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6<br />

Ajit Varma et al.<br />

parasite on other fungi. Basidiomycetes, e.g., include saprobes, mycorrhizaforming<br />

fungi, <strong>plant</strong> parasites, but also fungi which are parasites of other<br />

fungi. Hosts are both Basidiomycetes and Ascomycetes. Ultrastructural investigations<br />

of this kind of organismic interaction (Chap. 16) revealed two main<br />

types, the formation of colacosomes and the fusion between pathogen and<br />

host fungus cells. Colacosomes are unique organelles, which appear at the<br />

interface between parasite and host while fusion is based on specialized interactive<br />

cells (haustoria), which establish a direct cytoplasmic connection.<br />

Many microorganisms coexist with <strong>plant</strong>s in ways that do not lead to <strong>plant</strong><br />

disease, symbiosis, or other specific interactions. Some fungi or bacteria can<br />

be latent pathogens. Some have little or no influence on the <strong>plant</strong>, but may<br />

form toxic compounds that are damaging to grazing animals. Microorganisms<br />

that form more or less benign associations with <strong>plant</strong>s are generally<br />

termed ‘endophytes’ and are genetically diverse. A large number of fungal<br />

endophytes can be difficult to identify because they include a high proportion<br />

with sterile mycelia (Chap. 17). Overall, the roles of many of these organisms<br />

are poorly understood.<br />

The mechanisms for entry of endophytic organisms into <strong>plant</strong>s can be<br />

investigated using methodologies such as those applied to elucidate the<br />

cytoskeletal rearrangements of <strong>plant</strong> cells and fungal hyphae at the <strong>plant</strong>–<br />

microbe interface during colonization of roots by mycorrhizal fungi (Chap.<br />

18). Invading organisms have been shown to influence the expression of <strong>plant</strong><br />

genes for some filamentous structures within the cell cytoskeleton. Indirect<br />

immunofluorescence microscopy has been used to investigate the cytoskeleton<br />

of some mycorrhizal associations demonstrating the separation and<br />

invagination of the plasma membrane from the <strong>plant</strong> cell wall in response to<br />

growth of fungi inside the cell wall.<br />

Colonization of <strong>plant</strong>s by related and unrelated groups of microorganisms<br />

may occur simultaneously. For example, saprophytes, pathogens and mycorrhizal<br />

fungi may be associated with the same root systems and colonize roots<br />

to different degrees.Several species of arbuscular mycorrhizal fungi can simultaneously<br />

colonize the same sections of root,although they are generally separated<br />

in different cells or parts of the root cortex. Prior colonization by one<br />

organism can influence sequential colonization by other organisms. This<br />

occurs to varying degrees for different groups of <strong>plant</strong> endophytes, symbionts<br />

and pathogens. The relative extent to which roots become colonized by several<br />

species of arbuscular mycorrhizal fungi present in the same soil depends on<br />

the relative abundance of propagules of the fungi in the soil,the developmental<br />

stage of the hyphae associated with fungal propagules, the susceptibility of the<br />

roots to invasion and the physiological responses of the root to different<br />

species of fungi (Chap. 19). Investigations of the molecular communication<br />

between these fungi and their host <strong>plant</strong>s during root colonization and nutrient<br />

acquisition are now beginning to be understood in terms of gene expression<br />

in <strong>plant</strong>s and fungi. This provides a basis for predicting physiological

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