Neurospora possesses more cell types than are commonly recognized. We have been able to identify 28 morphologically distinct types. Having the cell types clearly defined will be important for genome annotation, describing new mutant phenotypes, and determining sites of gene expression

Bistis, George N.

Perkins, David D.

Read, Nick D.

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George N. Bistis

David D. Perkins

Nick D. Read

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Fungal Genetics Reports

Different cell types in Neurospora crassa

Kansas State University

Fungal Genetics Reports Volume 50 Article 8 Different cell types in Neurospora crassa George N. Bistis Drew University David D. Perkins Stanford University Nick D. Read University of Edinburgh Follow this and additional works at: https://newprairiepress.org/fgr This work is licensed under a Creative Commons Attribution-Share Alike 4.0 License. Recommended Citation Bistis, G. N., D.D. Perkins, and N.D. Read (2003) "Different cell types in Neurospora crassa," Fungal Genetics Reports: Vol. 50, Article 8. https://doi.org/10.4148/1941-4765.1154 This Regular Paper is brought to you for free and open access by New Prairie Press. It has been accepted for inclusion in Fungal Genetics Reports by an authorized administrator of New Prairie Press. For more information, please contact cads@k-state.edu. Different cell types in Neurospora crassa Abstract Neurospora possesses more cell types than are commonly recognized. We have been able to identify 28 morphologically distinct types. Having the cell types clearly defined will be important for genome annotation, describing new mutant phenotypes, and determining sites of gene expression. This regular paper is available in Fungal Genetics Reports: https://newprairiepress.org/fgr/vol50/iss1/8 Number 50, 2003 17Different cell types in Neurospora  crassaGeorge N. Bistis1, David D. Perkins2, and Nick D. Read31Department of B iology, Drew University, M adison, NJ 07940,   2Department of B iological Sciences, Stanford University,Stanford, CA 94305-5020, 3Department of Cell and Molecular Biology, University of Edinburgh, Rutherford Building,Edinburgh EH8 9QU, U.K.Fungal Genet. Newsl. 50:17-19Neurospora possesses more cell types than are commonly recognized. We have been able to identify 28morphologically distinct types. Having the cell types clearly defined  will be important for genome annotation, describing newmutant phenotypes, and determining sites of gene expression. ____________________________________________________________________________________ Neurospora is a morphologically complex multicellular organism with many more cell types than the unicellular yeastSaccharomyces.  Most workers are familiar with mycelia, macroconidia, perithecia, asci, and ascospores, but the diversity ofcell types produced by Neurospora may not be fully appreciated. Now that the products of specific genes can be localizedusing GFP and o ther fluorescent proteins, attention will be focused  increasingly on particular cell types that differ inmorphology, physiology, or developmental origin.  D istinguishing different cell types is also important for genome annotation.  For convenience, we need to use the terms ‘cell’ and ‘cell type’ rather loosely to cover both cellular elements such ashyphae and discrete cells such as spores (see discussion by Read, 1994).  The basic undifferentiated, totipotent cellular elementis the compartmentalized vegetative hypha at the colony periphery (the leader hypha).  Certain other cell types are comprisedof differentiated hyphae (e.g., fusion hyphae, ascogonia, trichogynes, ascogenous hyphae, asci, paraphyses, and periphyses). Atthe other extreme are highly differentiated nonhyphal cells such as ascospores, microconidia, and the different wall cells ofprotoperithecia and perithecia.Twenty-eight morphologically distinct cell types are listed and described below.  Designation of protoperithecia andmicroconidia as vegetative or sexual is arbitrary. Additional types or subtypes will no doubt be revealed.Vegetative phaseLeader (leading) hypha.  This is a wide, fast-growing hypha located at the colony periphery.  