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300<br />

Marjatta Raudaskoski, Mika Tarkka and Sara Niini<br />

the central vacuole becomes fragmented (Bonfante and Perotto 1995). These<br />

processes could also be controlled by rearrangements of MTs or MFs, or by<br />

both cytoskeletal elements.<br />

3.2 Fungal Hyphae<br />

In the research of cytoskeleton in endomycorrhiza, the structure and function<br />

of fungal MTs and MFs have gained less attention than those of <strong>plant</strong> cells. In<br />

the IIF microscopical investigation of nonsymbiotic hyphae of Glomus<br />

mossae, the MTs were visualized with tubulin and MFs with actin antibodies<br />

in the multinucleate hyphae originating from germinated spores (Åström et<br />

al. 1994). MTs extended in the cortical and central parts of the hyphae to the<br />

extreme hyphal tip, continued from the main hypha into a branch and the<br />

position of nuclei appeared to follow the MT tracks. Long MFs were also visualized<br />

in the hyphae (Åström et al. 1994). MTs and MFs could be involved in<br />

the intra- and intercellular nuclear movements and cytoplasmic streaming,<br />

respectively, recorded in living nonsymbiotic hyphae of different endomycorrhizal<br />

fungi (Bago et al. 1998; Giovannetti et al. 1999). The presence of both<br />

MTs and MFs in nonsymbiotic hyphae suggests that these structures could<br />

also play a significant role in hyphal morphogenesis associated with the formation<br />

of endomycorrhiza, such as the differentiation of the appressorium at<br />

the root <strong>surface</strong> at the beginning of the symbiosis and the formation of vesicular<br />

and arbuscular structures in the <strong>plant</strong> cell after the establishment of the<br />

symbiosis.<br />

4 Organization of Cytoskeleton in Ectomycorrhiza<br />

4.1 Root Cells<br />

The effect of ectomycorrhiza formation on the <strong>plant</strong> cell cytoskeleton is more<br />

difficult to investigate than that of endomycorrhiza or orchid mycorrhiza. In<br />

the ectomycorrhizal symbiosis, the fungal hyphae grow between the cortical<br />

cells of the host <strong>plant</strong>, forming a hyphal network for nutrient exchange called<br />

the Hartig net. The <strong>plant</strong> cells of the Hartig net have thick cell walls and accumulations<br />

of secondary metabolites such as phenols and starch. The thick cell<br />

walls inhibit rapid penetration of fixatives necessary for preservation of<br />

cytoskeletal elements, which is seen as lack of MTs or MFs from the published<br />

ultrastructural studies of ectomycorrhiza. Autofluorescence of secondary<br />

metabolites hampers the recording of cytoskeletal structures when they have<br />

been preserved during fixation. In spite of these difficulties, some knowledge<br />

of cytoskeletal structure has been obtained in Pinus sylvestris–Suillus bovinus<br />

ectomycorrhiza (Timonen et al. 1993; Niini and Raudaskoski 1998).

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