Oomycota and Zygomycota Phylum: Oomycota Phylum is similar to Chytridiomycota in that there are unicellular, short hyphal to mycelial members and in the latter mycelium is nonseptate except where reproductive structures occur. Differ from Chytridiomycota in that cell wall component is usually cellulose and life cycles are always gametic. Oogamous: Oogonium, with immobile egg, and sperms in antheridium are naked nuclei. Lecture 02 Phylum: Oomycota Phylum was once thought to represent the transition from aquatic fungi to terrestrial fungi, but is no longer thought to be related to fungi. In one classification scheme they are placed in the Kingdom Stramenopila with Phaeophyta, Chrysophyta, etc. All have same cell wall composition, zoospore morphology and food storage material is mycolaminarin. mycolaminarin It is believed Oomycota had lost their chloroplast (not most parsimonious event). Order: Saprolegniales We will go over two orders in this phylum: Saprolegniales: Both primary and secondary zoospores may occur in the same life cycle. Each oogonium always produce several eggs. Peronsporales: Only secondary zoospore is produced. Each oogonium always produce one egg. Zoospores Chytridiomycota zoospore Oomycota primary and secondary zoospore Order: Saprolegniales Life cycle is quite different from Chytridiomycota. Will use the genus Saprolegnia as representative of this order. Gametic life cycle: Only phylum to have this type of life cycle. Sperms are naked, nonmotile gametes. Only flagellated fungi with this characteristic. 1 Order: Saprolegniales Summary of Saprolegnia life Asexual Cycle cycle. Go cover asexual cycle first. Zoosporangia of Saprolegnia are not well differentiated from hyphae. Zoosporangia with primary zoospores. zoospores Zoosporangia delimited by septum at base and dense protoplasm. Order: Saprolegniales Dense Protoplasm Order: Saprolegniales Basal Septum Terminal Zoosporangium of Saprolegnia Order: Saprolegniales Cyst will germinate to give rise to secondary zoospore (laterally flagellated). Order: Saprolegniales Primary zoospores (posterior flagella) are released from zoosporangium. Encyst Zoospore swims for a period of time and encysts: encysts rounds up and cell wall forms around itself. Primary zoospore Cyst Order: Saprolegniales Secondary zoospore encyst. Cyst germinates to give rise to diploid mycelium. Secondary zoospore 2 Order: Saprolegniales Summary of Saprolegnia life cycle: Sexual stage. stage The sexual stage is more uniform than the asexual. Most species are homothallic as shown here. Sexual Cycle Order: Saprolegniales Formation of Typically, oogonia gametangia and antheridia form on same mycelium. Nuclei migrate into gametangia, followed by formation of septa to “trap” nuclei. Meiosis occurs at this time. Antheridium Oogonium Order: Saprolegniales Order: Saprolegniales Fertilization Tubes Plasmogamy Typically, oogonia and antheridia form on same mycelium. Order: Saprolegniales Karyogamy occurs in oogonium to form zygotes. zygotes The mature zygotes are referred to as oospores. oospores Oogonia with several eggs is characteristic of this order. Antheridia are always difficult to see in live or prepared material. Order: Saprolegniales Oospore germinates to form the diploid mycelium. The mycelium will give rise to both the zoosporangia and gametangia, i.e., antheridia and oogonia. 3 Order: Saprolegniales Variations occur in zoosporangia. Two common genera: Saprolegnia: Following release of zoospores, an internal proliferation occurs to produce another zoosporangium. Order: Saprolegniales Achyla: Primary zoospores Encysted zoospores are released and encyst, immediately. Following release of zoospores, a new zoosporangium forms below the septum and adjacent to old zoosporangium. Order: Peronosporales This order has some of the most well known pathogens. They include the genera Pythium and Phytophthora. Differs from the Saprolegniales in producing only secondary zoospores in a zoosporangium that is differentiated from hyphae, and one egg per oogonium. Order: Saprolegniales Proliferation of zoosporangium through previous zoosporangium Previous zoosporangium Order: Saprolegniales Encysted zoospores Achyla empty zoosporangia with encysted zoospores at tip. Note formation of new, lateral zoosporangium. Order: Peronosporales Zoosporangium of Phytophthora tropicalis releasing secondary zoospores. Note that the zoosporangium can readily be distinguished from hyphae. 