Exploring functional definitions of mycorrhizas: Are mycorrhizas always mutualisms?

Mycorrhizas are considered to be classic mutualisms. Here, we define mutualism as a reciprocal increase in fitness of the symbionts, and we review the evidence for mycorrhizal mutualism at the community, whole-plant, and cellular scales. It is difficult to use results of most mycorrhizal studies because (i) fungal contribution to nutrient uptake is not accurately estimated, (ii) increased growth is not necessarily correlated with increased plant fecundity or survival, especially in communities, and (iii) benefits that occur only at certain times of year, or under specific extreme conditions, may not be detected. To produce the nonmycorrhizal controls required to study mutualism in the field, soil microflora and fauna must be severely perturbed; therefore, it is virtually impossible to evaluate effects of mycorrhizas on plant fitness under realistic conditions. Using the evidence available, we conclude that mycorrhizas can occupy various positions along the continuum from parasitism to mutualism, depending on the specific plant and fungal genotypes and their abiotic and biotic environments. Although we discuss the possibility of defining mycorrhizas by some physiological characteristic, we conclude that mycorrhizas should be defined on a structural or developmental basis and that any requirement to demonstrate mutualism be eliminated.