The Fungi. Introduction. Introduction. Introduction
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1 The Fungi Instructor: George Wong Office: St. John, 612B Phone: X hawaii.edu Office Hour: Wednesdays,3:30 Introduction Once upon a time fungi, algae and bacteria were classified as plants. Reason we still study these organism in botany. Mycology: The discipline within botany that studies fungi. A polyphyletic group of organisms that have common characteristics in broad sense. Introduction In the strict sense: Eukaryotes Cell wall composed of chitin. Heterotrophic Saprobes Parasites Forms mutualistic symbiosis Introduction In the strict sense: Thallus is yeast and/or hypha Yeast: single celled fungus reproduce by budding or fission. Hypha (pl.=hyphae):: Short, filamentous unit of fungal thallus. Masses of hyphae form mycelium (pl.=mycelia). May be septate or coenocytic. Septate Mycelium: Cells attached end to end. Where cells are joined are septa (sing.=septum). Occurs at regular intervals: septum Coenocytic mycelium: Mycelium forms a more or less continuous protoplasm, with infrequent septa: Coenocytic hypha Septa occurrence usually associated with reproductive structures Mycelium 1
2 Budding Yeast Yeast and Dimorphic Fungus Yeast Fission Yeast: mitosis followed by cell division. Dimorphic: Fungi have both mycelial and yeast thalli Rhodotorula: Yeast Mycelium Tremella mesenterica: Dimorphic In the strict sense: Mode of nutrition, absorption Fungi live in their food. Enzyme(s) released to act on Insoluble food. Break-down (digestion) of food. Fungi absorb digested food. If food is soluble, digestion is not necessary. In the strict sense: Summary of absorption In the strict sense: Reproduce by spores: Reproductive structure made up of one to several cells. Variable in shape and size. Sexual or asexual. Fungi that share the characteristics just described are believed to be closely related. However, In the past, definition was broader: Eukaryotic, heterotrophic, spore producing organisms. Included numerous organisms that are no longer classified as fungi. Will look at all organisms, past and present, classified as fungi. 2
3 Concept of fungi has changed, greatly, over the years. Thus, various classification schemes have also arisen. In studying phylogeny of fungi, major difficulty is lack of fossil records. Extant species used as basis to formulate phylogenetic schemes. Recent molecular studies have led to sweeping changes in concepts of phylogeny. Phylogeny of Fungi Division of true fungi Kingdom: Protista Phylum: Myxomycota (Acellular Slime Molds): Characteristic: Assimilative stage = amoebae (1N) and plasmodium (2N). Assimilative stage lacks cell wall. Obtains nutrient via phagocytosis. Currently classified with Protozoans. Myxomycetes Physarum polycephalum used as representative of Myxomycetes. Spores are borne in a dark, lobed sporangium on a yellow stipe (~ mm tall). Spores are globose and spiny. Lobed Sporangium Stipe Branched filaments called capillitium (pl.=capillitia) is also in sporangium. Capillitium Spores 3
4 Spore germinates to produce haploid amoeba. Amoeba can convert into swarm cells and back, again. Water No Water Under unfavorable conditions, amoebae form microcysts and become dormant. Normally, amoebae grow and divide by fission as they feed by means of phagocytosis. A critical mass of amoebae or swarm cells is reached, different mating strains fuse in pairs to form zygote. Zygote divides, repeatedly by mitosis, without cytokinesis to form plasmodium. Plasmodium also may produce a resistant stage, the sclerotium. Under arid condition, plasmodium loses water and becomes dormant. sclerotium 4
5 Plasmodium migrates and continues to feed. When food becomes limiting, plasmodium becomes reproductive and forms sporangia. Plasmodium X-section Plasmodium becomes segmented into discrete units, each one will give rise to a single sporangium. Protoplasm Hypothallus: outer layer of plasmodium. Will form outer layer of sporangium. Differential pressure exerted by protoplasm shapes sporangium into morphology characteristic of species. Protoplasm is forced out of stalk into sporangium Meiosis occurs in sporangium to produce spores. Filamentous capillitium produced from waste material from vacuoles. Peridium Variations in Myxomycete Structures Mature, multi-lobed sporangia. Peridium Didymium iridis, a sporangium without lobes, a more typical sporangium. Powdery material on peridium is CaCO3 Several sporangia Single sporangium 5
