15 Fruiting Body Formation in Basidiomycetes
Establishment of the dikaryotic mycelium and formation of fruiting bodies are highly complex developmental programmes that are activated by a combination of environmental cues. A wide variety of proteins are expected to regulate and coordinate these progr
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Fruiting Body Formation in Basidiomycetes
JORDI F. PELKMANS1, LUIS G. LUGONES1, HAN A.B. WO¨STEN1
CONTENTS I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . II. Development of Emergent Structures . . . . . . . III. Regulation of Fruiting Body Formation . . . . . A. Environmental Signals . . . . . . . . . . . . . . . . . . . . B. Mating-Type Genes as Master Regulators . C. Other Regulatory Genes . . . . . . . . . . . . . . . . . . . D. Nuclear Positioning . . . . . . . . . . . . . . . . . . . . . . . IV. Structural Proteins and Enzymes Involved in Fruiting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. Hydrophobins . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. Lectins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. Haemolysins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. Oxidative Enzymes . . . . . . . . . . . . . . . . . . . . . . . E. Expansins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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I. Introduction Fruiting bodies have evolved in the fungal kingdom to produce and disperse sexual spores. These reproductive structures of the Homobasidiomycota generally contain specialised cells, basidia, in which the genetically different parental haploid nuclei fuse. The resulting diploid cells immediately undergo meiosis to form haploid basidiospores. In some cases, diploid nuclei are already formed in the vegetative mycelium, as in Armillaria species (Ullrich and Anderson 1978; Grillo et al. 2000). After discharge, germination of the haploid basidiospores results in homokaryotic mycelia. These mycelia can fuse and, depending on an often complex system of mating-type genes 1
Microbiology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands; e-mail: [email protected]
(see Freihorst et al. 2016), produce a heterokaryotic mycelium that is fertile. Consequently, it forms fruiting bodies under appropriate environmental conditions. Basidiomycetes behaving according to this scheme are in the majority and are called heterothallic (i.e. selfincompatible). From a teleological point of view, this makes sense because it ensures that the diploid basidia produce recombinant meiotic progeny. A minority of basidiomycetes (about 10 %; Whitehouse 1949) are homothallic (i.e. self-compatible). Recently, it was shown that unisexual reproduction also creates diversity (Ni et al. 2013), which would explain the existence of homothallic species from an evolutionary point of view. This review discusses the regulation of fruiting body formation in homobasidiomycetes and the role structural proteins and enzymes play in this process. Related topics such as morphogenesis, cytology and mathematical modelling are discussed in Wells and Wells (1982), Moore et al. (1985), Wes
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