodic soils

The abundance and distribution of species can be ascribed to both environmental heterogeneity and stress tolerance, with the latter measure sometimes associated with phenotypic plasticity. Although phenotypic plasticity varies predictably in response to common forms of stress, we lack a mechanistic understanding of the response of species to high saline‐sodic soils. We compared the phenotypic plasticity of three pairs of high and low saline‐sodic tolerant congeners from the families Poaceae (Leymus chinensis versus L. secalinus), Fabaceae (Lespedeza davurica versus L. bicolor) and Asteraceae (Artemisia mongolica versus A. sieversiana) in a controlled pot experiment in the Songnen grassland, China. The low tolerant species, L. secalinus and A. sieversiana exhibited higher plasticity in response to soil salinity and sodicity than their paired congeners. Highly tolerant species, L. chinensis and A. mongolica, had higher values for several important morphological traits, such as shoot length and total biomass under the high saline‐sodic soil treatment than their paired congeners. In contrast, congeners from the family Fabaceae, L. davurica and L. bicolor, did not exhibit significantly different plasticity in response to soil salinity and sodicity. All species held a constant reproductive effort in response to saline‐sodic soil stress. The different responses between low and high tolerant species offer an explanation for the distribution patterns of these species in the Songnen grassland. Highly tolerant species showed less morphological plasticity over a range of saline‐sodic conditions than their paired congeners, which may manifest as an inability to compete with co‐occurring species in locations where saline‐sodic soils are absent.     

Chloroplast microsatellites reveal that metallicolous populations of the Mediterranean shrub Cistus ladanifer L have multiple origins

Cistus ladanifer L. (Cistaceae) is a Mediterranean shrub covering different kinds of soils in the Western Mediterranean area. This species has colonised several metalliferous areas (serpentine outcrops as well as human-polluted sites) throughout its distribution range, and is therefore an interesting species to study the possible effects on genetic diversity and differentiation produced by the colonisation of areas polluted with heavy metals. The genetic structure of 33 natural populations distributed across its entire natural distribution range (Morocco, Portugal and Spain) and growing on either metalliferous or non-metalliferous soils was investigated using chloroplast microsatellites. Population genetic parameters were estimated and genetic groups were identified using Bayesian inference. In addition, we compared the genetic diversity and differentiation among metallicolous and non-metallicolous populations within each Bayesian-defined group. The cpSSR data suggested that metallicolous populations of Cistus ladanifer have arisen through multiple independent evolutionary origins within two different chloroplast lineages. Evidence that the soil type provoked genetic bottlenecks in metallicolous populations or genetic differentiation among metallicolous and non-metallicolous populations was not observed. Historical factors are the main cause of the present genetic structure of C. ladanifer. The nature of tolerance to heavy metals as a species-wide trait in this shrub is discussed.     

A comparative study of metal levels in plants from two contrasting lead-mine sites

Summary An investigation into the levels of metals in plants growing on metalliferous soils was carried out. The exchangeable metal concentration of soils from Tyndrum and Trelogan was found to be extremely variable from sample to sample and the influence of soil pH on the exchangeable lead and zinc concentration is discussed. Large differences in heavy metal levels were found between species and may be indicative of different mechanisms of tolerance to lead and zinc excess. Large differences in calcium levels between plant tissues were also found. These may result from the use of calcium in ameliorating lead and zinc toxicity in some species.     

Climate and Soil Type Together Explain the Distribution of Microendemic Species in a Biodiversity Hotspot

The grasshopper genus Caledonula, endemic to New Caledonia, was studied to understand the evolution of species distributions in relation to climate and soil types. Based on a comprehensive sampling of 80 locations throughout the island, the genus was represented by five species, four of which are new to science, of which three are described here. All the species have limited distributions in New Caledonia. Bioclimatic niche modelling shows that all the species were found in association with a wet climate and reduced seasonality, explaining their restriction to the southern half of the island. The results suggest that the genus was ancestrally constrained by seasonality. A molecular phylogeny was reconstructed using two mitochondrial and two nuclear markers. The partially resolved tree showed monophyly of the species found on metalliferous soils, and molecular dating indicated a rather recent origin for the genus. Adaptation to metalliferous soils is suggested by both morphological changes and radiation on these soils. The genus Caledonula is therefore a good model to understand the origin of microendemism in the context of recent and mixed influences of climate and soil type.     

