Growth, Differentiation and Sexuality

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296 R. Debuchy and B.G. Turgeon locus has been found. This locus will be described below in self-incompatible, self-compatible, and asexual species, respectively. 1. Loculoascomycetes a) Self-Incompatible Loculoascomycetes The simplest idiomorphs are those of selfincompatible Loculoascomycetes (Fig. 15.2). For all taxa in this group, MAT1-1 and MAT1-2 idiomorphs consist of one ORF corresponding to genes encoding the α1 and HMG proteins, respectively (Table 15.1). Following standard nomenclature, described above, the genes have been designated MAT1-1-1 and MAT1-2-1. The size of the idiomorphs varies with species, but both encoded MAT genes have the same orientation with respect to their chromosome. For C. heterostrophus, the first described for this class, each ORF occupies almost all of the idiomorph sequence (Turgeon et al. 1993). Expression analysis revealed that both MAT1-1-1 and MAT1-2-1 transcripts start and stop in common sequences flanking the idiomorphs, resulting in transcripts that are almost twice the size of the corresponding ORF (Leubner-Metzger et al. 1997). Multiple transcripts, resulting from the use of at least two transcription start sites and alternative splicing of 5 ′ UTR introns, are produced. The 5 ′ UTR of the MAT messages contain μORFs that could control translation of the downstream coding sequence. However, the transcription initiation sites and the 5 ′ UTR region of the MAT genes have been deleted in both parents without penalty to fertility, indicating that these regions do not contain information essential for mating (Wirsel et al. 1998). By contrast, the 3 ′ UTR of the idiomorphic genes contains structural information that is necessary for completion of the sexual cycle. The molecular function of this sequence present in the 3 ′ common flanking sequence of MAT1-1 and MAT1-2 idiomorphs is as yet unknown. This one gene/one idiomorph structure has also been found in C. ellisii, C. carbonum, C. victoriae,andC. intermedius (Yun et al. 1999). The MAT idiomorphs of Didymella rabiei (sexual state of Ascochyta rabiei; Barve et al. 2003), Mycosphaerella graminicola (sexual form of Septoria tritici; Waalwijk et al. 2002) and Phaeosphaeria nodorum (sexual state of Stagonospora nodorum; Bennett et al. 2003) also contain a single ORF that starts and terminates in the idiomorph, but the sizes of the idiomorphic sequences are larger than those of C. heterostrophus (Table 15.1). Transcription initiation sites have not been established for these fungi. It is possible that, in contrast to C. heterostrophus, the promoter is localized in the MAT-specific sequences, raising the possibility that, unlike C. heterostrophus, each MAT gene responds to specific transcription regulatory factors. b) Self-Compatible Loculoascomycetes Self-compatible Loculoascomycetes display various arrangements of MAT1-1-1 and MAT1-2-1 genes in their haploid genome (Fig. 15.2 and Table 15.2). C. cymbopogonis carries complete MAT1-1-1 and MAT1-2-1 counterparts, in contrast to C. luttrellii and C. homomorphus that contain MAT gene fusions (Yun et al. 1999). These fusions consist of either MAT1-1-1::MAT1-2-1 (C. luttrellii) or MAT1-2-1::MAT1-1-1 (C. homomorphus) recombinant genes. Possible evolution of these self-compatible species from self-incompatible ancestors is detailed in Sect. IV. A more complex structure has been found in C. kusanoi (Yun et al. 1999). This fungus contains a complete MAT1-1-1 homologlinkedtoaMAT1-1-1::MAT1-2-1 chimeric gene that is devoid of the α1 encoding region but contains the HMG encoding sequence. The MAT locus from a self-compatible species outside of the genus Cochliobolus, Mycosphaerella zeae-maydis (Didymella zeae-maydis), has been characterized (Yun 1998). In this case, M. zeae-maydis MAT has complete versions of the C. heterostrophus MAT counterparts tandemly arranged, head to tail, separated by about one kilobase of non-coding sequence. A total of 106 diverse isolates of Stemphylium have been examined to determine the structure of the mating-type loci (Inderbitzin et al. 2005). Isolates were found to contain both separate and linked MAT genes. Where separate, MAT loci contain either a MAT1-1-1 or MAT1-2-1 gene in the same 5 ′ ->3 ′ orientation as in Cochliobolus, between upstream ORF1 and downstream BGL1 genes (Fig. 15.3). Self-compatible isolates were isolated from field ascomata or conidia and demonstrated to undergo self-mating in the laboratory. These isolates had two types of MAT locus configuration. In the first type, the two MAT genes are linked. The MAT1-1-1 gene and both flanks are inverted and connected to MAT1-2-1 oriented in the original MAT gene direction (as in Cochliobolus), approx. 200 bp upstream of the MAT1-2-1
Fig. 15.2. General structure of the MAT locus in selfincompatible and self-compatible Euascomycetes. Arrows indicate position and orientation of the genes. Black Mating Types in Euascomycetes 297 arrows indicate genes found in all studied species in each class. Red arrows indicate genes that are absent in some representative in each class
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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
