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20 Meiosis in Mycelial Fungi D. Zickler 1 CONTENTS I. Introduction ......................... 415 II. Entering Meiosis: Mycelial Fungi Devote Significant Resources to Make Sure that the Two Nuclei that Will Fuse Before Entering Meiosis Have Identical Genomes .......... 416 A. Karyogamy and Premeiotic Replication . . 417 B. Premeiotic “Checking and Cleaning” Mechanisms....................... 418 C. A “Checking” Mechanism thatOperatesAfterKaryogamy ........ 419 III. Meiotic Recombination ................. 419 A. Initiation ......................... 419 B. From Initiation toRecombinationProducts ........... 420 C. Meiotic Exchanges Are Highly Regulated . 421 IV. Homologue Recognition and Pairing ...... 422 A. Mycelial Fungi Offer Unique Opportunities for the Analysis ofHomologousPairing .............. 423 B. HowDoesPairingOccur?............. 423 C. The Bouquet Stage: a Specific ConfigurationofMeiosis ............. 424 D. Chromosome Interlocking: a Universal Complication of Pairing . . . . . 425 V. The Synaptonemal Complex and Synapsis . . 425 A. Synaptonemal Complex Formation . . . . . 426 B. Ascomycetes Exhibit Several Peculiarities in the Synaptonemal Complex Formation andMorphology ................... 426 C. Synaptonemal Complex andtheRecombinationProcess........ 427 D. Recombination Nodules, the Substructures of the Synaptonemal Complex that Correlate with Crossover andNoncrossoverExchanges.......... 427 VI. Meiotic Chromosome Segregation or how to Resolve Sister-Chromatid Cohesion in Two Steps .................. 428 A. Chromosome and Sister-Chromatid Segregation Are Mediated bytheCohesinComplex.............. 429 B. Other Proteins Important for Sister-Chromatid Cohesion andSegregation.................... 430 VII. From Meiosis to Sporulation ............. 430 VIII. Concluding Remarks ................... 431 References ........................... 432 1 Université Paris-Sud, Institut de Génétique et Microbiologie, Bâtiment 400, 91405 Orsay cedex, France I. Introduction Meiosis is a highly conserved process, occupying a central role in most eukaryote life cycles. The meiotic process, in contrast to the mitotic process, reduces the diploid cellular genome complement by half, as required for sexual reproduction and fecundation. The reduction in chromosome number is achieved by one round of DNA replication, followed by two rounds of divisions with no intervening replication. The absolute need for each gamete to inherit a copy of the genome is achieved bythefactthatpaternalandmaternalhomologous chromosomes segregate to opposite poles during the first meiotic division, whereas their sister chromatids segregate at the second division. Mycelial fungi provide several positive attributes for studying meiosis. First, they have a brief life cycle during which several hundred meiocytes and the resulting gametes (asco- and basidiospores) can be analyzed. Second, the four products of a single meiosis are held together in a single cell (ascus or basidium). This allows the determination of the genetic constitution of each of the DNA strands involved in meiosis. Moreover, in species with linearly arranged ascospores, the position of each ascospore reflects the preceding nuclear divisions. When in some of these species a mitotic division occurs after meiosis, each of the resulting eight haploid nuclei represents the genetic character of one of the eight DNA strands of the chromosomes produced by meiosis (e.g., Perkins 1974, 1997). The popularity of Neurospora crassa, Sordaria fimicola, Saccharomyces cerevisiae, Schizosaccharomyces pombe, Aspergillus nidulans, Ascobolus immersus, Podospora anserina, Sordaria macrospora, Schizophyllum commune and Coprinus cinereus (now called Coprinopsis cinerea) for a wide variety of genetic studies on meiotic recombination reflects these exceptional advantages (review in Esser and Kuenen 1967; Whitehouse 1982; Lamb 1996). The Mycota I Growth, Differentation and Sexuality Kües/Fischer (Eds.) © Springer-Verlag Berlin Heidelberg 2006
416 D. Zickler Third, a large pool of meiotic mutants have been isolated in A. nidulans, N. crassa, P. anserina, S. macrospora and C. cinereus, and molecular tools are now available to characterize them (see below). Fourth, publication of the genome sequences of five ascomycetes (A. nidulans, Fusarium graminearum, N. crassa, Magnaporthe grisea, P. anserina) and four basiomycetes (C. cinerea, Cryptococcus neoformans, Phanerochaete chrysosporium, Ustilago maydis) provides a powerful basis for genome comparison with other organisms, and especially with the budding and fission yeasts, in which a large spectrum of meiotic genes are already characterized (e.g., Borkovich et al. 2004). Homologies among genes involved in the meiotic and sporulation processes establish rapid ways to identify genes in the fungus of interest (even when the sexual cycle is unknown, e.g., Pöggeler 2002). They offer also a starting point for the identification of fungal-specific genes and/or evolutionary gene requirements. Fifth, the small chromosomal size of most fungi was long considered a handicap for the general requirement for good chromosome morphology to study meiosis. However, since chromosome organization can be investigated using tools such as three-dimensional reconstruction from serial sections, fluorescence in situ hybridization (FISH) and green fluorescent protein (GFP) tagging, fungi became model systems for investigating both meiotic pairing and segregation (cf. references below, in corresponding paragraphs). Finally, everyone