It consists of an apical, tip-growing hyphal compartment interconnected with subapical compartments separated by perforated septa that providecontinuity by allowing the passage of nuclei, other organelles, and cytoplasm. The leader hyphae undergo subapical branchingand their growth contributes to the increase in colony diameter (Robertson 1965). Trunk hypha.  This is located in the colony interior.  It is wide and composed of hyphal compartments that typically becomehighly vacuolate.  These hyphal compartments are typically shorter than those of leader hyphae because of the greaterfrequency of septa.  The septal pores of trunk hyphae are frequently occluded.Fusion hypha.  This is typically a narrow, dichotomously branching hypha that arises from a trunk hypha.  It exhibits positivetropisms, homing towards other fusion or trunk hyphae and anastomosing with them (Hickey et al., 2002).Aerial hyphae.  In contrast to the hyphae described above which grow in or on the surface of solid or liquid media, aerialhyphae grow away from the medium surface into the air.  This is believed  to be a result of forming hydrophobic hydrophobinproteins on their surfaces  (Sietsma and W essels, 1994). Macroconidiophore .  The specialized hypha that gives rise to macroconidia (Springer and Yanofsky, 1989).Macroconidium (blastoconidium).  Large, multinucleate (typically 3-6 nuclei) asexual spore that grows and develops byrepeated  budding of the apical cell of the conidiophore (Springer and Yanofsky, 1989).  Important as a dispersal agent and as amale fertilizing agent. Macroconidium (arthroconidium).  Large asexual, multinucleate (typically 3-6 nuclei) spore that arises by fragmentation of theconidiophore (Springer and Yanofsky, 1989).  Important as a dispersal agent and as a male fertilizing agent.Macroconidial germ tube.  Specialized hypha that emerges from the macroconidium and subsequently undergoes branchingand differentiates into the  vegetative hyphae of the young colony. Macroconidial (conidial) anastomosis tube.  Specialized hypha that emerges from the macroconidium but which is narrowerthan the germ tube.  It grows only a short distance and exhibits positive tropisms to other conidial anastomosis tubes withwhich it fuses (M.G. Roca and Read, unpubl.).Microconidiophore.  The specialized hypha that gives rise to microconidia (Maheshwari, 1999).Microconidium.  Small, uninucleate spore that primarily functions as a male fertilizing agent (M aheshwari, 1999). 1Bistis et al.: Different cell types in Neurospora crassaPublished by New Prairie Press, 201718 Fungal Genetics NewsletterMicroconidial germ tube.  Specialized  hypha that emerges from a microconidium but exhibits only limited growth(Maheshwari, 1999).Ascospore germ tube.  Specialized  hypha that emerges from an ascospore (usually one at each end) and subsequentlyundergoes branching and differentiates into the vegetative hyphae of the young colony.Sexual phaseAscogonium.  A wide, coiled septate hypha that arises as a specialized branch of a trunk hypha (Read, 1983).  It acts as the‘female organ’ during fertilization.Enveloping (or ascogonial investing) hyphae. Slender and highly branched hyphae that arise from the ascogonium orsurrounding vegetative hyphae, and which envelope and interweave around the ascogonium (Read, 1983).  The fusedenveloping hyphae develop into the different cells of the protoperithecial wall (Read, 1994).Trichogyne.  Specialized narrow and often branched hypha that emerges from hyphal compartments of the ascogonium.  Itexhibits a positive tropism toward cells of opposite mating type, with which it fuses and obtains fertilizing nuclei (Bistis,1981).Ascogenous hyphae.  Specialized hyphae that arise from the ascogonium and contain nuclei of bo th mating types.  Thesehyphae generate asci from specialized cells within their croziers (Read and Beckett, 1995; Raju, 1980, 1992).Terminal (apical) crozier cell.  Specialized apical compartment of the hook-shaped crozier. This cell, which is haploid anduninucleate, grows around toward the basal crozier cell with which it fuses to recover the dikaryotic state (Read and B eckett,1996; Raju, 1980, 1992). Basal (stalk) crozier cell.  Specialized  third compartment of a crozier. This cell, which is haploid and uninucleate, fuses withthe terminal crozier cell to recover the dikaryotic state (Beckett and Read, 1996; Raju, 1980, 1992).  Penultimate crozier cell (ascus m other cell).  