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Introduction to Fungi, Third Edition

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SCHIZOSACCHAROMYCETALES<br />

253<br />

about 5 a.m. and the spores appear mature by<br />

8 a.m. However, maximum spore discharge does<br />

not occur until about 8 p.m. Outside the ascus,<br />

ascospores or yeast cells may continue budding<br />

and the fungus can be grown saprotrophically<br />

as a yeast in agar or liquid culture. The yeast<br />

cells are often pigmented due <strong>to</strong> the presence of<br />

b-carotene and several other carotenoid pigments<br />

(van Eijik & Roeymans, 1982). Young<br />

leaves can be infected from such yeast cells and<br />

it has been shown that a culture derived from<br />

a single ascospore can cause infection resulting<br />

in the formation of a fresh crop of asci, so that<br />

T. deformans is homothallic. In this respect it differs<br />

from some other species, e.g. T. epiphylla,<br />

where the fusion of yeast cells, presumably of<br />

different mating types, is necessary before infection<br />

can occur (Kramer, 1987). In T. deformans the<br />

binucleate condition is established at the first<br />

nuclear division of a yeast cell placed on a peach<br />

leaf, and the two daughter nuclei remain associated<br />

in the germ tube which penetrates the<br />

cuticle (Fig. 9.2j). In other Taphrina species, the<br />

germ tube penetrates through s<strong>to</strong>mata but is<br />

unable <strong>to</strong> breach the intact cuticle (Taylor &<br />

Birdwell, 2000).<br />

9.2.2 Growth hormones<br />

In infections of peach leaves with T. deformans,<br />

the dis<strong>to</strong>rtions of the host tissue are associated<br />

with division and hypertrophy of the cells of the<br />

palisade mesophyll. In liquid cultures, especially<br />

on media containing tryp<strong>to</strong>phane, considerable<br />

quantities of the auxin-type phy<strong>to</strong>hormone<br />

indole acetic acid (IAA) have been demonstrated.<br />

A number of different cy<strong>to</strong>kinins are also<br />

produced by several species of Taphrina in culture<br />

(Kern & Naef-Roth, 1975; Tudzynski, 1997).<br />

Together, these hormones promote processes of<br />

cell division, enlargement and differentiation in<br />

plants, and leaves infected with T. deformans<br />

show higher levels of auxins and cy<strong>to</strong>kinins<br />

than uninfected leaves (Sziráki et al., 1975). It is<br />

therefore tempting <strong>to</strong> assume that the fungus<br />

produces these substances also in planta.<br />

However, this has not been formally proven<br />

yet, and the Taphrina peach system seems <strong>to</strong><br />

have been less thoroughly examined than the<br />

interaction between Plasmodiophora brassicae and<br />

cabbage plants (see p. 63).<br />

9.2.3 Control of Taphrina deformans<br />

Taphrina deformans is by far the most serious<br />

pathogen among the Taphrinales, and it occurs<br />

wherever peach or almond trees grow. It is not<br />

yet entirely clear how T. deformans overwinters;<br />

Butler and Jones (1949) and Smith et al. (1988)<br />

considered it unlikely that the mycelial form is<br />

involved because leaves harbouring mycelium<br />

are shed in the autumn. It is more probable<br />

that yeast cells arising from discharged ascospores<br />

survive saprotrophically on the surface of<br />

twigs or in bud scales. Between November and<br />

March, the yeast cells develop thick walls and in<br />

spring, as the peach buds open, they produce<br />

germ tubes which penetrate the young leaves.<br />

The first symp<strong>to</strong>ms of infection can be seen as<br />

soon as the buds break, but no further infection<br />

occurs from about early July onwards. This may<br />

be because T. deformans has a relatively low<br />

temperature maximum of 26 30°C (Butler &<br />

Jones, 1949).<br />

Good chemical control of T. deformans can<br />

be achieved by spraying with Bordeaux mixture<br />

in autumn after leaf fall, in order <strong>to</strong> reduce the<br />

population of yeast cells on the twigs. Another<br />

spray in early spring, at the time of bud swelling,<br />

will give improved control of infection because<br />

the time span in which Taphrina can infect<br />

is limited. Dithiocarbamates or other simple<br />

fungicides are commonly used in spring<br />

(Smith et al., 1988). Other diseases caused by<br />

Taphrina can be controlled in a similar way if<br />

necessary.<br />

9.3 Schizosaccharomycetales<br />

The classification of the Schizosaccharomycetales<br />

has been the subject of controversial<br />

discussions, but the emerging consensus is<br />

that there is only one genus with three species,<br />

S. japonicus, S. oc<strong>to</strong>sporus and S. pombe (Kurtzman<br />

& Robnett, 1998; Vaughan-Martini & Martini,<br />

1998a; Barnett et al., 2000). All three species grow

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