who has observed fungal chromosomes knows that Barbara McClintock was right when she said in her 1961 conference exposé “You may think they (chromosomes of N. crassa) are small when I show you pictures of them. But when you look at them, they get bigger and bigger and bigger” (cited in Perkins 1992). The major purpose of the present review is to explore similarities and differences in the better known organisms, in order to identify key conserved features of the meiotic process. Recent studies based on genetic, molecular and cytological approaches, combined with a rapidly growing arsenal of mutations, have yielded a great deal of new information about how pairing and synapsis relate to other processes such as recombination and segregation in budding and fission yeasts as well as in C. cinereus and S. macrospora. In fact, the major task of the prolonged meiotic prophase is orderly recognition plus juxtaposition of the homologous chromosomes and establishment of stable connections via crossing-over. Throughout this period, events at the DNA level and events at the chromosome axis level are spatially and temporally coordinated. Most of the review will therefore be dedicated to these critical steps. Differences found when mutants of different organisms are analyzed shed light on our understanding of meiosis, and although only few mycelial fungi are used as model organisms, it is clear that they underline already both the diversity and the potentiality of these organisms. A clear example is given by the variety of gene silencing mechanisms developed by some fungi before and during meiosis. Differences underline also thefactthatinthisvastgroupoforganismsthere may be interesting “obscure” species awaiting the opportunity to give important clues. Interestingly also, mycelial fungi have developed a great variety of methods for separating or bringing together the pairs of nuclei that are issued from both meiotic divisions and, when present, the postmeiotic mitosis. Such movements imply a strict regulation of how spindles are located, which is also discussed here. Finally, an understanding of the meiotic process is not only pivotal to furthering research on fertility but also has important implications for fungal and human disease curing. II. Entering Meiosis: Mycelial Fungi Devote Significant Resources to Make Sure that the Two Nuclei that Will Fuse Before Entering Meiosis Have Identical Genomes Mycelial fungi are self-fertile (also called selfcompatible or homothallic) or self-sterile, cross-fertile (self-incompatible or heterothallic). In primary homothallic species, a homokaryotic mycelium established from a single haploid nucleus has the potentiality to progress through both the dikaryotic stage and meiosis to complete the sexual cycle. In secondary homothallic species (also called “pseudoheterothallic”), a fertile dikaryotic mycelium is established from a single spore carrying two haploid nuclei of different mating types. In heterothallic species, mating between homokaryotic mycelia of different mating types is required to complete the sexual cycle (Fig. 20.1A; see also Debuchy and Turgeon, and Casselton and Challen, Chaps. 15 and 17, respectively, this volume). In all cases, there is a more (cf. mycelial basidiomycetes) or less (cf. mycelial
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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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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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Page 370 and 371: 364 L.A. Casselton and M.P. Challen
Page 382 and 383: 376 M. Feldbrügge et al. different
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Page 386 and 387: 380 M. Feldbrügge et al. et al. 20
Page 388 and 389: 382 M. Feldbrügge et al. for unbia
Page 390 and 391: 384 M. Feldbrügge et al. In U. may
Page 392 and 393: 386 M. Feldbrügge et al. Neurospor
Page 394 and 395: 388 M. Feldbrügge et al. Bernards
Page 396 and 397: 390 M. Feldbrügge et al. O’Donne
Page 398 and 399: 19 The Emergence of Fruiting Bodies
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Page 402 and 403: (Manachère 1988), C. cinereus (Tsu
Page 404 and 405: emergence of fruiting bodies. In th
Page 406 and 407: Rather, development is arrested in
Page 408 and 409: 10 nm thick is highly insoluble and
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Page 412 and 413: expression in the stipe suggests th
Page 414 and 415: Cooper DNW, Boulianne RP, Charlton
Page 416 and 417: Lu BC (1974) Meiosis in Coprinus. R
Page 418 and 419: Takagi Y, Katayose Y, Shishido K (1
Page 422 and 423: Fig. 20.1. A-E Diagrammatic represe
Page 424 and 425: etween dispersed DNA repeats. In N.
Page 426 and 427: Fig. 20.2. Meiotic recombination. S
Page 428 and 429: mechanism whereby homologues locate
Page 430 and 431: An important component of the bouqu
Page 432 and 433: observed when the mammal LE compone
Page 434 and 435: A. Chromosome and Sister-Chromatid
Page 436 and 437: ascus plus ascospore morphogenesis
Page 438 and 439: syndrome)discoveredasageneinvolvedi
Page 440 and 441: Kitajima TS, Kawashima SA, Watanabe
Page 442 and 443: Rossignol J-L, Faugeron G (1995) MI
Page 444 and 445: Biosystematic Index Absidia glauca
Page 446 and 447: violaceum 153 Microsporum 149 Mimos
Page 448 and 449: Subject Index ABC transporter 382 a
Page 450 and 451: fluffy 270, 276 formin 8, 10, 13, 1
Page 452 and 453: MAT protein interaction 308, 315 ma
Page 454: sporangiophore 234, 242, 246-252, 2
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