A comparison of published estimates of mycorrhizal and dark septate endophyte (DSE) colonisation from various ecosystems suggests that DSE may be as abundant as mycorrhizal fungi as judged by the proportion of host plants colonised in mixed plant communities, or by the extent of colonisation in sampled root systems. While many strides have been made in understanding the ecological significance of the mycorrhizal fungi, our knowledge about the role of DSE fungi is in its infancy. In order to provide a framework of testable hypotheses, we review and discuss the most likely functions of this poorly understood group of root-associated fungi. We propose that, like mycorrhizal symbioses, DSE-plant symbioses should be considered multifunctional and not limited to nutrient acquisition and resultant positive host growth responses. Admittedly, many mycorrhizal and endophyte functions, (e.g. stress tolerance, pathogen or herbivore deterrence) are likely to be mediated by improved nutritional status and increased fitness of the host. Accordingly, it is pivotal to establish whether or not the DSE fungi are involved in host nutrient acquisition, either from inorganic and readily soluble sources, or from organic and recalcitrant sources. Facilitation by DSE of the use of organic nitrogen, phosphorus and sulphur sources by plants is a topic that warrants further attention and research. Even in the absence of a clear nutrient uptake function, the extensive DSE colonisation that occurs is likely to pre-emptively or competitively deter pathogens by minimising the carbon available in host rhizosphere environment. The DSEs’ high melanin levels and their potential production of secondary metabolites toxic or inhibitory to herbivores are also likely to be factors influencing host performance. Finally, the broad host ranges speculated for most DSE fungi thus far suggest that they are candidates for controlling plant community dynamics via differential host responses to colonisation. We emphasise the need for simple experiments that allow unravelling of the basic biological functions of DSE fungi when they colonise their hosts

Plant tissues host a variety of fungi. One important group is the dark septate endophytes (DSEs) that colonize plant roots and form characteristic intracellular structures – melanized hyphae and microsclerotia. The DSE associations are common and frequently observed in various biomes and plant taxa. Reviews suggest that the proportion of plant species colonized by DSE equal that colonized by AM and microscopic studies show that the proportion of the root system colonized by fungi DSE can equal, or even exceed, the colonization by AM fungi. Despite the high frequency and suspected ecological importance, the effects of DSE colonization on plant growth and performance have remained unclear. Here, we draw from over a decade of experimentation with the obscure DSE symbiosis and synthesize across large bodies of published and unpublished data from Arabidopsis thaliana and Allium porrum model systems as well as from experiments that use native plants to better resolve the host responses to DSE colonization. The data indicate similar distribution of host responses in model and native plant studies, validating the use of model plants for tractable dissection of DSE symbioses. The available data also permit empirical testing of the environmental modulation of host responses to DSE colonization and refining the “mutualism-parasitism-continuum” paradigm for DSE symbioses. These data highlight the context dependency of the DSE symbioses: not only plant species but also ecotypes vary in their responses to populations of conspecific DSE fungi – environmental conditions further shift the host responses similar to those predicted based on the mutualism-parasitism-continuum paradigm. The model systems provide several established avenues of inquiry that permit more detailed molecular and functional dissection of fungal endophyte symbioses, identifying thus likely mechanisms that may underlie the observed host responses to endophyte colonization.

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Dark septate endophytes (DSE) are common and abundant root-colonizing fungi in the native tallgrass prairie. To characterize DSE fungi were isolated from roots of mixed tallgrass prairie plant communities. Isolates were grouped according to morphology, and the grouping was refined by ITS-RFLP and/or sequencing of the ITS region. Sporulating species of Periconia, Fusarium, Microdochium and Aspergillus were isolated along with many sterile fungi. Leek resynthesis was used to quickly screen for DSE fungi among the isolates. Periconia macro-spinosa and Microdochium sp. formed typical DSE structures in the roots; Periconia produced melanized intracellular microsclerotia in host root cortex, whereas Microdochium produced abundant melanized inter- and intracellular chlamydospores. To further validate the results of the leek resynthesis growth responses of leek and a dominant prairie grass, Andropogon gerardii, were assessed in a laboratory resynthesis system. Leek growth mainly was unresponsive to the inoculation with Periconia or Microdochium, whereas Andropogon tended to respond positively. Select Periconia and Microdochium isolates were tested further for their enzymatic capabilities and for ability to use organic and inorganic nitrogen sources. These fungi tested positive for amylase, cellulase, polyphenol oxidases and gelatinase. Periconia isolates used both organic and inorganic nitrogen sources. Our study identified distinct endophytes in a tallgrass prairie ecosystem and indicated that these endo-phytes can use a variety of complex nutrient sources, suggesting facultative biotrophic and saprotrophic habits.

Root colonization by arbuscular mycorrhizae (AM) and dark septate endophytic (DSE) fungi in nitrogen amended and unamended mixed tallgrass prairie communities were analyzed monthly over two growing seasons. Roots were stained with Trypan blue and Sudan IV and fungal structures quantified using the modified magnified intersections method. Root length colonized (RLC) by DSE exceeded AM colonization during early part of the growing season. Fungal colonization varied among the years and was greater in 2003 than in 2002. Seasonal variation among the months within a growing season was observed in 2002 but not in 2003 for both AM and DSE. AM fungi were most abundant during the peak growing season of dominant C4 vegetation while DSE were most abundant during the early part of the growing season. Hyperparasitism of AM hyphal coils by melanized septate fungi was frequently observed and increased with AM coil frequency. Although nitrogen amendment had altered the plant community composition, it had no impact on the colonization by AM or DSE fungi.