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252 L.M. Corrochano and P. Galland in the graviperception of plants, plays a substantial role also in the gravitropism of Gigaspora rosea, because lanthanum (a Ca 2+ blocker) and EGTA (a Ca 2+ chelator) induce hyphal branching, and atthesametimealsoinhibitionofgravitropism (Berbara et al. 2002). The gravisusceptor of these mycorrhizal fungi are presently unknown. In view of the fact that these oleaginous fungi contain numerous lipid droplets, it appears reasonable to assume that the buoyancy of these droplets mediates graviperception. We found that vertically growing hyphae of Gigaspora margarita possess an apical complex of lipid globules (CLG) that is similar to the one found in Phycomyces (unpublished data). Even in the CLG of Gigaspora, the lipid globules are in continual motion. Lateral hyphae growing horizontally do not contain apical lipid globules and are agravitropic. Thus, lipid globules occur only in segments that grow vertically upward. The close correlation between gravitropic orientation and the occurrence of lipid globules clearly indicates their role as gravisusceptors (Döring, unpublished data). IV. Conclusions Light and gravity are environmental stimuli responsible for the final appearance of fungi. For many decades, research on fungal photomorphogenesis and gravitropic responses has concentrated on the description and characterization of the responses to light and gravity. Genetics and molecular biology have been employed only with limited examples, but nevertheless with great success. The isolation of blind mutants has allowed the identification of relevant proteins, like the photoreceptor WC-1 from Neurospora, confirming the importance of a detailed genetic analysis in the dissection of a photoresponse. Additionally, the Neurospora genome sequence has allowed the identification of several putative photoreceptors whose functions are still unknown. The photoreceptor WC-1 binds DNA promoters of light-inducible genes, establishing a very simple transduction chain. Fungal homologs of the WC-1 photoreceptor have been identified in other ascomycetes and basidiomycetes, but their function may not be exactly the same as that of their Neurospora counterparts, based on the absence of some functional domains and differences in their pattern of expression. Neurospora photobiology is now considered a model for other fungi, but only future research will either confirm this, or will establish differences in fungal photobiology. Undoubtedly, the continuing sequencing of fungal genomes will increase the number of putative elements in light transduction pathways, and these willserveasstartingpointsforfutureresearchin fungal photobiology. The combination of physiology with genetics and molecular biology should be employed for a fuller understanding of fungal photomorphogenesis, from the primary photoreception to the final developmental response. The molecular basis of fungal gravitropism, however, is still rudimentary. Several examples of gravisusceptors and mechanisms for gravity sensing have been suggested and presented here. We hope that the combination of genetics and molecular biology, with detailed cytological characterization of model systems, will help to unravel the complexities of fungal gravitropism. Light and gravity interact in complex ways to direct sporangiophore growth in Phycomyces, possibly at the level of the photoreceptor system itself. It is thus possible that light and gravity will share some of the elements of the transduction chain in fungi. Only future research will confirm the possible relationship between light and gravity as major signals for fungal development, growth, and appearance. References Adams TH, Wieser JK, Yu JH (1998) Asexual sporulation in Aspergillus nidulans. Microbiol Mol Biol Rev 62:35–54 Aguirre J, Ríos-Momberg M, Hewitt D, Hansberg W (2005) Reactive oxygen species and development in microbial eukaryotes. Trends Miocrobiol 13:111–118 Alvarez MI, Benito EP, Campuzano V, Eslava AP (1992) Genetic loci of Phycomyces blakesleeanus. In:O’Brien SJ (ed) Genetic maps. Locus maps of complex genomes, 6th edn. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY, pp 3120–3126 Ambra R, Grimaldi B, Zamboni S, Filetici P, Macino G, Ballario P (2004) Photomorphogenesis in the hypogeous fungus Tuber borchii: isolation and characterization of Tbwc-1, the homologue of the blue-light photoreceptor of Neurospora crassa. Fungal Genet Biol 41:688–697 Arpaia G, Loros JJ, Dunlap JC, Morelli G, Macino G (1993) The interplay of light and the circadian clock. Independent dual regulation of clock-controlled gene ccg- 2(eas). Plant Physiol 102:1299–1305 Arpaia G, Loros JJ, Dunlap JC, Morelli G, Macino G (1995) Lightinduction ofthe clock-controlledgeneccg-1 is not transduced through the circadian clock in Neurospora crassa. Mol Gen Genet 247:157–163 Arpaia G, Cerri F, Baima S, Macino G (1999) Involvement of protein kinase C in the response of Neurospora crassa to blue light. Mol Gen Genet 262:314–322
