Growth, Differentiation and Sexuality

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370 L.A. Casselton and M.P. Challen Kües et al. 2002). Several interesting genes have been characterised (see Chaps. 15 and 19, this volume) but we are a long way from understanding the downstream pathways in any of the fungi that we have described above. Progress is likely to change dramatically now that the genome sequences of these fungi are becoming available (http://www.broad.mit.edu/annotation/fungi/fgi/), and microarray technology will aid identification of major subsets of genes whose transcription alters in response to mating. As yet, few genes have been identified that are direct targets of the pheromone response factor (other than the mating type genes themselves) or the homeodomain protein heterodimer. Fortuitously, a direct target gene of the b heterodimer of U. maydis resides in the a2 mating type locus, lga2 (Romeis et al. 1997) but its function appears to be irrelevant to mating (Bortfeld et al. 2004). The lga2 promoter has a sequence motif that binds the b proteins, and this is remarkably similar to the binding site of the corresponding a1/a2 heterodimer of S. cerevisiae.ThebproteinsofU. maydis are much longer than the yeast proteins and, unlike a1/α2, the b proteins are transcriptional activators. Another b-target gene (frb52), encoding a putative DNA polymerase, was identified in a major screen to detect genes activated or repressed by the b heterodimer (Brachmann et al. 2001). Several interesting gene functions were identified that act downstream of b, one being a MAPK (Kpp6) required for fungal penetration of the host tissues (Brachmann et al. 2003). In C. cinereus, a potential target of the A protein heterodimer was identified by the classical genetic approach of looking for mutants blocked in clamp cell formation. The clp1 gene is induced by activating the A pathway, and has an essential sequence motif in its promoter that resembles the b protein binding sites in lga2 and frb52 (Inada et al. 2001). Expressed off a constitutive promoter, clp1 is sufficient to activate the entire clamp cell programme, showing that although there may be many promoters that bind the HD1/HD2 heterodimer, clp1 may be the only critical target required to activate the pathway in C. cinereus. Makino and Kamada (2004) have identified several mutants that are defective in nuclear migration; these could be potential targets of the predicted pheromone response factor Hmg1. The Pcc1 protein has been implicated in regulating clamp cell fusion (see above) but the pcc1 mutant has an unusual phenotype, which is why it was identified; it constitutively produces unfused clamp cells, as demonstrated by Murata et al. (1998). These authors suggested that Pcc1 might act as a repressor of the clamp cell pathway. This may well be an essential, if indirect function of an activated pheromone response. The A and B regulated pathways alternate during dikaryotic cell growth and are mutually exclusive. The A pathway leads to tip cell extension and nuclear division, whereas the B pathway requires cell–cell recognition which, as in mating S. cerevisiae cells, is likely to require cell cycle arrest before cell fusion. IV. Concluding Remarks The mating type genes are not conserved between basidiomycetes and ascomycetes (see Chap. 15, this volume, and Hiscock and Kües 1999 for a detailed account of both ascomycete and basidiomycete mating types) but this does not mean that the pathways they regulate are different. The mating typelocusissimplyastrategyforpreventingconstitutive sexual development by requiring mates to bring together the full complement of necessary genes (see Casselton 2002); the genes that are sequestered there are the result of evolutionary accident. Pheromone signalling is an essential part of mating in both groups of fungi, and we see a strong conservation in the response induced by pheromone signalling, once we look beyond the apparent differences in life style. There are only two mating types in ascomycetes, and the pheromones and receptors are not mating type genes – they are regulated by the mating type genes (see Coppin et al. 1997, and Chap. 16, this volume). As in U. maydis and C. neoformans, pheromones can act as chemoattractants, and are secreted into the surrounding environment where binding to a compatible receptor may induce the formation of mating structures that aid cell fusion. S. cerevisiae, U. maydis and C. neoformans all change shape in response to pheromone stimulation, increasing in size or forming pegs or filaments that orient towards the source of pheromone and promote cell fusion. In the filamentous ascomycetes such as Neurospora crassa, both mating types may differentiate male and female cells, the female cells being the protoperithecia which, when fertilised, develop into the fruiting bodies in which the sexual cycle is completed. Emerging from the protoperithecia is a filament known as the trichogyne
that is attracted by pheromone stimulation to fuse with the conidia that act as male gametes (Chap. 16, this volume). The conidial nucleus must migrate through the trichogyne cells (Chap. 20, this volume), in much the same way that the donor nucleus migrates through recipient monokaryotic cells in C. cinereus mating (Fig. 17.2). At the base of the trichogyne in N. crassa, nuclear sorting occurs to establish binucleate cells (ascogenous hyphae) that divide in exactly the same way as do dikaryotic cells of homobasidiomycetes and C. neoformans. These cells contain one nucleus from each mate, and tip cell division involves a structure analogous to the clamp connection (crozier; Chap. 20, this volume). It is not surprising that, in N. crassa, pheromone signalling has been shown to be essential after trichogyne fusion (Kim and Borkovich 2004) – it is required to maintain the dikaryophase. Whydosomanyofthesefungihavealong dikaryophase? Diploidy is not a normal feature of