Growth, Differentiation and Sexuality

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Signals in Growth and Development
11 Autoregulatory Signals in Mycelial Fungi U. Ugalde 1 CONTENTS I. Introduction ......................... 203 II. Germination ......................... 204 III. Colony Morphogenesis ................. 205 IV. Asexual Development .................. 207 V. Sexual Development ................... 208 VI. Dimorphism ......................... 210 VII. Conclusions .......................... 210 References ........................... 211 I. Introduction The mycelium is a very successful colony form capable of orchestrating the development of different cell types in response to the microenvironments which it encounters. This attribute in particular has led to the realisation that the mycelium is governed by a sophisticated chemosensitive system.Manyoftheelementsofthissystemhaveprobably been already inventoried after several decades of intense research, but the functional connections between them remain largely undiscovered. On the one hand, many reported colony functions have been ascribed to as yet unidentified chemical signals, whilst on the other, many thousands of molecules have been purified and characterised, of which relatively few have been credited with a biological role. Those signals recognised so far comprise two large categories. First, those directed at relations with other organisms, such as hosts and including elicitors and inhibitors, competitors involving antibiotics, and predators where mycotoxins are implicated (for further information, see The Mycota, Vols. VA, X and XI). The second category consists of in-house signals which formerly fell under the generic term 1 Biochemistry Laboratory II, Faculty of Chemistry, University of The Basque Country, P.O. Box 1072, 20080 Donostia-San Sebastian, Spain hormones (Gooday 1994), and are now separately considered. They include pheromones, which facilitate the interaction of compatible gametes, and developmental hormones regulating the formation or maintenance of differentiated multicellular structures or protoorgans, such as ascomata or basidiomata. These will not be dealt with in this chapter, but individually in Chaps. 12 (Pheromone action in the fungal groups Chytridiomycota, and Zygomycota, and in the Oomycota), 17 (Mating type genes of the basidiomycetes) and 16 (Fruiting body development in ascomycetes) of this volume respectively. In addition to these important in-house signals, there is an array of endogenous cues which convey information on environmental conditions or the status of cells within the mycelium, thus ensuring a coordinated colony function. They are transmitted by extracellular metabolites, and their signalling action results in adaptations which are advantageous to the colony as a living unit. In the past, these signals have been referred to as autoinducers, autoinhibitors, quorum sensing factors and morphogens. From a functional viewpoint, they all share the feature of regulating transitions between alternative morphogenetic and/or functional programmes, hence the proposal to term them collectively as autoregulators. Signallingmoleculesperforming equivalent tasks had already been assigned this term when first discovered in filamentous bacteria (Horinouchi and Beppu 1992). However, the range of activities covered in mycelial fungi can be presumed to be wider and more complex in light of the information now available on secondary metabolites and fungal development (Calvo et al. 2002). This review will focus on those autoregulatory signals which are involved in germination, colony morphogenesis, and asexual and sexual development in mycelial fungi. Finally, the incidence of autoregulators in dimorphism will also be covered. The Mycota I Growth, Differentation and Sexuality Kües/Fischer (Eds.) © Springer-Verlag Berlin Heidelberg 2006
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The Mycota Edited by K. Esser
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The Mycota A Comprehensive Treatise
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Karl Esser (born 1924) is retired P
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VIII Series Preface Class: Saccharo
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Addendum to the Series Preface In e
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XIV Volume Preface to the First Edi
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XVI Volume Preface to the Second Ed
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XVIII Contents Reproductive Process
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XX List of Contributors André Flei
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Vegetative Processes and Growth
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4 K.J. Boyce and A. Andrianopoulos
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22 L.J. García-Rodríguez et al. I
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24 L.J. García-Rodríguez et al. F
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26 L.J. García-Rodríguez et al. 2
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28 L.J. García-Rodríguez et al. M
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30 L.J. García-Rodríguez et al. a
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32 L.J. García-Rodríguez et al. t
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34 L.J. García-Rodríguez et al. H
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36 L.J. García-Rodríguez et al. T
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38 S.D. Harris dle organization tha
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40 S.D. Harris 2001; Borkovich et a
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42 S.D. Harris terized in A. nidula
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44 S.D. Harris astral microtubules
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46 S.D. Harris entry, and the CDK N
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48 S.D. Harris and Hamer 1997), the
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50 S.D. Harris Murray AW (2004) Rec
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4 Apical Wall Biogenesis J.H. Siets
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fungi can be regarded as “tube-dw
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In agreement with an essential role