Baima S, Macino G, Morelli G (1991) Photoregulation of the albino-3 gene in Neurospora crassa. JPhotochem Photobiol B11:107–115 Ballario P, Macino G (1997) White collar proteins: PASsing the light signal in Neurospora crassa. Trends Microbiol 5:458–462 Ballario P, Vittorioso P, Magrelli A, Talora C, Cabibbo A, Macino G (1996) White collar-1, a central regulator of blue light responses in Neurospora, isazincfinger protein. EMBO J 15:1650–1657 Ballario P, Talora C, Galli D, Linden H, Macino G (1998) Roles in dimerization and blue light photoresponse of the PAS and LOV domains of Neurospora crassa white collar proteins. Mol Microbiol 29:719–729 Bejarano ER, Avalos J, Lipson ED, Cerdá-Olmedo E (1991) Photoinduced accumulation of carotene in Phycomyces. Planta 183:1–9 Bell-Pedersen D (2000) Understanding circadian rhythmicity in Neurospora crassa: from behavior to genes and back again. Fungal Genet Biol 29:1–18 Bell-Pedersen D, Dunlap JC, Loros JJ (1996) Distinct cisacting elements mediate clock, light, and developmental regulation of the Neurospora crassa eas (ccg-2)gene. Mol Cell Biol 16:513–521 Berbara R, Fonseca H, Daft M (2002) The role of calcium in the germination and germinative hyphal growth of Gigaspora rosea Nicolson and Schenk. http:// plantbio.berkeley.edu/∼bruns/abstracts/berbara Bergman K, Eslava AP, Cerdá-Olmedo E (1973) Mutants of Phycomyces with abnormal phototropism. Mol Gen Genet 123:1–16 Bergo V, Spudich EN, Spudich JL, Rothschild KJ (2002) A Fourier transform infrared study of Neurospora rhodopsin: similarities with archaeal rhodopsins. Photochem Photobiol 76:341–349 Berrocal-Tito G, Sametz-Baron L, Eichenberg K, Horwitz BA, Herrera-Estrella A (1999) Rapid blue light regulation of a Trichoderma harzianum photolyase gene. J Biol Chem 274:14288–14294 Berrocal-Tito G, Rosales-Saavedra T, Herrera-Estrella A, Horwitz BA (2000) Characterization of blue-light and developmental regulation of the photolyase gene phr1 in Trichoderma harzianum. Photochem Photobiol 71:662–668 Bieszke JA, Braun EL, Bean LE, Kang S, Natvig DO, Borkovich KA (1999a) The nop-1 gene of Neurospora crassa encodes a seven transmembrane helix retinalbinding protein homologous to archaeal rhodopsins. Proc Natl Acad Sci USA 96:8034–8039 Bieszke JA, Spudich EN, Scott KL, Borkovich KA, Spudich JL (1999b) A eukaryotic protein, NOP-1, binds retinal to form an archaeal rhodopsin-like photochemically reactive pigment. Biochemistry 38:14138–14145 Blumenstein A, Vienken K, Tasler R, Frankenberg- Dinkel N, Fischer R (2005) No sex in red light – a role for phytochromes in Aspergillus nidulans. Fungal Genet Newslett Suppl 52:42 Borkovich KA, Alex LA, Yarden O, Freitag M, Turner GE, Read ND, Seiler S, Bell-Pedersen D, Paietta J, Plesofsky N et al. (2004) Lessons from the genome sequence of Neurospora crassa: tracingthepathfromgenomic blueprint to multicellular organism. Microbiol Mol Biol Rev 68:1–108 Photomorphogenesis and Gravitropism 253 Borriss H (1934) Beiträge zur Wachstums- und Entwicklungsphysiologie der Fruchtkörper von Coprinus lagopus. Planta 22:28–69 Brown LS, Dioumaev AK, Lanyi JK, Spudich EN, Spudich JL (2001) Photochemical reaction cycle and proton transfers in Neurospora rhodopsin. J Biol Chem 276:32495– 32505 Buller AHR (1909) Researches on Fungi, vol 1. Longman Green, London Buller AHR (1922) Researches on Fungi, vol 2. Longman Green, London Busch S, Eckert SE, Krappmann S, Braus GH (2003) The COP9 signalosome is an essential regulator of development in the filamentous fungus Aspergillus nidulans. Mol Microbiol 49:717–730 Calvo AM, Hinze LL, Gardner HW, Keller NP (1999) Sporogenic effect of polyunsaturated fatty acids on development of Aspergillus spp. Appl Environ Microbiol 65:3668–3673 Calvo AM, Gardner HW, Keller NP (2001) Genetic connection between fatty acid metabolism and sporulation in Aspergillus nidulans. J Biol Chem 276:25766–25774 Campuzano V, Galland P, Alvarez MI, Eslava AP (1996) Blue-light receptor requirement for gravitropism, autochemotropism and