thefilamentousfungallifecycle,andnuclearfusion generally occurs only in cells specialised for meiosis. The dikaryophase provides a way of amplifying acompatiblepairofnucleisothatlargenumbersof meiotic products can finally be derived from them. For example, a single fruiting body of C. cinereus may produce 10 7 meiocytes (Pukkila and Casselton 1991), all having a diploid nucleus derived from the same dikaryotic nuclear pair. In filamentous ascomycetes, the dikaryophase is of limited duration but in mushrooms it probably represents the predominant mycelial state in nature, and can exist for hundreds of years, colonising entire woodlands (Smith et al. 1992). Acknowledgements. We are particularly grateful to June Kwon-Chung for describing the C. neoformans life cycle, Tim James for sharing unpublished data, and Joe Heitman for helpful discussion. L.A. Casselton is supported by a Leverhulme Trust Emeritus Fellowship and work at Warwick HRI is supported by the Biotechnology and Biological Sciences Research Council of the UK. References Alic M, Letzring C, Gold MH (1987) Mating system and basidiospore formation in the lignin-degrading basidiomycete Phanerochaete chrysosporium.ApplEnviron Microbiol 53:1464–1469 AndersonCM,WillitsDA,KostedPJ,FordEJ,Martinez- Espinoza AD, Sherwood JE (1999) Molecular analysis of the pheromone and pheromone receptor genes of Ustilago hordei. Gene 240:89–97 Mating Type Genes in Basidiomycetes 371 Asante-Owusu RN, Banham AH, Böhnert HU, Mellor EJC, Casselton LA (1996) Heterodimerization between two classes of homeodomain proteins in the mushroom Coprinus cinereus brings together potential DNA-binding and activation domains. Gene 172:25–31 Badalyan SM, Polak E, Hermann R, Aebi M, Kües U (2004) Role of peg formation in clamp cell fusion of homobasidiomycete fungi. J Basic Microbiol 44:167–177 Bakkeren G, Kronstad JW (1993) Conservation of the bmating-type gene-complex among bipolar and tetrapolar smut fungi. Plant Cell 5:123–136 Bakkeren G, Kronstad JW (1994) Linkage of mating-type loci distinguishes bipolar from tetrapolar mating in basidiomycetous smut fungi. Proc Natl Acad Sci USA 91:7085–7089 Banham AH, Asante Owusu RN, Göttgens B, Thompson SAJ, Kingsnorth CS, Mellor EJC, Casselton LA (1995) An N-terminal dimerization domain permits homeodomain proteins to choose compatible partners and initiate sexual development in the mushroom Coprinus cinereus. Plant Cell 7:773–783 Banuett F (1995) Genetics of Ustilago maydis, a fungal pathogen that induces tumors in maize. Annu Rev Genet 29:179–208 Bölker M, Urban M, Kahmann R (1992) The a mating type locus of U. maydis specifies cell signaling components. Cell 68:441–450 Bortfeld M, Auffarth K, Kahmann R, Basse CW (2004) The Ustilago maydis a2 mating-type locus genes Iga2 and rga2 compromise pathogenicity in the absence of the mitochondrial p32 family protein Mrb1. Plant Cell 16:2233–2248 Brachmann A, Weinzierl G, Kämper J, Kahmann R (2001) Identification of genes in the bW/bE regulatory cascade in Ustilago maydis. Mol Microbiol 42:1047–1063 Brachmann A, Schirawski J, Müller P, Kahmann R (2003) An unusual MAP kinase is required for efficient penetration of the plant surface by Ustilago maydis. EMBO J 22:2199–2210 Brown AJ, Casselton LA (2001) Mating in mushrooms: increasing the chances but prolonging the affair. Trends Genet 17:393–400 Buller AHR (1931) Researches on Fungi, vol 4. Longmans, Green, London Casselton LA (2002) Mate recognition in fungi. Heredity 88:142–147 Casselton LA, Olesnicky NS (1998) Molecular genetics of mating recognition in basidiomycete fungi. Microbiol Mol Biol Rev 62:55–70 Christensen JJ (1931) Studies on the genetics of Ustilago zeae. Phytopathol Z 4:129–188 Coppin E, Debuchy R, Arnaise S, Picard M (1997) Mating types and sexual development in filamentous ascomycetes. Microbiol Mol Biol Rev 61:411–428 Cummings WJ, Celerin M, Crodian J, Brunick LK, Zolan ME (1999) Insertional mutagenesis in Coprinus cinereus: use of a dominant selectable marker to generate tagged, sporulation-defective mutants. Curr Genet 36:371–382 Davidson RC, Moore TDE, Odom AR, Heitman J (2000) Characterization of the MF α pheromone of the human fungal pathogen Cryptococcus neoformans. Mol Microbiol 38:1017–1026 Day PR (1960) The structure of the A mating-type locus in Coprinus lagopus. Genetics 45:641–650
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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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Page 332 and 333: 16 Fruiting-Body Development in Asc
Page 334 and 335: phae originating from the base of a
Page 336 and 337: Table. 16.1. (continued) Fruiting B
Page 338 and 339: (Raju 1992). When male-sterile muta
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Page 342 and 343: egular intervals; both effects requ
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Page 348 and 349: (Catlett et al. 2003). Eleven genes
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Page 352 and 353: through MAP kinase modules to cell-
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Page 356 and 357: Balestrini R, Mainieri D, Soragni E
Page 358 and 359: point mutation of the beta subunit
Page 360 and 361: Mitchell TK, Dean RA (1995) The cAM
Page 362 and 363: YamashiroCT,EbboleDJ,LeeBU,BrownRE,
Page 364 and 365: 358 L.A. Casselton and M.P. Challen
Page 382 and 383: 376 M. Feldbrügge et al. different
Page 384 and 385: 378 M. Feldbrügge et al. Fig. 18.3
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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
Page 400 and 401: 2003; Kües et al. 2004) are the fi
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
Page 410 and 411: it has been suggested that hydropho
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 420 and 421: 20 Meiosis in Mycelial Fungi D. Zic
Page 422 and 423: Fig. 20.1. A-E Diagrammatic represe
Page 424 and 425: 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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