6 Variations in Myxomycete Structures Trichia favoginea, an example of a sessile sporangium. Variations in Myxomycete Structures Fuligo septica, a very large aethalium. Variations in Myxomycete Structures Hemitrichia serpula forms a plasmodiocarp fruiting structure. Variations in Myxomycete Structures Branched capillitium with spiral ornamentation. Variations in Myxomycete Structures Trichia favoginea elaters (=capillitium). Flagellated Fungi Term used here in descriptive sense and does not have any taxonomic significance. We will go over in detail, two phyla: Chytridiomycota Oomycota Briefly cover Hyphochytridiomycota 6
7 Flagellated Fungi Tree indicates that Oomycota and Hyphochytridiomycota are distantly related from true fungi. Chytridiomycota is classified with true fungi. Characteristics of Flagellated Fungi Thallus may be unicellular, have short hyphal fragments or mycelial. Chytridiomycota is classified with true fungi. Hyphae and mycelium, when present are coenocytic (=nonseptate). Zoospores produced in zoosporangium. Characteristics of Flagellated Fungi Based on Zoospore morphology and cell wall biochemistry, the three divisions can readily separated: Chytridiomycota: Single, posterior, whiplash flagellum. Cell wall composition mostly chitin. Oomycota: Two, posterior or more commonly lateral flagella; one tinsel type and the other whiplash type. Cell wall is mostly cellulose. Chytridiomycota zoospore Zoospores Oomycota primary and secondary zoospore Characteristics of Flagellated Fungi Based on Zoospore morphology and cell wall biochemistry, the three divisions can readily separated: Hyphochytridiomycota: Single, anterior, tinsel type flagellum. Cell wall composition celluluose. Division: Chytridiomycota Phylum is composed of a single class: Chytridiomycetes and two orders: Chytridiales: Thallus is simple, unicellular to short hyphal strands. Zoospores with loose aggregation of ribosome. Without a nuclear cap. Blastocladiales: Thallus always mycelial. Zoospores with a nuclear cap. 7
8 Chytridiomycota Zoospores Division: Chytridiomycota Morphologically, the Chytridiomycota does not appear related to the true fungi. However: Cell wall biochemistry is identical to Ascomycota and Basidiomycota and similar to Zygomycota. Sequencing of rdna also supports this relationship. Blastocladiales zoospore Chytridiales zoospore Division: Chytridiomycota Morphologically, the Chytridiomycota does not appear related to the true fungi. However: Cell wall biochemistry is identical to Ascomycota and Basidiomycota and similar to Zygomycota. Sequencing of rdna also supports this relationship. Order: Chytridiales Rhizophydium sp.: An example of a simple, unicellular species. Order: Chytridiales Order: Blastocladiales will be used as the representative life cycle for this order. Will begin life cycle with the gametothallus Rhizophydium on pine pollen: Four zoosporangia emerging from pollen. 8
9 Young gametothallus that has not yet produce gametangia. Mature gametothallus with terminal male gametangia and subterminal female gametangia. Anisogametes produced. Male and female gametes morphologically similar, with female gametes three times larger than male. Female gametangiium Male gametangiium Terminal male gametangium is orange and female is subterminal and colorless. Release of anisogametes. Female gametes release sirenin that attracts male gametes. Male produce parisin that attracts female gametes. Syngamy of gametes produce zygote that germinates to produce sporothallus with zoosporangia and resistant sporangia. Zoosporangia produce diploid zoospores that asexually reproduce the sporothallus. zoosporangia resistant sporangia Zoosporangia are colorless while resistant sporangia are dark brown, with thick, pitted walls. 9
10 Resistant sporangia undergo meiosis and produce haploid zoospores, after a dormancy period. Dormancy broken when sporothallus dies off and favorable condition for gametothallus growth occurs. 10
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