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Kopecka and Gabriel 1992). They als
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nearly all the label was present in
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ingredients such as wall components
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in membrane enlargement and exocyto
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sis occurs, coinciding with a gradi
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pling of two (1-3)-alpha-glucan seg
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Sietsma JH, Wessels JGH (1977) Chem
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5 The Fungal Cell Wall J.P. Latgé
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polymers (chitosan) and glucuronic
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Whereas the structural branched β1
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their structural role in the cell w
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eight chitin synthases of A. fumiga
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Characterization of the chs4 mutant
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Fig. 5.8. Experimental data and hyp
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Fig. 5.9. Elongation of the mannan
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end of linear β1,3 glucans, and tr
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Fig. 5.12. Signal transduction in f
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shock,lowosmolarityaswellasotherfac
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ditions. Accordingly, enzymes and r
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Calonge TM, Arellano M, Coll PM, Pe
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Hiura N, Nakajima T, Matsuda K (198
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Martin-Yken H, Dagkessamanskaia A,
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Saporito-Irwin SM, Birse CE, Sypher
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6 Septation and Cytokinesis in Fung
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Septation in Fungi 107 Table. 6.1.
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Fig. 6.1. Selection of a cell divis
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Calderone, Chap. 5, this volume, an
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permissive temperature, multiple se
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eports suggested such a function fo
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understood mechanism of mitotic exi
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Implication in cytokinesis in Sacch
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Trinci APJ, Morris NR (1979) Morpho
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124 N.L. Glass and A. Fleissner ter
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126 N.L. Glass and A. Fleissner 188
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128 N.L. Glass and A. Fleissner Fig
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130 N.L. Glass and A. Fleissner sub
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132 N.L. Glass and A. Fleissner dur
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134 N.L. Glass and A. Fleissner G1
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136 N.L. Glass and A. Fleissner Bri
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138 N.L. Glass and A. Fleissner Mat
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8 Heterogenic Incompatibility in Fu
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Fungal Heterogenic Incompatibility
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B. Genetic Control The genetic back
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active phenotype het-s. The neutral
Page 164 and 165: Table. 8.1. (continued) Fungal Hete
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Page 170 and 171: ility controlled by multiple allele
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Page 176 and 177: Semi-Incompatibilität. Z Indukt Ab
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Page 180 and 181: Vilgalys RJ, Miller OK (1987) Matin
Page 182 and 183: 168 B.C.K. Lu (Esser et al. 1980; K
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Page 186 and 187: 172 B.C.K. Lu et al. 2004). It is l
Page 188 and 189: 174 B.C.K. Lu (MMP), through ruptur
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Page 192 and 193: 178 B.C.K. Lu drial fission during
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Page 202 and 203: 10 Senescence and Longevity H.D. Os
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Page 208 and 209: Fig. 10.3. Copper delivery to the c
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Page 216 and 217: 204 U. Ugalde II. Germination The p
Page 218 and 219: 206 U. Ugalde Fig. 11.2.A-C Drawing
Page 220 and 221: 208 U. Ugalde duction (Schimmel et
Page 222 and 223: 210 U. Ugalde position, possibly in
Page 224 and 225: 212 U. Ugalde Champe SP, Rao P, Cha
Page 226 and 227: 12 Pheromone Action in the Fungal G
Page 228 and 229: sibly due to displacement of the hy
Page 230 and 231: all these compounds, the B-derivate
Page 232 and 233: shows the same activity in M. muced
Page 234 and 235: C. Oomycota In the non-mycotan phyl
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Page 240 and 241: Elliott CG, Knights BA (1981) Uptak
Page 242 and 243: constitutively transcribed but its
Page 244 and 245: 234 L.M. Corrochano and P. Galland
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254 L.M. Corrochano and P. Galland
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Reproductive Processes
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264 R. Fischer and U. Kües 2. Chla
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266 R. Fischer and U. Kües resourc
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268 R. Fischer and U. Kües ular le
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270 R. Fischer and U. Kües pathway
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272 R. Fischer and U. Kües and con
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274 R. Fischer and U. Kües Timberl
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276 R. Fischer and U. Kües PsiB le
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278 R. Fischer and U. Kües Table.
