The genetics of metal tolerance in vascular plants.

In many parts of the world soils are detrimental to plant growth owing to elevated levels of metal ions, caused either by natural processes or by the result of man's activities. Many plants have evolved ecotypes or varieties that are able to grow more-or-less normally on these soils. This paper reviews our knowledge of the genetics of this phenomenon. The nature of tolerance and the problems of its measurement are discussed. Tolerance is frequently measured by an index produced by comparing growth in a contaminated environment with growth in a control environment. It is argued that this measurement is inappropriate for many genetical studies, and that it is frequently more useful to use growth at a single critical level of metal as a measure of tolerance. Polygenic inheritance provides a null hypothesis that has to be tested in a genetical analysis. Examples of major genes for tolerance to aluminium, arsenic, boron, cadmium, copper and manganese are discussed. Even where major genes have been demonstrated, it is probable that other minor genes, 'modifiers', are present as well. Because of the nature of tolerance as a character, dominance and epistasis are likely to vary with the level of metal at which an analysis is performed. Tolerance is generally found to be dominant at some levels of the metal. Studies which have mapped tolerance genes, particularly to aluminium and salt, are discussed. The specificity of tolerance is a matter of some confusion. Some studies indicate that tolerances evolve independently to different metals, but others have suggested that tolerance to one metal may often confer a degree of tolerance to some other metals. Very little is known about the molecular genetics of tolerance, and the mechanisms of tolerance to most metals. The possible role of phytochelatins and metallothionein-like proteins in metal tolerance is discussed. The distribution of tolerance in natural populations suggests that tolerance is a disadvantage in uncontaminated environments, but how this 'cost' arises is not known. There is some evidence that the disadvantage to tolerance may be associated more with the modifiers of tolerance than with the primary tolerance gene. The study of the genetics of tolerance is of importance in planning breeding programmes to produce tolerant crops for use in areas where metal contamination is a limiting factor in productivity. It can also assist in understanding the mechanisms of tolerance, as exemplified by the study of the mechanism of arsenic tolerance in Holcus lanatus. Important areas for further research are discussed. Contents Summary 541 I. Introduction 542 II. Introduction 542 III. Transmission genetics of tolerance 544 IV. Specificity of tolerance 550 V. Molecular genetics of tolerance 552 VI. Ecological genetics of tolerance 553 VII. Conclusions 555 Acknowledgements 556 References 556.