A new dawn – the ecological genetics of mycorrhizal fungi

Many human activities, such as ore mining and smeltering, sewage sludge treatment and fossil fuel consumption, result in toxic soil concentrations of 'heavy metals' (Al, Cd, Co, Cr, Cu, Hg, Mn, Ni, Pb, Ti, Zn and others) (Gadd, 1993). There are also natural soils, such as serpentine, with levels of heavy metals that inhibit or preclude the growth of many plants and soil micro-organisms. However, certain plants and microorganisms do grow in these metalliferous sites. Understanding the physiology, ecology and evolution of tolerance to elevated soil metal concentrations is important in an applied setting, and is also of interest in theoretical biology. Applied importance relates to the improvement of forest health in areas subject to increasing pollution, rehabilitation of severely polluted sites by phytostabilization of metals, and metal removal using hyperaccumulating plants (Krämer, 2000; Ernst, 2000). Areas of theoretical interest include the evolution of local adaptation (Sork et al ., 1993) and how it is shaped by the combined influences of natural selection, gene flow and genetic architecture, as well as metal influences on various species interactions (Pollard, 2000). A paper appears on pages 367–379 in this issue by Jan Colpaert and coworkers which adroitly combines the disparate fields of physiology, genetics and ecology to answer several outstanding questions concerning heavy metal tolerance in mycorrhizal fungi.
Mycorrhizal fungi, which interact mutualistically with the majority of plant species, are well known for improving the P status of their hosts (Smith & Read, 1997). Some mycorrhizal fungi are also able to mobilize N and P from organic substrates and to provide plants with improved micronutrient and water acquisition, pathogen resistance, and a variety of other benefits (Smith & Read, 1997). One of these additional benefits is the amelioration of toxicity in metalliferous soils.     

Metal accumulation and tolerance in wetland plants

This paper briefly reviews the progress in studies of wetland plants in terms of heavy metal pollution. The current research mainly includes the following areas: (1) metal uptake, translocation, and distributions in wetland plants and toxicological effects on wetland plants, (2) radial oxygen loss (ROL) of wetland plants and its effects on metal mobility in rhizosphere soils, (3) constitutional metal tolerance in wetland plants, and (4) mechanisms of metal tolerance by wetland plants. Although a number of accomplishments have been achieved, many issues still remain unanswered. The future research effort is likely to focus on the ROL of wetland plants affecting metal speciation and bioavailability in rhizosphere soils, and the development of rhizosphere management technologies to facilitate and improve practical applications of phytoremediation of metalpolluted soils.     

Metal accumulation and tolerance in wetland plants

This paper briefly reviews the progress in studies of wetland plants in terms of heavy metal pollution. The current research mainly includes the following areas: (1) metal uptake, translocation, and distributions in wetland plants and toxicological effects on wetland plants, (2) radial oxygen loss (ROL) of wetland plants and its effects on metal mobility in rhizosphere soils, (3) constitutional metal tolerance in wetland plants, and (4) mechanisms of metal tolerance by wetland plants. Although a number of accomplishments have been achieved, many issues still remain unanswered. The future research effort is likely to focus on the ROL of wetland plants affecting metal speciation and bioavailability in rhizosphere soils, and the development of rhizosphere management technologies to facilitate and improve practical applications of phytoremediation of metal-polluted soils.     

When the same is not the same: phenotypic variation reveals different plant ecological strategies within species occurring in distinct Neotropical savanna habitats

Variations in abiotic characteristics such as soil water availability and fertility impose different selective pressures on plant populations. This may produce intraspecific variability in functional traits, even at a fine spatial scale. We investigated whether functional traits related to water-use efficiency, resource-retention strategy, soil nutrient acquisition, and fire tolerance differ in species that occur in two different habitats of Brazilian Cerrado: rocky savannas and savanna woodlands. Rocky savannas occur over sandstone, quartzite outcrops and have shallow nutrient-poor and low-moisture rocky soils, while savanna woodlands occur over well-drained and deep soils with frequent fire regimes. We measured nine functional traits of 40 tree species that occur in both habitats. Rocky savanna individuals exhibited a greater water-use efficiency strategy. The resource-retention strategy in rocky savanna individuals was corroborated by lower adult maximum height. However, despite the lower nutrient availability in rocky savanna soils, we only detected lower leaf phosphorus content in individuals from this habitat. Furthermore, individuals from both habitats had equally thick bark, suggesting that the fire-defense strategy is related to a stable, rather than plastic trait. Overall, our results highlight the central role of contrasting soil water availability patterns in driving phenotypic plasticity within species. We conclude that savanna species are responding to water and nutrient availabilities, via plasticity in traits related to the resource-retention strategy, and preparing for future fires, via uniformly thick bark. Wide plant distribution in contrasting habitats is possible for species that can shift ecological strategies to survive in nutrient- and water-limited habitats such as rocky savannas.