Population genetics of ectomycorrhizal fungi: from current knowledge to emerging directions

Abstract

Ectomycorrhizal (EM) fungi are major microbial components of boreal, temperate and Mediterranean forests, as well as some tropical forest ecosystems. Nearly two decades of studies have clarified many aspects of their population biology, based on several model species from diverse lineages of fungi where the EM symbiosis evolved, i.e. among Hymenomycetes and, to a lesser extent, among Ascomycetes. In this review, we show how tools for individual recognition have changed, shifting from the use of somatic incompatibility reactions to dominant and non-specific markers (such as random amplified polymorphic DNA (RAPD) and amplified fragment length polymorphism (AFLP)) and, more recently, to co-dominant and specific markers (such as microsatellites and single nucleotide polymorphisms (SNPs)). At the same time, the theoretical focus has also changed. In earlier studies, a major aim was the description of genet size and popul/ation strategy. For example, we show how some studies supported or challenged the simple, classical model of colonization of new forest stands by ruderal (R) species, propagating by spores and forming small genets, progressively replaced in older forests by more competitive (C) species, propagating by mycelial growth and forming larger genets. By contrast, more recent studies give insights into some genetic traits, such as partners' assortment (allo- versus autogamy), genetic structure of populations and gene flow that turn out to depend both on distance and on whether spores are animal- or wind-dispersed. We discuss the rising awareness that (i) many morphospecies contain cryptic biological species (often sympatric) and (ii) trans- and inter-continental species may often contain several biological species isolated by distance. Finally, we show the emergence of biogeographic approaches and call for some aspects to be developed, such as fine-scale and long-term population monitoring, analyses of subterranean populations of extra-radical mycelia, or more model species from the tropics, as well as from the Ascomycetes (whose genetic idiosyncrasies are discussed). With the rise of the '-omics' sciences, analysis of population structure for non-neutral genes is expected to develop, and forest management and conservation biology will probably profit from published and expected work.