Plant defense using toxic inorganic ions: conceptual models of the defensive enhancement and joint effects hypotheses

The concept of plant defense using toxic mineral elements originated as an explanation for extremely elevated concentrations of some elements (termed hyperaccumulation) in some plant tissues. The Defensive Enhancement Hypothesis suggests that hyperaccumulation evolved because, after an initial defensive benefit accrued from a relatively low initial concentration, increased concentration of an element provided increased plant fitness and drove evolution of higher element concentrations until hyperaccumulation was achieved. The Joint Effects Hypothesis postulates that additive or synergistic effects between element-based defenses, or between toxic element and organic chemical defenses, may have contributed to the evolution of hyperaccumulation. By lessening the concentration of an element necessary to provide an initial defensive benefit to a plant, joint effects could decrease the level of an element that provides an initial defensive benefit, allowing additive or synergistic defensive enhancement to take effect. Recent experimental tests have demonstrated defense at relatively low element concentrations, and tests of metal/metal and metal/organic compound combinations have shown joint effects. These hypotheses suggest how hyperaccumulator plants may have evolved in response to plant-herbivore interactions, and suggest that toxic element levels below those used to define hyperaccumulation may be ecologically effective.