4 Phylum: Microsporidia Until the 1990’s this genus was classified as a very early eukaryote: Anaerobic, intracellular parasite of arthropods. Genus lacks mitochondria. Was originally assumed to have evolved before acquisition of mitochondria. Based on sequencing of protein tree of beta-tubulins is a fungus! Phylum: Microsporidia Spore germination: Anchoring Polar tube disk ruptured Pierce Pierce host host cell cell if if nearby nearby Phylum: Zygomycota Once thought to be more closely related to flagellated fungi and classified in Phycomycetes (literally algal fungi) with flagellated fungi. Cell wall material, however, is composed mostly of chitin. This and other fungal characters places this taxon with the fungi, in the strict sense. Phylum: Microsporidia Until the 1990’s this genus was classified as a very early eukaryote: Placement among other phyla of fungi uncertain at this point. Phylum: Zygomycota This phylum represents the first group of terrestrial fungi, but still has some characteristics in common with the flagellate fungi: Mycelium is coenocytic. Although asexual spores are not motile, they are still produced in a sporangium. Above characteristics not found in other terrestrial fungi Phylum: Zygomycota Characteristic that defines phylum is formation of the zygospore during sexual reproduction. Zygospore formation usually results from the fusion of isogametangia: isogametangia gametangia that are morphologically identical. The life cycle of Rhizopus stolonifer will be used as the representative of this phylum. 5 Rhizopus stolonifer Lifecycle Rhizopus stolonifer Lifecycle Rhizopus stolonifer life cycle includes both sexual and asexual reproduction. Sporangium development Stolon Nuclei migration, up sporangiophore Rhizoids Rhizoids form with contact with substrate Will cover asexual cycle first. Rhizopus stolonifer Lifecycle Nuclei migrate into swollen tip Developing Sporangium Columella forms trapping nuclei in future sporangium Rhizopus stolonifer Lifecycle Sporangium ruptures, releasing air-borne spores Germination Rhizopus stolonifer Lifecycle Mature Sporangium Cell wall forms around each nucleus to form sporangiospores or just spores Columella Rhizopus stolonifer Lifecycle Asexual reproduction will continue as long as there is available food. Sexual reproduction will not take place unless a different mating strain is present. The two mating strains are designated as “+” (plus) and “-” (minus). Mycelium 6 Rhizopus stolonifer Lifecycle Two mating strains are allowed to grow together. “-” (Minus) Mycelium “+” (Plus) Mycelium Rhizopus stolonifer Lifecycle Plus and minus Progametangia grow together due to phermonal response. Nuclei migrate to tips of progametangia. Suspensors Gametangia Sexual reproduction will take place where mating strains meet in center. Rhizopus stolonifer Lifecycle Gametangia fuse, followed by plasmogamy and karyogamy to form multinucleate zygote. zygote Zygote Rhizopus stolonifer Lifecycle Developing Germination of sporangium zygospore forms a sporangium containing “+” and “-” sporangiospores. Germinating zygospore Rhizopus stolonifer Lifecycle Zygote develops a thick, pitted wall referred to as the zygospore. zygospore Zygospore has a long dormancy period before meiosis and germination occurs. Does not disperse! Zygospore Zygorhynchus molleri Sporangiophore A species that is homothallic and produces anisogametangia. 7 Phylum: Zygomycota Phylum: Zygomycota There is a great deal of variation in the asexual stage of the Zygomycota: Rhizopus is representative of simplest and what is believed to be the earliest sporangia from which other variations are derived. Syncephalastrum produces spores in cylindrical sporangiole. sporangiole Phylum: Zygomycota Cunninghamella produces a one spored sporangiole and is believed to be the most recently derived condition. 8