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Table. 8.1. (continued) Fungal Hete
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is a complex genetic trait controll
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indicate how speciation may be init
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ility controlled by multiple allele
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dependonthematingtypegenes,wasobser
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6. Relation with Histo-Incompatibil
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Semi-Incompatibilität. Z Indukt Ab
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Micali CO, Smith ML (2003) On the i
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Vilgalys RJ, Miller OK (1987) Matin
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168 B.C.K. Lu (Esser et al. 1980; K
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170 B.C.K. Lu enterthePCDpathway,wi
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172 B.C.K. Lu et al. 2004). It is l
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174 B.C.K. Lu (MMP), through ruptur
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176 B.C.K. Lu Fig. 9.1. Effects of
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178 B.C.K. Lu drial fission during
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180 B.C.K. Lu Although the cytologi
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182 B.C.K. Lu be arrested at diffus
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184 B.C.K. Lu Harris MH, Thompson C
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186 B.C.K. Lu oxygen species, in Kl
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10 Senescence and Longevity H.D. Os
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the amplification of plDNA is not a
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GRISEA is an orthologue of the yeas
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Fig. 10.3. Copper delivery to the c
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have been identified, for example,
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Kück U, Stahl U, Esser K (1981) Pl
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Signals in Growth and Development
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204 U. Ugalde II. Germination The p
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206 U. Ugalde Fig. 11.2.A-C Drawing
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208 U. Ugalde duction (Schimmel et
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210 U. Ugalde position, possibly in
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212 U. Ugalde Champe SP, Rao P, Cha
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12 Pheromone Action in the Fungal G
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sibly due to displacement of the hy
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all these compounds, the B-derivate
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shows the same activity in M. muced
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C. Oomycota In the non-mycotan phyl
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following sequence of events has be
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the standard Mendelian segregation
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Elliott CG, Knights BA (1981) Uptak
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constitutively transcribed but its
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234 L.M. Corrochano and P. Galland
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Page 254 and 255: 244 L.M. Corrochano and P. Galland
Page 270 and 271: Reproductive Processes
Page 272 and 273: 264 R. Fischer and U. Kües 2. Chla
Page 274 and 275: 266 R. Fischer and U. Kües resourc
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Page 278 and 279: 270 R. Fischer and U. Kües pathway
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Page 286 and 287: 278 R. Fischer and U. Kües Table.
Page 288 and 289: 280 R. Fischer and U. Kües tein ki
Page 290 and 291: 282 R. Fischer and U. Kües nitroge
Page 292 and 293: 284 R. Fischer and U. Kües tion, t
Page 294 and 295: 286 R. Fischer and U. Kües Busch S
Page 296 and 297: 288 R. Fischer and U. Kües Jeffs L
Page 298 and 299: 290 R. Fischer and U. Kües Pöggel
Page 300 and 301: 292 R. Fischer and U. Kües Yamashi
Page 302 and 303: 294 R. Debuchy and B.G. Turgeon tha
Page 306 and 307: 298 R. Debuchy and B.G. Turgeon Tab
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Page 310 and 311: 302 R. Debuchy and B.G. Turgeon mai
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Page 318 and 319: 310 R. Debuchy and B.G. Turgeon Fig
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Page 324 and 325: 316 R. Debuchy and B.G. Turgeon 2.
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Page 330 and 331: 322 R. Debuchy and B.G. Turgeon Lee
Page 332 and 333: 16 Fruiting-Body Development in Asc
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Page 336 and 337: Table. 16.1. (continued) Fruiting B
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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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