ethylene response in Phycomyces. Photochem Photobiol 63:686–694 Carattoli A, Kato E, Rodriguez-Franco M, Stuart WD, Macino G (1995) A chimeric light-regulated amino acid transport system allows the isolation of blue light regulator (blr) mutantsofNeurospora crassa. ProcNatl Acad Sci USA 92:6612–6616 Carlile MJ (1970) The photoresponses of fungi. In: Halldal P (ed) Photobiology of microorganisms. Wiley, London, pp 309–344 Casadevall A, Perfect JR (1998) Cryptococcus neoformans. ASM Press. Washington, DC Casas-Flores S, Ríos-Momberg M, Bibbins M, Ponce- Noyola P, Herrera-Estrella A (2004) BLR-1 and BLR-2, key regulatory elements of photoconidiation and mycelial growth in Trichoderma atroviride. Microbiology 150:3561–3569 Cashmore AR (2003) Cryptochromes: enabling plants and animals to determine circadian time. Cell 114:537–543 Cerda-Olmedo E (2001) Phycomyces andthebiologyoflight and color. FEMS Microbiol Rev 25:503–512 Cerdá-Olmedo E, Corrochano LM (2001) Genetics of Phycomyces and its responses to light. In: Häder DP, Lebert M (eds) Photomovement. Comprehensive Series in Photosciences, vol 1. Elsevier, Amsterdam, pp 589–620 Cerdá-Olmedo E, Lipson ED (eds) (1987) Phycomyces.Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY Chen C, Dickman MB (2002) Colletotrichum trifolii TB3 kinase, a COT1 homolog, is light inducible and becomes localized in the nucleus during hyphal elongation. Eukaryot Cell 1:626–633 Cheng P, Yang Y, Heintzen C, Liu Y (2001a) Coiled-coil domain-mediated FRQ-FRQ interaction is essential for its circadian clock function in Neurospora. EMBOJ 20:101–108 Cheng P, Yang Y, Liu Y (2001b) Interlocked feedback loops contribute to the robustness of the Neurospora circadian clock. Proc Natl Acad Sci USA 98:7408–7413
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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,
Page 212 and 213: Kück U, Stahl U, Esser K (1981) Pl
Page 214 and 215: Signals in Growth and Development
Page 216 and 217: 204 U. Ugalde II. Germination The p
Page 218 and 219: 206 U. Ugalde Fig. 11.2.A-C Drawing
Page 220 and 221: 208 U. Ugalde duction (Schimmel et
Page 222 and 223: 210 U. Ugalde position, possibly in
Page 224 and 225: 212 U. Ugalde Champe SP, Rao P, Cha
Page 226 and 227: 12 Pheromone Action in the Fungal G
Page 228 and 229: sibly due to displacement of the hy
Page 230 and 231: all these compounds, the B-derivate
Page 232 and 233: shows the same activity in M. muced
Page 234 and 235: C. Oomycota In the non-mycotan phyl
Page 236 and 237: following sequence of events has be
Page 238 and 239: the standard Mendelian segregation
Page 240 and 241: Elliott CG, Knights BA (1981) Uptak
Page 242 and 243: constitutively transcribed but its
Page 244 and 245: 234 L.M. Corrochano and P. Galland
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
Page 276 and 277: 268 R. Fischer and U. Kües ular le
Page 278 and 279: 270 R. Fischer and U. Kües pathway
Page 280 and 281: 272 R. Fischer and U. Kües and con
Page 282 and 283: 274 R. Fischer and U. Kües Timberl
Page 284 and 285: 276 R. Fischer and U. Kües PsiB le
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 304 and 305: 296 R. Debuchy and B.G. Turgeon loc
Page 306 and 307: 298 R. Debuchy and B.G. Turgeon Tab
Page 308 and 309: 300 R. Debuchy and B.G. Turgeon Fig
Page 310 and 311: 302 R. Debuchy and B.G. Turgeon mai
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304 R. Debuchy and B.G. Turgeon mol
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306 R. Debuchy and B.G. Turgeon gen
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308 R. Debuchy and B.G. Turgeon int
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310 R. Debuchy and B.G. Turgeon Fig
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312 R. Debuchy and B.G. Turgeon pro
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314 R. Debuchy and B.G. Turgeon B.
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316 R. Debuchy and B.G. Turgeon 2.
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318 R. Debuchy and B.G. Turgeon in
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320 R. Debuchy and B.G. Turgeon be
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322 R. Debuchy and B.G. Turgeon Lee
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16 Fruiting-Body Development in Asc
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phae originating from the base of a
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Table. 16.1. (continued) Fruiting B
