BACKGROUNDPlants that accumulate metal and metalloid trace elements to extraordinarily high concentrations in their living biomass have inspired much research worldwide during the last decades. Hyperaccumulators have been recorded and experimentally confirmed for elements such as nickel, zinc, cadmium, manganese, arsenic and selenium. However, to date, hyperaccumulation of lead, copper, cobalt, chromium and thallium remain largely unconfirmed. Recent uses of the term in relation to rare-earth elements require critical evaluation. SCOPE Since the mid-1970s the term 'hyperaccumulator' has been used millions of times by thousands of people, with varying degrees of precision, aptness and understanding that have not always corresponded with the views of the originators of the terminology and of the present authors. There is therefore a need to clarify the circumstances in which the term 'hyperaccumulator' is appropriate and to set out the conditions that should be met when the terms are used. We outline here the main considerations for establishing metal or metalloid hyperaccumulation status of plants, (re)define some of the terminology and note potential pitfalls. CONCLUSIONS Unambiguous communication will require the international scientific community to adopt standard terminology and methods for confirming the reliability of analytical data.
SUMMARYHeavy metal uptake, accumulation and tolerance were investigated in five British populations of the metallophyte Thlaspi caerulescens from metalliferous sites from the north and south Pennines orefields. Analysis of field samples sbowed mean shoot Zn, Pb and Cd concentrations of up to 21 000, 660 and 164 //g g"' respectively, A solution culture experiment designed to investigate both tolerance and metal accumulation is reported. Indices of tolerance of five populations to 12 metals (Ag, A!, Cd, Co, Cr, Cu, Fe, Mn, Mo, Ni, Pb, Zn) showed few population differences but unexpectedly high tolerance to metals not present at elevated concentrations in the parent soils. This u-as paralleJed b> exceptionaWy high upt;ikes of all metals studied. Zn, Cd, Co, Mn and Ni were readily transported to the shoot whereas Al, Cr Cu, Fe and Pb were predominantly immobilized in the roots. The data suggest common mechanisms of absorption and transport of several metals in this species.
Plants that have evolved to survive on metal-rich soilsmetallophytes-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?
Understanding the relative importance of the abiotic environment and species interactions in determining the distribution and abundance of organisms has been a challenge in ecological research. Serpentine substrata are stressful environments for plant growth due to multiple limitations, collectively called the "serpentine syndrome". In the present review, our aim is not only to describe recent work in serpentine ecology, but also to highlight specific mechanisms of species tolerance and adaptation to serpentine soils and their effects on community structure and ecosystem functioning. We present hypotheses of the development of serpentine endemism and a description of functional traits of serpentine plants together with a synthesis of species interactions in serpentine soils and their effects on community structure and ecosystem productivity. In addition, we propose hypotheses about the effects of the 'serpentine syndrome' on ecosystem processes including productivity and decomposition.
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