Growth, Differentiation and Sexuality

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310 R. Debuchy and B.G. Turgeon Fig. 15.8. Amino-acid alignment of the HMG domain of deduced MAT1-2-1 proteins of Euascomycetes. P. brassicae (CAA06843), R. secalis (CAD62166), N. crassa (AAA33598), S. macrospora (CAA71624), P. anserina (CAA45520), C. globosum (Broad Institute), G. moniliformis (AAC71056), F. oxysporum (BAA28611), G. zeae (AAO42810), C. eucalypti (AAF00498), M. grisea (BAC65090), P. tenuipes (BAC66503), in vitro, whose core is 5 ′ -CAAAG-3 ′ (Philley and Staben 1994). This binding sequence is identical to the binding site of the SOX-TCF_HMG box, suggesting that SOX-TCF and MATA-HMG boxes may function in a very similar way, although these pro- C. parasitica (AAK83343), E. nidulans (AAQ07985), C. heterostrophus (CAA48464), C. luttrellii (AAD33439), C. homomorphus (AAD33441), C. kusanoi (AAD33442), C. cymbopogonis (AAD33447), A. alternata (BAA75908), A. rabiei (Barve et al. 2003), P. nodorum (AAO31742), M. graminicola (AAL30836). Fraction of sequences that must agree for shading:0.8 teins are placed in separate families. MAT1-1-3 proteins display several features that distinguish them from MAT1-2-1, and suggest that they could have very different DNA-binding properties. The structures of the HMG boxes in MAT1-1-3 and MAT1-2-1
appear very dissimilar (see consensus in Figs. 15.7 and 15.8). Phylogenetic analysis of the HMG domains from the MATA family shows that MAT1-2-1 proteins form a distinct subfamily that does not contain any other HMG proteins, except MAT1-1-3 from P. brassicae and R. secalis (data not shown). Moreover, SMR2 from P. anserina was shown to interact with FMR1 (Arnaise et al. 1995), and similar phenotypes conferred by FMR1 and SMR2 mutations support the idea that these two proteins cooperate in the regulation of their target genes (Zickler et al. 1995; Arnaise et al. 1997, 2001). These data support the idea that MAT1-1-3 proteins bind to DNA in a very different way from that of MAT1- 2-1/SOX-TCF proteins. Further investigations on DNA-binding modalities of fungal HMG transcription factors would be required to substantiate classification of these proteins, by function. IV. Evolution of Mating Types Two fascinating questions attend any discussion of mating-type gene evolution. The first is how dimorphic sex chromosomes or dissimilar regions (idiomorphs) of a chromosome evolved, and the second is how different reproductive lifestyles, i.e., self- or non-self-compatibility, evolve. The evolutionary origins of the dissimilar MAT idiomorphs of self-incompatible Ascomycetes are unknown at this point, but data are accumulating on the matter. One hypothesis is that the small pockets of identity in the otherwise unlike MAT idiomorphs reflect common ancestry (Coppin et al. 1997). Conceivably, these pockets are remnants of a series of mutagenic events in a single ancestral gene(s). The mutations, coupled with recombination suppression, might have led to the highly divergent extant MAT genes that now encode different products (Turgeon et al. 1993). A similar scenario has been proposed for the evolution of the Y chromosome. Indeed, acquisition of sex-determining genes, recombination isolation, and addition of genes not involved in mating are shared between fungal mating-type loci and mammalian X and Y chromosomes; it has been suggested that these elements may be early steps linking the evolution of sex chromosomes in diverse organisms (Lahn and Page 1999; Fraser et al. 2004). This section focuses on how Ascomycetes change from one reproductive lifestyle to another (self-incompatibility to self-compatibility, and vice versa). Mating Types in Euascomycetes 311 A. Phylogenetic Analyses of Mating Type 1. Loculoascomycetes a) Cochliobolus Because MAT genes control the reproductive process, comparison of their sequences reflects life history and should reveal mechanisms underlying changes in reproductive mode. A combination of molecular and phylogenetic data have been used to determine the direction of evolution of reproductive lifestyle in Cochliobolus (Yun et al. 1999) and in Stemphylium (Inderbitzin et al. 2005). In both cases, examination of MAT structure in many species from diverse geographical locations, plus phylogenetic treatments, support the hypothesis that the direction is likely from self-incompatible to self-compatible. To address the issue of which mode of fungal sexual reproduction is ancestral in Cochliobolus, Yun et al. (1999) compared extant MAT sequences from self-incompatible and self-compatible species. Structural organization of MAT loci of self-incompatible C. heterostrophus, C. carbonum, C. victoriae, C. ellisii, C. sativus and C. intermedius species is highly conserved; each strain carries asingleMAT gene, either MAT1-1-1 or MAT1- 2-1 (Fig. 15.2). By contrast, all self-compatible Cochliobolus species carry both MAT genes in one genome, but as described in Sect. III. and Fig. 15.2, the structural organization of each locus is unique. In two cases (C. luttrellii and C. homomorphus), the genes are fused into a single ORF; the gene order in C. luttrellii is reversed in C. homomorphus. In the remaining two cases, the genes are not fused. In C. kusanoi, the organization is 5 ′ MAT1-2-13 ′ –3 ′ MAT1-1-15 ′ , and part of the sequence between the genes is similar to a portion of the β-glucosidasegenenormallyfound3 ′ of both MAT genes in self-incompatible C. heterostrophus (Fig. 15.3). To the 5 ′ of MAT1-2-1 is a perfect inverted repeat of a 561-bp region containing 123 bp of the 5 ′ end of the MAT1-1-1 ORF fused to the 5 ′ end of MAT1-2-1, and 145 bp of a different fragment of the β-glucosidase gene, separated from each other by 293 bp. C. cymbopogonis carries both homologs of the self-incompatible idiomorphs, but these are not closely linked. It is not known if they reside on the same or different chromosomes. Thus, the MAT genes have close physical association in three Cochliobolus self-compatible species, but not in the fourth. The fused MAT genes in self-compatible species
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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 268 and 269: 258 L.M. Corrochano and P. Galland
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Page 272 and 273: 264 R. Fischer and U. Kües 2. Chla
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Page 278 and 279: 270 R. Fischer and U. Kües pathway
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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
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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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Page 346 and 347: ductionofeitherpheromoneisdirectlyc
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Page 360 and 361: Mitchell TK, Dean RA (1995) The cAM
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Page 364 and 365: 358 L.A. Casselton and M.P. Challen
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362 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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