Chapter 1 The Biosphere Chapter 2 The Anthroposphere Introduction Air Pollution Water Pollution Soil Plants Chapter 3 Soils and Soil Processes Introduction Weathering Processes Pedogenic Processes Chapter 4 Soil Constituents Introduction Trace Elements Minerals Organic Matter Organisms in Soils Chapter 5 Trace Elements in Plants Introduction Absorption Translocation Availability Essentiality and Deficiency Toxicity and Tolerance Speciation Interaction Chapter 6 Elements of Group 1 (Previously Group Ia) Introduction Lithium Rubidium Cesium Chapter 7 Elements of Group 2 (Previously Group IIa) Beryllium Strontium Barium Radium Chapter 8 Elements of Group 3 (Previously Group IIIb) Scandium Yttrium Lanthanides Actinides Chapter 9 Elements of Group 4 (Previously Group IVb) Titanium Zirconium Hafnium Chapter 10 Elements of Group 5 (Previously Group Vb) Vanadium Niobium Tantalum Chapter 11 Elements of Group 6 (Previously Group VIb) Chromium Molybdenum Tungsten Chapter 12 Elements of Group 7 (Previously Group VIIb) Manganese Technetium Rhenium Chapter 13 Elements of Group 8 (Previously Part of Group VIII) Iron Ruthenium Osmium Chapter 14 Elements of Group 9 (Previously Part of Group VIII) Cobalt Rhodium Iridium Chapter 15 Elements of Group 10 (Previously Part of Group VIII) Nickel Palladium Platinum Chapter 16 Elements of Group 11 (Previously Group Ib) Copper Silver Gold Chapter 17 Trace Elements of Group 12 (Previously of Group IIb) Zinc Cadmium Mercury Chapter 18 Elements of Group 13 (Previously Group IIIa) Boron Aluminum Gallium Indium Thallium Chapter 19 Elements of Group I4 (Previously Group IVa) Silicon Germanium Tin Lead Chapter 20 Elements of Group 15 (Previously Group Va) Arsenic Antimony Bismuth Chapter 21 Elements of Group 16 (Previously Group VIa) Selenium Tellurium Polonium Chapter 22 Elements of Group 17 (Previously Group VIIa) Fluorine Chlorine Bromine Iodine

Trace elements in soils and plants

Our understanding of plant mineral nutrition comes largely from studies of herbaceous crops that evolved from ruderal species characteristic of nutri­ ent-rich disturbed sites (52). With the development of agriculture, these ancestral species were bred for greater productivity and reproductive output at high nutrient levels where there was little selective advantage in efficient nutrient use. This paper briefly reviews the nature of crop responses to nutrient stress and compares these responses to those of species that have evolved under more natural conditions, particularly in low-nutrient envi­ ronments. I draw primarily upon nutritional studies of nitrogen and phos­ phorus because these elements most commonly limit plant growth and because their role in controlling plant growth and metabolism is most clearly understood (51). Other more specific aspects of nutritional plant ecology not discussed here include ammonium/nitrate nutrition (79), cal­ cicole/calcifuge nutrition (51,88), heavy metal tolerance (4), and serpentine ecology (133).

The Mineral Nutrition of Wild Plants

Fifteen or more elements present in rocks and soils normally in very small amounts are essential for plant and/or animal nutrition. By the nature of their low abundance in natural uncontaminated earth materials or plants, they are known as trace elements, minor elements or micro-nutrients. Boron, copper, iron, manganese, molybdenum, silicon, vanadium and zinc are required by plants; copper, cobalt, iodine, iron, manganese, molybdenum, selenium and zinc by animals. In addition essential roles of arsenic, fluorine, nickel, silicon, tin and vanadium have in recent years been established in animal nutrition.

Trace Elements in Soils and Plants

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.

Accumulators and excluders ?strategies in the response of plants to heavy metals

Zinc (Zn) is an essential component of thousands of proteins in plants, although it is toxic in excess. In this review, the dominant fluxes of Zn in the soil-root-shoot continuum are described, including Zn inputs to soils, the plant availability of soluble Zn(2+) at the root surface, and plant uptake and accumulation of Zn. Knowledge of these fluxes can inform agronomic and genetic strategies to address the widespread problem of Zn-limited crop growth. Substantial within-species genetic variation in Zn composition is being used to alleviate human dietary Zn deficiencies through biofortification. Intriguingly, a meta-analysis of data from an extensive literature survey indicates that a small proportion of the genetic variation in shoot Zn concentration can be attributed to evolutionary processes whose effects manifest above the family level. Remarkable insights into the evolutionary potential of plants to respond to elevated soil Zn have recently been made through detailed anatomical, physiological, chemical, genetic and molecular characterizations of the brassicaceous Zn hyperaccumulators Thlaspi caerulescens and Arabidopsis halleri.

Zinc in plants

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.

R.G. Turner

Papers

Heavy metal tolerance in plants