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Growth, Differentiation and Sexuality

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14 Asexual Sporulation in Mycelial Fungi<br />

R. Fischer 1 ,U.Kües 2<br />

CONTENTS<br />

I. Introduction ......................... 263<br />

II. Spore Formation ...................... 263<br />

A. VariabilityofSpores<br />

<strong>and</strong>SporeProduction ............... 263<br />

B. Ecological Aspects of Spore Production . . 264<br />

C. Sexual Versus Asexual Reproduction . . . . 266<br />

III. Endogenous <strong>and</strong> Environmental Factors<br />

Trigger Spore Formation ................ 266<br />

IV. Genetics of Asexual Spore Formation ...... 268<br />

A. Aspergillus nidulans ................. 268<br />

1.Cytology........................ 268<br />

2.Genetics........................ 268<br />

a) Seven-Transmembrane<br />

<strong>and</strong> Other Signal Receptors . . . . . . 269<br />

b) HeterotrimericG-Proteins ....... 270<br />

c) MAP-Kinase <strong>and</strong> Other Signalling<br />

Modules ..................... 271<br />

d) cAMPSignalling............... 272<br />

e) SmallG-Proteins............... 272<br />

f) Transcription Factors<br />

<strong>and</strong>OtherRegulators ........... 273<br />

g) TargetGenes.................. 275<br />

h) Interdependence of Vegetative<br />

<strong>Growth</strong><strong>and</strong>Development........ 275<br />

B. OtherAscomycetes.................. 276<br />

C. Basidiomycetes..................... 280<br />

1.Cytology........................ 280<br />

2.Genetics........................ 281<br />

D. ConservedGeneticPathways .......... 283<br />

V. Concluding Remarks ................... 284<br />

References ........................... 285<br />

I. Introduction<br />

Thefilamentousgrowthformistypicalformany<br />

fungi. By indefinite polarized growth, fungi are able<br />

to populate enormous areas of a substrate, even<br />

though most of them are non-motile organisms.<br />

Renowned are the gigantic Armillaria individuals<br />

covering several hundreds of hectares of for-<br />

1 Max-Planck-Institute for Terrestrial Microbiology, Marburg, <strong>and</strong><br />

Institute for Applied Life Sciences, Applied Microbiology, University<br />

of Karlsruhe, Hertzstr. 16, 76187 Karlsruhe, Germany<br />

2 Institut für Forstbotanik, Abteilung Molekulare Holzbiotechnologie,<br />

Georg-August-Universität Göttingen, Büsgenweg 2, 37077 Göttingen,<br />

Germany<br />

est grounds in Northern America (Ferguson et al.<br />

2003). However, growth of hyphae requires constant<br />

feeding with organic matter from the environment,<br />

although the mycelium can cope with short<br />

distances of nutrient-depleted areas by means of<br />

hyphal transport (Cairney 2005). One strategy with<br />

which sessile fungi adapt to conditions of limited<br />

resources at a given place is spore formation. Typically,<br />

different types of spores, sexually or asexually<br />

derived, are generated, depending on the purpose<br />

of the spores. Asexual spores are obtained from mitosis<br />

(mitospores). Usually, they are produced in<br />

high numbers as genetically identical propagules<br />

of the individual they come from. Various modes<br />

of production <strong>and</strong> different types of asexual spores<br />

have evolved in fungi, emphasizing the high significance<br />

they have within fungal life cycles, for fungal<br />

life styles <strong>and</strong> for survival in the environment.<br />

Often, asexual spores fulfil not only one distinct<br />

function but have several different roles. Asexual<br />

spores can serve as a means of reproduction <strong>and</strong><br />

dispersal to new substratum, for the purpose of<br />

mating with other individuals <strong>and</strong> for perennation<br />

as resting bodies under unfavourable conditions<br />

(Kendrick 1979a,b; Kirk et al. 2001; see also Andrews<br />

<strong>and</strong> Harris, The Mycota, Vol. IV, Chap. 1).<br />

II. Spore Formation<br />

A. Variability of Spores <strong>and</strong> Spore Production<br />

Asexual fungal spores are produced by different<br />

mechanisms (Kendrick 1979a,b; Esser 2001; Kirk<br />

et al. 2001).<br />

1. In thallic spore formation, arthrospores or<br />

oidia are generated by fragmentation of<br />

pre-existing hyphal filaments. If production<br />

of a specific hyphal branch directly precedes<br />

fragmentation, spores are also called arthroconidia.<br />

The Mycota I<br />

<strong>Growth</strong>, Differentation <strong>and</strong> <strong>Sexuality</strong><br />

Kües/Fischer (Eds.)<br />

© Springer-Verlag Berlin Heidelberg 2006

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