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280 R. Fischer and U. Kües tein ki
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282 R. Fischer and U. Kües nitroge
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284 R. Fischer and U. Kües tion, t
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286 R. Fischer and U. Kües Busch S
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288 R. Fischer and U. Kües Jeffs L
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290 R. Fischer and U. Kües Pöggel
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292 R. Fischer and U. Kües Yamashi
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294 R. Debuchy and B.G. Turgeon tha
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296 R. Debuchy and B.G. Turgeon loc
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298 R. Debuchy and B.G. Turgeon Tab
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300 R. Debuchy and B.G. Turgeon Fig
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302 R. Debuchy and B.G. Turgeon mai
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304 R. Debuchy and B.G. Turgeon mol
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306 R. Debuchy and B.G. Turgeon gen
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308 R. Debuchy and B.G. Turgeon int
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310 R. Debuchy and B.G. Turgeon Fig
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312 R. Debuchy and B.G. Turgeon pro
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314 R. Debuchy and B.G. Turgeon B.
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316 R. Debuchy and B.G. Turgeon 2.
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318 R. Debuchy and B.G. Turgeon in
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320 R. Debuchy and B.G. Turgeon be
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322 R. Debuchy and B.G. Turgeon Lee
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16 Fruiting-Body Development in Asc
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phae originating from the base of a
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Table. 16.1. (continued) Fruiting B
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(Raju 1992). When male-sterile muta
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1. nsd (never in sexual development
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egular intervals; both effects requ
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the balance between sexual and asex
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ductionofeitherpheromoneisdirectlyc
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(Catlett et al. 2003). Eleven genes
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negative mutation of KREV-1 resulte
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through MAP kinase modules to cell-
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The fact that fruiting-body formati
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Balestrini R, Mainieri D, Soragni E
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point mutation of the beta subunit
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Mitchell TK, Dean RA (1995) The cAM
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YamashiroCT,EbboleDJ,LeeBU,BrownRE,
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358 L.A. Casselton and M.P. Challen
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376 M. Feldbrügge et al. different
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378 M. Feldbrügge et al. Fig. 18.3
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380 M. Feldbrügge et al. et al. 20
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382 M. Feldbrügge et al. for unbia
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384 M. Feldbrügge et al. In U. may
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386 M. Feldbrügge et al. Neurospor
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388 M. Feldbrügge et al. Bernards
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390 M. Feldbrügge et al. O’Donne
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19 The Emergence of Fruiting Bodies
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2003; Kües et al. 2004) are the fi
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(Manachère 1988), C. cinereus (Tsu
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emergence of fruiting bodies. In th
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Rather, development is arrested in
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10 nm thick is highly insoluble and
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it has been suggested that hydropho
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expression in the stipe suggests th
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Cooper DNW, Boulianne RP, Charlton
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Lu BC (1974) Meiosis in Coprinus. R
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Takagi Y, Katayose Y, Shishido K (1
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20 Meiosis in Mycelial Fungi D. Zic
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Fig. 20.1. A-E Diagrammatic represe
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etween dispersed DNA repeats. In N.
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Fig. 20.2. Meiotic recombination. S
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mechanism whereby homologues locate
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An important component of the bouqu
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observed when the mammal LE compone
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A. Chromosome and Sister-Chromatid
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ascus plus ascospore morphogenesis
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syndrome)discoveredasageneinvolvedi
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Kitajima TS, Kawashima SA, Watanabe
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Rossignol J-L, Faugeron G (1995) MI
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Biosystematic Index Absidia glauca
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violaceum 153 Microsporum 149 Mimos
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Subject Index ABC transporter 382 a
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fluffy 270, 276 formin 8, 10, 13, 1
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MAT protein interaction 308, 315 ma
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sporangiophore 234, 242, 246-252, 2
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