3/8/17 fungus - any of those plants without leaves, flowers or green coloring matter, growing on other plants or on decaying matter and including mushrooms, toadstools and molds. Oxford American Dictionary, 1980 fungus - any chemoheterotrophic eukaryote that reproduces with spores and has cell walls at some stage of life. Plant Biology 2nd Ed. (2006) Rost, Barbour, Stocking, Murphy chemoheterotrophic - an organism that obtains both energy and carbon from organic sources. Some fungi obtain nutrition from living organisms such as mycorrhizae and lichens. Others are saprobes - organisms that feed on dead organic material. Main differences between plants and fungi: 1) both have cell walls but the fungi use chitin for cell walls while plants use cellulose; 2) plants are photoautotropic, fungi are chemoheterotrophic. Because fungi have a rigid cell wall, that cannot engulf food as animal cells can. They secrete digestive enzymes and transport the products (sugars, amino acids, peptides) through the plasma membrane. Kingdom Fungi - monophyletic group of eukaryotic heterotrophs with chitinous cell walls. dikaryomycete - has a dikaryotic mycelium stage with pairs of haploid nuclei in each septate compartment. coenomycetes - lack a dikaryotic stage and have aseptate hyphae. dikaryomycetes coenomycetes Ascomycota Glomeromycota Basidiomycota Zygomycota and Chytridiomycota polymer of an N-acetylglucosamine Phyla in Kingdom Fungi Fungal septa present in dikaryomycetes Chytridiomycota - artificial (non monophyletic) phylum containing all fungi that have swimming reproductive cells. Zygomycota - fungi with zygospores, aseptate hyphae, mitosporangia and lack of swimming cells. dikaryomycetes Glomeromycota monophyletic, members engage in arbuscular endomycorrhizae Septa of Ascomycetes Dolipore septa of Basidiomycetes coenomycetes Ascomycota Glomeromycota Basidiomycota Zygomycota and Chytridiomycota zygospore 1 3/8/17 most fungi develop a mycelium composed of branching hyphae Phyla (continued) Ascomycota - monophyletic phylum, fungi that produce sexual spores in asci, mycelia have septate hyphae, there is a dikaryotic stage in sexual reproduction, and they spread primarily by releasing conidia (asexual mitospores). Basidiomycota - all members extrude myospores from a basidium, mycelia are septate and a dikaryotic stage is present. septate aseptate Mycelium - branching network of hyphae ascospores in asci basidium Some fungi live as single cells (yeasts) Some are dimorphic consisting of hyphae or yeasts Hyphae allow the fungus to grow towards food sources by chemotropism (remember that a tropism is oriented growth in response to a directional signal). budding cell proteins, starch Hyphae grow by tip growth enzymes amino acids, sugar, etc. water solutes flow PRESSURE growth septum tip of hypha Fungi compete with bacteria for food and bacteria existed for 1 billion years before fungi. So how do fungi compete? yeasts Most fungi produce asexual spores to multiply, such as these mitospores from Penicillium conidia Although fungal cell division is slower (90 minutes for yeast, 20 minutes for bacteria). Fungi are able to grow toward food sources. Also they are able to redistribute the acquired nutrients through the mycelium and produce spores above the surface for dispersal. conidiophore Fungi secrete acid (H+) that inhibits bacterial growth. Fungi secrete antibiotics including alcohol that inhibit or kill bacteria. and sexual spores for diversity, 2 3/8/17 Are fungi closer to animals or plants? Oomycetes kingdom Fungi Acrasids kingdom Animalia Mycetozoa Original kingdom Fungi kingdom Plantae The original kingdom Fungi contained slime molds and oomcyetes (egg fungi). But this classification is not monophyletic and therefore does not make sense. Within the kingdom Fungi the phyla (or division) names end with mycota as in Basidiomycota or Ascomycota. Within phyla the class names end with mycetes such as Hymenomycetes the largest class in the Basidiomycota. In the Zygomycota, compatible hyphae fuse together to form a large cell that becomes the zygospore. In the Basidiomycota, basidiospores are exuded from the basidium zygospore In the Ascomcota, ascospores are contained in a cell called the ascus. One problem with using the morphology of sexual structures to classify fungi is that sexual reproduction has never been observed for thousands of species of fungi. basidium Sexual reproduction in the dikaryomcetes 95% of named fungal species are dikaryomycetes, what makes them sucessful? 