Chapter 19 : Hyphal Fusion

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This chapter focuses on hyphal fusion in filamentous ascomycete and basidiomycete species, with emphasis on the model ascomycete fungus, . It reviews the different types of hyphal fusion, its mechanistic basis, and the varied functions that it serves, and it compares hyphal fusion with processes of cell fusion in fungi and other eukaryotic species. Hyphal fusion between spores and spore germlings during colony initiation is very common. In members of the Ascomycota and Basidiomycota, hyphal fusion occurs during mating-cell fusion and during the formation and maintenance of the dikaryon during the sexual phase of the life cycle. Future comparison of different types of hyphal fusion at different stages during the fungal life cycle will be important to distinguish molecular components universally involved in cell fusion from those that are specific to individual cell fusion pathways. Many of the processes required for hyphal fusion in filamentous fungi during vegetative growth are also required during cell fusion processes in general, including signaling by diffusible substances, directed cell growth or movement towards each other, attachment of the two cell types to one another, production and targeting of enzymes to the attachment site, and fusion of the plasma membranes of the interacting cells. Understanding the molecular basis of hyphal fusion during vegetative growth in filamentous fungi may provide a paradigm for self-signaling and self-fusion mechanisms in eukaryotic microbial species, as well as provide a useful model for somatic cell fusion events in complex, multicellular species.

Citation: Read N, Fleißner A, Roca M, Glass N. 2010. Hyphal Fusion, p 260-273. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch19

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Image of FIGURE 1

. (A) Conidial anastomosis tubes (CATs) (c) that have formed directly from macroconidia (s) and fused with each other. Note that the germ tubes (g) are wider than the CATs. Bar = 5 μm. (From M. G. Roca, C.E. Jeffree, and N. D. Read, unpublished data.) (B) CATs (c) that have formed from germ tube (g) tips, grown towards each other, and made contact. Bar = 5 μm. (From Roca, Jeffree, and Read, unpublished.) (C) CATs (c) that have formed subapically from germ tubes (g) and have fused. Bar = 5 μm. (From , with permission.) (D) The CAT homing assay. The two conidia had germinated, and their CATs were homing towards each other (0 min). The left-hand germling was repositioned (here shown 4 min after repositioning). The CAT tips then changed their orientation to home back towards each other (15 and 21 min) before making contact (25 min) and subsequently fusing (not shown). The left-hand conidium remained trapped throughout the entire 25-min period without apparent inhibition of CAT growth, homing, or fusion. The position of the trap in the germ tube (g) is represented by the crosshair in the circle. Note that the germ tube is significantly wider than either CAT. Bar = 10 μm. (From , with permission.) (E) Hyphal fusion in a mature colony that has resulted in a complex interconnected hyphal network. Bar = 100 μm. (From K.M. Lord and N. D. Read, cover image for 2008 issues of , with permission.) (F) A comparison of the morphology of anastomosis between fusion hyphae in a mature wild-type colony (note fusion pores [asterisks]) and a fusion mutant () in which fusion does not occur. Hyphae imaged by confocal microscopy after staining with calcofluor white M2R. Bar = 10 μm. (From Fleißner et al., 2005, with permission.) (G) Trichogyne that has homed towards, and wrapped around, a macroconidium (arrow) of opposite mating type. Bar = 20 μm. (From H.C. Kuo, C.E. Jeffree, and N. D. Read, unpublished data).

Citation: Read N, Fleißner A, Roca M, Glass N. 2010. Hyphal Fusion, p 260-273. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch19
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Image of FIGURE 2

(A) Diagram of crozier cell fusion in a typical ascomycete species. (B) Diagram of clamp cell fusion in a basidiomycete species. See “Hyphal Fusion in a Mature Colony” for details.

Citation: Read N, Fleißner A, Roca M, Glass N. 2010. Hyphal Fusion, p 260-273. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch19
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Image of FIGURE 3

Working model of the signaling involved during vegetative hyphal fusion in (see the text for details). The self-signaling ligand and receptor responsible for the process of self-fusion are unknown. GPIG1, GPIT1, GPIP1, GPIP2, and GPIP3 are involved in the GPI protein anchoring pathway ( ). It is not known which stage(s) of vegetative hyphal fusion these proteins regulate. HAM-2 is a predicted transmembrane protein ( ), although it is not known which cellular membrane it is associated with. NRC1-MEK2-MAK2 (Li et al., 2005; ), MIK1-MEK1-MAK1 ( ), and the OS4-OS5-OS2 ( ) are three MAP kinase pathways, and PP1 ( ) is the transcription factor predicted to be at the base of the MAK2 pathway. Upstream elements of the MAK1 and OS2 MAP kinase pathways and the stage(s) of vegetative hyphal fusion that they regulate are unknown. Mutations in suppress the vegetative hyphal fusion defect of ( ). SO is an ascomycete-specific WW domain protein ( ).

Citation: Read N, Fleißner A, Roca M, Glass N. 2010. Hyphal Fusion, p 260-273. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch19
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Genes required for hyphal fusion in and their roles in the fusion process

Citation: Read N, Fleißner A, Roca M, Glass N. 2010. Hyphal Fusion, p 260-273. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch19

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