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Thallus Development in the Chytridiomycota

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Thallus Development in the Chytridiomycota Thallus Development in the Chytridiomycota Rabern Simmons, PEET trainee University of Maine Mycological Society of America Annual Meeting July 29, 2009 Purpose • Provide a glossary for teachers and students of the Chytridiomycota, or chytrids – Slightly different vocabulary than other fungal groups – Allow students to accurately explain what they see – Provide coherent discourse of ideas To meaningfully construct this glossary . • Define morphological terms • Show examples and compare morphologies • Show evidence that similarity of form does not necessitate relatedness – Morphological trends occur in multiple lineages Chytridiomycota • True fungi that produce zoospores with single smooth, posterior flagellum • Molecular phylogenies hypothesize 2 classes – Monoblepharidomycetes • 1 order – Chytridiomycetes • 6 described orders • 3 clades with no ordinal description Traditional Morphological Taxonomy • Characters viewed by light microscopy (LM) – Possession of operculum • Lid-like portion of zoosporangium – Amount of thallus devoted to zoosporangium – Development of zoosporangium from zoospore cyst – Number of zoosporangia per thallus I. Thallus Morphology Holocarpic Eucarpic Olpidium sp. (currently incertae sedis) Most common chytrid form. Sparrow, 1960. Aquatic Phycomycetes. Holocarpic “chytrid” Zoosporangia Plant cell wall with zoospore discharge tubes Resting sporangia of fungus Olpidium brassicae in root cells (LM). Used with permission of D J S Barr. Eucarpic chytrids Rhizoids – Defining feature Robust and Sparse, fine and Isodiametric blunt-ended tapered . Just to name a few. Rhizoids May have subsporangial swelling, or apophysis Chytriomyces aureus JEL72 Unidentified Unidentified on chitin (Hoffman Cladochytriales on agar Spizellomycetales modulation contrast LM) (phase contrast LM) on agar (LM) II. Development of the Zoospore Cyst Endogenous Exogenous Zoospore cyst JEL369 Clydaea vesicula on agar Barr81 Powellomyces hirtus on agar (phase contrast LM) (Hoffman modulation contrast LM) Endogenous Growth: Swelling of zoospore cyst Rhizoidal axis Zoospore with flagellum JEL539 Maunachytrium keaense on agar (phase contrast LM) Exogenous Growth: Nuclear migration Zoospore zc cyst (zc) Zoosporangium Rhizoidal zc axis JEL129 Entophlyctis luteolus on agar (phase contrast LM) Exogenous-Endogenous Growth e.g. Blyttiomyces helicus Cytoplasm initially exits Cytoplasm re-enters zoospore cyst zoospore cyst into germ tube during maturation of zoosporangium Zoospore cyst Endogenous development Exogenous development Blyttiomyces helicus Apiculus: Apex of initial zoospore cyst On spruce pollen (LM) III. Number of zoosporangia Monocentric Polycentric Papilla Zoosporangium Rhizoids Rhizomycelium Rhizoids Zoosporangia JEL 95 Powellomyces sp. on agar JEL109 Polychytrium aggregatum on agar (Hoffman modulation contrast LM) (phase contrast LM) Monocentric Chytrids • Holocarpic or Eucarpic • Endogenous or Exogenous • Many Chytridiomycetes • Select Monoblepharidomycetes – Holdfast – anchor to substrate – Loss of flagellum Monocentric Monoblepharidomycete with holdfast Harpochytrium hedinii on agar (phase contrast LM) Emerson & Whisler 1968. Cultural studies on Oedogoniomyces and Harpochytrium, and a proposal to place them in a new order of aquatic phycomycetes. Monocentric, Holocarpic Monoblepharidomycete Previous cell wall Hyaloraphidium curvatum on agar (phase contrast LM) Polycentric chytrids • Giving rise to multiple zoosporangia • Eucarpic and Exogenous – Rare exceptions • Several Chytridiomycetes • Majority of Monoblepharidomycetes Polycentric Chytrids Rhizomycelium is coenocytic or rarely septate JEL109 Polychytrium aggregatum JEL610 Cladochytrium sp. on agar (phase contrast LM) on agar (phase contrast LM) Pseudosepta in Gonapodya sp. (Monoblepharidomycetes) Gonapodya polymorpha on agar (phase contrast LM) Polycentricity continued Terminal or intercalary zoosporangia & swellings JEL109 Polychytrium aggregatum on agar M53 Monoblepharis sp. JEL611 Nowakowskiella sp. (phase contrast LM) On agar (phase contrast LM) on agar (phase contrast LM) More polycentrics on agar JEL137 Physocladia obscura JEL342 Catenomyces persicinus M36 Gonapodya polymorpha M6 Gonapodya prolifera Polycentric Monoblepharidomycete w/ holdfast & w/o rhizomycelium Oedogoniomyces spp. on agar (phase contrast LM) Emerson & Whisler 1968. Cultural studies on Oedogoniomyces and Harpochytrium, and a proposal to place them in a new order of aquatic phycomycetes. Problems with morphological taxonomy 1. Morphology in situ not necessarily the same as pure culture Lobulomyces poculatus eucarpic, endogenous, monocentric “Veil” of “Veil” may be absent cell wall Empty zoosporangium Developing zoosporangium with operculum on agar on spruce pollen (phase contrast LM) (phase contrast LM) Problems with morphological taxonomy 1. Morphology in situ not necessarily the same as pure culture 2. Morphology may differ on various nutrient media JEL95 Powellomyces sp. eucarpic, monocentric, generally exogenous zc zc zc zc? on agar (Hoffman modulation contrast LM) Problems with morphological taxonomy 1. Morphology in situ not necessarily the same as pure culture 2. Morphology may differ on various nutrient media 3. Morphology is not indicative of phylogeny Example 1: Molecular clades with known monocentric exogenous development Tree adapted from James et al. 2006. A molecular phylogeny of the flagellated fungi (Chytridiomycota) and description of a new phylum (Blastocladiomycota). Example 2: Molecular clades with known polycentric growth Tree adapted from James et al. 2006. A molecular phylogeny of the flagellated fungi (Chytridiomycota) and description of a new phylum (Blastocladiomycota). Conclusions • Large degree of variation seen in “simple, basal fungi” – Makes them interesting, but difficult to explain • Morphology remains useful for species identification – L. poculatus, B. helicus, etc. • However, morphological similarity is not necessarily an indication of relatedness – Must emphasize, or else we are defining but not educating!!!.
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