Second compartment of a crozier. This cell is initially dikaryotic and haploid. Itthen becomes diploid after karyogamy has occurred, and develops into the ascus (Read and Beckett, 1996; Raju, 1980, 1992).Ascus.  The specialized  unbranched nonseptate hypha within which eight ascospores develop .  Its nucleus is initially diploidbut subsequently undergoes meiosis and an extra mitotic division to produce eight haploid nuclei. Each nucleus becomesenclosed within an ascospore wall (Read and Beckett, 1996;  Raju, 1980, 1992). Ascospore.  A thick-walled, pigmented spore with characteristic longitudinal ribs on its surface.  Formed in the ascus followingmeiosis. Initially uninucleate but later multinucleate. Haploid, with nuclei derived from one of the products of meiosis (Raju,1980, 1992).   Paraphyses.  Specialized multinucleate and branched hyphae possessing thin walls and interspersed between asci within theperithecium.  T hey arise  from centrum pseudoparenchyma cells and  from in between asci (Read and  Beckett, 1985).  Centrum pseudoparenchymal cells.  Pseudoparenchymal cells derived from the fusion of the innermost layer of envelopinghyphae.  These thin-walled cells are initially very swollen in the protoperithecium but subsequently become flattened and formthe innermost layer of the two layered perithecial wall (Read and Beckett, 1986; Read, 1994).Outer protoperithecial and perithecial wall cells (not neck).  Thick-walled, pseudoparenchymatous cells with that are derivedfrom the fusion of the outer layer of enveloping hyphae (Read and Beckett, 1985; Read, 1994).Fringe hyphae (hair cells).  Slender, sometimes branched, hyphae that extend outward from protoperithecial and perithecialwall cells (Read, 1983).  Typically shorter than trichogynes Periphyses.  Specialized  short, slender and branched hyphae which line the neck canal.  They possess multiple, multinucleatehyphal compartments, and their tips bend towards the centre of the neck canal.  They differentiate into thepseudoparenchymatous neck wall cells around of rim of the ostiolar pore (Read and Beckett, 1985).Wall cell of perithecial neck.  Pseudoparenchymatous cells which, being derived  from periphyses, are distinct from wall cellsof the lower bulbous region of the perithecium (Read and Beckett, 1985).ReferencesBistis, G. N. 1981. Chemotropic interactions between trichogyenes and conidia of opposite mating-type in Neurospora  crassa . Mycologia 73: 959-975.Hickey, P. C., D. J. Jacobson, N. D. Read, and N . L. Glass  2002. Live-cell imaging of vegetative hyphal fusion in Neurosporacrassa .  Fungal Genet .Biol. 37: 109-119.2Fungal Genetics Reports, Vol. 50 [2003], Art. 8https://newprairiepress.org/fgr/vol50/iss1/8DOI: 10.4148/1941-4765.1154Number 50, 2003 19Maheshwari, R. 1999. Microconidia of Neurospora  crassa .  Fungal Genet. Biol. 26: 1-18.Raju, N. B . 1980. Meiosis and ascospore genesis in N eurospora.  Eur. J. Cell Biol. 23: 208-223.Raju, N. B . 1992. Genetic control of the sexual cycle  in Neurospora.  M ycol. Res.  96: 241-262.Read, N. D. 1983. A scanning electron microscopic study of the external features of perithecium development in Sordariahumana .  Can. J .  Bot.  61: 3217-3229.Read, N. D. 1994. Cellular nature and multicellular morphogenesis of higher fungi.  In Shape and Form in Plants and Fungi.(D. Ingram and A. Hudson, eds.), pp. 254-271.  Academic Press, London.Read, N. D., and A. Beckett  1985. The anatomy of the mature perithecium in Sordaria humana and its significance for fungalmulticellular development.  Can. J . Bot.  63: 281-296.Read, N. D., and A. B eckett  1996. Ascus and ascospore morphogenesis.  Mycol. Res. 100: 1281-1314.Robertson, N. F. 1965 . The mechanism of cellular extension and branching. In The Fungi: An Advanced Treatise. Vol. 1.  (G.C. Ainsworth and A. S. Sussman, eds.). Academic Press, New York. pp. 613-623.Sietsma, J. H., and J. G. H. W essels  1994. Apical wall biogenesis.  In The Mycota Vol 1, Growth, Differentiation andSexuality (J. G. H. Wessels and F. Meinhardt, eds.), pp. 129-141. Springer-Verlag, Berlin.Springer, M. L., and C. Yanofsky  1989. A morphological and genetic analysis of conidiophore development in Neurosporacrassa .  Genes Devel.  3: 559-571.3Bistis et al.: Different cell types in Neurospora crassaPublished by New Prairie Press, 2017

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Different cell types in Neurospora crassa

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