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6 Wood Discoloration

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3.8 Competition and Interactions Between Organisms 83<br />

Fig.3.5. Major forms of mycorrhizas.<br />

Ek ectotrophic, En endotrophic, VA<br />

vesicular-arbuscular (modified from<br />

Hock and Bartunek 1984)<br />

The ectotrophic mycorrhiza (ectomycorrhiza) occurs predominantly on<br />

conifers and hardwoods of the boreal and temperate zone, particularly associated<br />

with Pinales, Fagales and Salicales. In many conifers and in beech<br />

and oak, the association is obligatory, and in other trees like elms it is facultative<br />

(Müller and Löffler 1992). The predominant part of the mycelium grows<br />

at the surface of side roots and forms a dense mycelial coat at the root tips.<br />

The hyphae penetrate between the cells of the outer root tissue by dissolving<br />

the middle lamellae, and coat the cells completely as “Hartig net” (Kottke<br />

and Oberwinkler 1986). The colonized roots do no longer possess root hairs;<br />

instead hyphae or rhizomorphs radiate into the soil.<br />

In the endotrophic mycorrhiza (endomycorrhiza) of the orchids, only a loose<br />

hyphal net is formed around the root, and the hyphae settle inside the cells<br />

in the root bark area. As an intermediate, the ectendotrophic mycorrhiza is<br />

particularly present on roots of 1 to 3-year-old conifers, whereby the hyphae<br />

penetrating into the bark cells degenerate with ageing.<br />

The most frequent form, the vesicular arbuscular mycorrhiza (VAM) occurs<br />

associated with over 200,000 wild and cultivated angiosperms, in addition,<br />

with Ginkgo biloba, Taxus baccata and Sequoia gigantea and S. sempervirens<br />

(Werner 1987), as well as predominant form in tropical forests. In the VAM,<br />

the unseptate hyphae extend inside the root cells bubble-shaped (vesicles)<br />

or branch out tree-shaped (arbuscules). The arbuscules develop by hyphal<br />

branching and become enclosed by the peri-arbuscular membranes from the<br />

plant (Bothe and Hildebrandt 2003).<br />

The benefit for the trees is the improved nutrient (amino acids) and mineral<br />

(N, P, K, Mg, Cu, Zn, Fe) support and the better water supply (Smith and<br />

Read 1997) due to the larger absorption area. Soils with frequently occurring<br />

ectomycorrhiza are commonly characterized by a lower nutrient content, the<br />

trees growing there would not be competitive without mycorrhizas (Schönhar<br />

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