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(Raju 1992). When male-sterile muta
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1. nsd (never in sexual development
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egular intervals; both effects requ
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the balance between sexual and asex
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ductionofeitherpheromoneisdirectlyc
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(Catlett et al. 2003). Eleven genes
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negative mutation of KREV-1 resulte
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through MAP kinase modules to cell-
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The fact that fruiting-body formati
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Balestrini R, Mainieri D, Soragni E
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point mutation of the beta subunit
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Mitchell TK, Dean RA (1995) The cAM
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YamashiroCT,EbboleDJ,LeeBU,BrownRE,
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358 L.A. Casselton and M.P. Challen
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376 M. Feldbrügge et al. different
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378 M. Feldbrügge et al. Fig. 18.3
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380 M. Feldbrügge et al. et al. 20
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382 M. Feldbrügge et al. for unbia
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384 M. Feldbrügge et al. In U. may
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386 M. Feldbrügge et al. Neurospor
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388 M. Feldbrügge et al. Bernards
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390 M. Feldbrügge et al. O’Donne
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19 The Emergence of Fruiting Bodies
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2003; Kües et al. 2004) are the fi
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(Manachère 1988), C. cinereus (Tsu
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emergence of fruiting bodies. In th
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Rather, development is arrested in
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10 nm thick is highly insoluble and
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it has been suggested that hydropho
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expression in the stipe suggests th
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Cooper DNW, Boulianne RP, Charlton
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Lu BC (1974) Meiosis in Coprinus. R
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Takagi Y, Katayose Y, Shishido K (1
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20 Meiosis in Mycelial Fungi D. Zic
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Fig. 20.1. A-E Diagrammatic represe
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etween dispersed DNA repeats. In N.
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Fig. 20.2. Meiotic recombination. S
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mechanism whereby homologues locate
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An important component of the bouqu
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observed when the mammal LE compone
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A. Chromosome and Sister-Chromatid
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ascus plus ascospore morphogenesis
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syndrome)discoveredasageneinvolvedi
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Kitajima TS, Kawashima SA, Watanabe
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Rossignol J-L, Faugeron G (1995) MI
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Biosystematic Index Absidia glauca
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violaceum 153 Microsporum 149 Mimos
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Subject Index ABC transporter 382 a
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fluffy 270, 276 formin 8, 10, 13, 1
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MAT protein interaction 308, 315 ma
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sporangiophore 234, 242, 246-252, 2
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