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spore germination, and also containing a dye
(bromocresol purple). For application to stump surfaces,
the sachets were diluted with water, allowing the
spores to germinate, and the dye enabled foresters
to see that the stump surfaces had been treated.
More recently, the spores have been produced as dry
powder formulations by commercial companies.
This biocontrol system proved highly effective in pine
forests over much of Europe and North America. But
in Britain it had one significant limitation – P. gigantea
could be used only to protect pine stumpsand was
not effective in protecting Sitka spruce, which is the
most common plantation tree in Britain. Now it seems
that there is a simple explanation for this: the strains
of P. giganteaused to control the disease in Britain are
endemic strains, specialized to colonize pine stumps,
whereas the strains used in Scandinavia have a broader
host range and can protect spruce as well as pines.
In Finland a commercial preparation of P. gigantea,
marketed as Rotstop™, effectively controls H. annosum
on spruce. The UK Forestry Commission is currently
investigating the possibility of introducing such strains
into Britain.

Mycoparasites: fungi that parasitize other
fungi

The fungi that parasitize other fungi can be grouped
into two broad categories: necrotrophic mycoparasites,
which invade and destroy other fungal cells and then
feed on the dead cell contents, and biotrophic myco-
parasites, which can establish a specialized feeding
relationship, usually by producing haustoria to pene-
trate and absorb nutrients from living fungal hyphae.
These two types of mycoparasite are equivalent to the
necrotrophic and biotrophic parasites of plants, dis-
cussed in Chapter 14.

Biotrophic mycoparasites

There are several types of biotrophic mycoparasite
with different feeding mechanisms ( Jeffries & Young
1994), but the most common and distinctive group are
the haustorial biotrophs. These fungi penetrate living
host hyphae to produce a haustorium inside the host
wall but separated from the cell contents by a host cell

244 CHAPTER 12

Fig. 12.9Possible mode of action of Phlebiopsis giganteain controlling Heterobasidion annosumin recently felled pine
stumps. 1, P. giganteais inoculated onto freshly exposed stump surfaces and prevents colonization of the stumps from
airborne basidiospores of H. annosum(shown by the double lines). 2, When P. giganteahas colonized the dying stump
and root tissues it prevents the spread of H. annosumfrom existing foci of infection in the root zone. 3, P. gigantea
might also prevent H. annosumfrom growing up to the stump surface to sporulate.

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