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Introduction to Fungi, Third Edition

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432 HYMENOASCOMYCETES: HELOTIALES (INOPERCULATE DISCOMYCETES)<br />

Fig15.2 Apothecia of Sclerotinia tuberosa rising from<br />

subterranean sclerotia formed on rhizomes of Anemone<br />

nemorosa.<br />

15.2.2 Monilinia fructigena and M. laxa<br />

Monilinia fructigena is the cause of a brown fruit<br />

rot of apples, pears, plums and other s<strong>to</strong>ne fruits<br />

(Byrde & Willetts, 1977). Although the apothecial<br />

state is only rarely formed, the disease is<br />

common and is transmitted by means of conidia.<br />

Apples and pears showing brown rot bear buffcoloured<br />

pustules of conidia often in concentric<br />

zones (Fig. 15.3a). Sporulation is stimulated by<br />

light, and adjacent zones correspond <strong>to</strong> daily<br />

periods of illumination. The conidia are blas<strong>to</strong>conidia<br />

formed in chains which extend in length<br />

at their apices by budding of the terminal conidium.<br />

Occasionally more than one bud is formed,<br />

and this results in branched chains (Fig. 15.3b).<br />

Conidiogenesis of this type is characteristic<br />

of the anamorphic genus Monilia. Infection of<br />

the fruit is commonly through wounds caused<br />

mechanically or by insects such as codling<br />

moth, wasps or earwigs (Croxall et al., 1951;<br />

Xu & Robinson, 2000). Fruits left lying on the<br />

ground are the source of infection in the following<br />

season. During the winter, infected fruits<br />

become mummified, and the shrivelled fruit<br />

thoroughly colonized by mycelium is interpreted<br />

as the sclerotium. In the following year the<br />

sclerotium may produce further conidial pustules.<br />

Infections can develop as a post-harvest<br />

disease in s<strong>to</strong>red apples, and in some varieties<br />

a twig infection (spur canker) may also occur.<br />

A similar group of diseases of apple and<br />

plum is caused by Monilinia laxa which also has<br />

a Monilia conidial state. In addition <strong>to</strong> fruit rot,<br />

this species causes blossom and shoot blight,<br />

in which infected fresh shoots wilt and become<br />

coated by conidial pustules. Monilinia fructicola<br />

causes brown rot especially of peaches and nectarines<br />

in North and South America, South Africa,<br />

Australia and the Far East, but has not been<br />

reported from Europe. It produces the apothecial<br />

state more readily than the other species (Holtz<br />

et al., 1998), and ascospores released from overwintered<br />

mummified fruits can be a source of<br />

inoculum in the field (Tate & Wood, 2000). These<br />

and a fourth species of the brown fruit rot<br />

complex, M. polystroma, can be distinguished by<br />

means of morphological features and DNA<br />

sequences (van Leeuwen et al., 2002).<br />

15.2.3 Sclerotinia sclerotiorum<br />

This species causes a range of diseases (Sclerotinia<br />

rot, white mould, stalk break) in over 400<br />

cultivated and wild plant species belonging <strong>to</strong><br />

some 75 different families (Boland & Hall, 1994).<br />

The most important crop plants affected are<br />

sunflower, soybean and oilseed rape, with crop<br />

losses approaching 100% under conditions favourable<br />

<strong>to</strong> the disease. Sclerotia are formed on<br />

decaying crop debris and remain viable in a<br />

dormant state in the soil for many years, especially<br />

if deep-ploughed. Sclerotia located within<br />

the <strong>to</strong>p 3 cm of soil germinate <strong>to</strong> produce hyphae<br />

or apothecia in spring. Plants can be infected<br />

either from mycelium or from ascospores; there is<br />

no macroconidial state. Phialidic microconidia<br />

are formed and probably serve as spermatia.<br />

Infection by S. sclerotiorum is often initiated by<br />

a saprotrophic phase on dead leaves or petals<br />

during which mycelial biomass is generated,<br />

prior <strong>to</strong> the attack on the living plant tissue.<br />

Above-ground shoots and, <strong>to</strong> a lesser extent, roots<br />

can be infected, and infections of living tissues<br />

are strongly necrotrophic. This necrotrophic<br />

phase is followed by further saprotrophic<br />

growth and the formation of sclerotia. The<br />

infection cycle in S. sclerotiorum is therefore<br />

tri-phasic. Reviews of the general biology and<br />

pathology of S. sclerotiorum have been written by

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