Bryophytes, including the mosses, liverworts, and hornworts, occur in a variety of habitats with high concentrations of metals and have other characteristics that are advantageous for studies of metal tolerance. Mosses may evolve genetically specialized, metal-tolerant races less frequently than flowering plants. Some species of mosses appear to have inherently high levels of metal tolerance even in individuals that have not been subjected to natural selection in contaminated environments. Scopelophila cataractae, one of the so-called copper mosses, not only tolerates extremely high concentrations of metals in its substrates, but requires these substrates for optimum growth. This species should be included in mechanistic studies of tolerance at the cellular and molecular levels. - 274-3474. evolve in just a few generations, and high levels of tolerance tend to be metal-specific. Tolerance mechanisms are varied and the genetic basis for infraspecific differences in tolerance may also be varied. Major genes with modifiers seem to be involved in several species but additional genetic analyses are needed.
Tolerance in Widespread MossesMany species of bryophytes are known to tolerate habitats with high concentrations of environmental pollutants (7) and the effects of metals such as copper, zinc, and lead have been most thoroughly studied under experimental conditions. Genetically specialized ecotypes tolerant of copper have been described in the liverwort, Solenostomum crenulatum (8) Other studies on several species of mosses suggest that in addition to, or sometimes instead of, the evolution of genetically adapted ecotypes, inherently broad tolerances permit plants to colonize and thrive in metal-contaminated environments. Shaw (9) studied tolerance of copper, zinc, cadmium, and nickel in plants from eight populations of Funaria hygrometrica. There was strong evidence of ecotypic differentiation for copper tolerance; populations could be clearly segregated as either tolerant or relatively nontolerant. Tolerant plants were restricted to copperenriched sites. These observations provide solid, if indirect, evidence of responses to natural selection for tolerance at contaminated sites.The same populations were genetically variable with regard to cadmium and zinc tolerance, but the relationships between tolerance levels and concentrations of these metals in the natural habitats of populations were less clear than for copper. For cadmium, plants from one population that originated at a cadmium-enriched site were indeed relatively tolerant of this metal, but plants from several other populations growing at sites without cadmium enrichment were also tolerant. Plants that were tolerant of copper also tended to be tolerant of cadmium, even if cadmium was not present in high concentration in their substrates. This observation suggests cotolerances between metals, with obvious implications for mechanistic studies.Plants also exhibited variation in tolerance of zinc, but there appeared to be no relationship between toleran...