The evolutionary implications of exploitation in mycorrhizas1

Some views of mutualism, where the fitness of two symbiotic partners is higher in association than when apart, assume that they necessarily evolve towards greater benefit for the partners. Most mutualisms, however, seem prone to conflicts of interest that destabilize the partnership. These conflicts arise in part because mutualistic outcomes are conditional, depending upon complex interactions between environmental, developmental, and genotypic factors. Mutualisms are also subject to exploitation or cheating. Although various compensating mechanisms have been proposed to explain how mutualism can be maintained in the presence of exploiters, none of these mechanisms can eliminate exploitation. In this paper we explore various compensating mechanisms in mycorrhizas, examine the evidence for exploitation in mycorrhizas, and conclude that mycorrhizal mutualisms exhibit characteristics that are more consistent with a concept of reciprocal parasitism. We propose that researchers should not assume mycorrhizas are mutualistic based upon structural characteristics or limited functional studies showing bilateral exchange and should view mycorrhizas as occupying a wider range on the symbiotic continuum, including commensalism and antagonism. We recommend that comparative studies of mycorrhizas incorporate other types of root associations that have traditionally been considered antagonistic.