1) a dikaryotic stage 2) septate hyphae. coenomycetes (lower fungi) Chytridiomycota Zygomycota dikaryomycetes (higher fungi) Sexual life cycle of a dikaryomycetes with a dikaryotic stage. By having 2 different nuclei in each cell, the number of meiotic events can be huge leading to high diversity of progeny. karyogamy zygote nuclei diploid Glomeromycota Ascomycota Basidiomycota dikaryotic stage n+n dikaryotic stage septate hyphae 5% of named fungal species 2n n + n mycelium n + n meiosis 1n 95% of named fungal species Fig. 20-11, p. 344 spores plasmogamy 1n mating mycelia haploid 3 3/8/17 Septate hyphae may be an advantage in producing more complex fungal structures. It may facilitate differentiation (different tissues need different genetic programs). The septum may also help in damage control. proteins, starch karyogamy - the fusion of two nuclei haploid - 1n, having one complete set of chromosomes. enzymes diploid - 2n amino acids, sugar, etc. water This example is from the zygomycota solutes flow plasmogamy - fusion of protoplasts, not accompanied by nuclear fusion PRESSURE growth tip of hypha septum Phylum Chytridiomycota - swimming reproductive cells mitosporangium c Phylum Chytridiomycota 1000 named species. none cause human diseases except for “swimmers itch” there are amphibian diseases caused by chytrids mitospores (zoospores) asexual loop d gametes e Here is a chytrid sporangium growing on pollen grains plasmogamy rhizoids zygote meiospores (zoospores) karyogamy and meiosis resting meiosporangium Phylum Zygomycota - fungi with zygospores, aseptate hyphae, mitosporangia and lack of swimming cells. Phylum Zygomycota - life cycle of Rhizopus stolonifer 1100 zygomycete species a few human diseases - zygomycosis occurs mainly in immunocompromised individuals. Transmission is usually by inhalation of spores. 4 3/8/17 Some common features of both zygomyces and chytridiomyces both meiospores and mitospores are made in sporangia the meiosporagium is a resting phase both make aseptate hyphae Phylum Glomeromycota - form mycorrhizae 157 species Differences: in the Zygomycota neither meiospores nor mitospores are motile Spore fusion takes place in the Chytridiomycota but is replaced by fusing hypha in the Zygomycota. 80% of plants make symbiotic mycorrhizal interactions with glomeromycetes form endomycorrhizae - fungal interactions “within the cell” arbuscules - “little trees”, hyphal proliferations within the cell wall of plant cells but not within the plasma membrane vesicles - swollen hyphal tips that store nutrients The fungus contributes water, nutrients such as phosphate to the plant and may make other - yet to be discovered - contributions. Sexual reproduction is unknown in glomeromycetes, they make huge mitospores (up to 1mm). Phylum Glomeromycota cortex root tissues epidermis Ascomycota and Basidiomycota soil root hair spore vesicle arbuscule hypha cortex cell fungal mycelium Ascomycota 32,000 named species Many human diseases: athelete’s foot, jock itch, ringworm Many plant diseases: apple scab, corn leaf blight, Dutch elm disease, powdery mildew Many benefits to humans also: bread, wine, beer, penicillin, morels (Morchella), truffles (Tuber), soy sauce, bleu cheese. ascospores (meiospores) mitospores are produced in the haploid generation both mating types make antheridia and ascogonia meiosporangia are called asci ascus conidia (mitospores) b n conidiophore m a ascogonium mycelia c l karyogamy and meiosis occur in the ascus Meiosis antheridium zygote Diploid (2n) d Haploid (1n) Plasmogamy Dikaryotic (n + n) 1n hypha meiospores are produced in the ascus and released k Karyogamy ascogenous ascus crozier hypha g j i e f n+n hypha h ascoma 5 3/8/17 Different fruiting bodies (ascoma) of the Ascomycota Single dikaryotic hyphae can lead to multiple asci Apothecia 1 ascogenous hypha (n + n) Perithecia Cleistothecia hymenium b a fertilized ascogonium c d e ascospores 3 2 1 h etc. asci 2 asci perithecial wall f g asci Ascomycetes and symbiosis The vast majority of lichens are composed of an ascomycete + either a cyanobacteria or an algae. Most lichens reproduce asexually by releasing soredia (singular is soredium). crustose - flat and crusty Soredia are a few hyphae wrapped around photosynthetic cells. foliose - leaflike fruiticose - branching crustose lichens Lichens also reproduce asexually through isidia (singular is isidium). isidia are fragments of lichens sterigma Basidiomycota life cycle no mitospores! Basidiomycetes use meiospores to reproduce and diversify a basidium 1n mycelia g Meiosis b basidioma is dikaryotic isidia karyogamy and meiosis occur in the basidium basidiospores are not enclosed in a cell basidiospores (meiospores) i h Diploid (2n) zygote f Haploid (1n) Dikaryotic (n + n) Plasmogamy n + n fusion cell Karyogamy c basidium n+n mycelium gill e basidioma d 6 3/8/17 Poisonous mushrooms (Basidiomycetes) The Destroying Angel (Amanita bisporigera). It is an ectomycorrhizal fungus, so it can be found near trees. It makes amatoxin and as little as 1/2 of a mushroom cap can be fatal. Symptoms appear 5 to 24 hours after ingestion, symptoms include vomiting, cramps, diarrhea, delirium, and convulsions. By the time symptoms appear, liver and kidney damage has occurred. Fly agaric (Amanita muscaria) It is poisonous and psychoactive (produces muscimol). It was used as an intoxicant in Siberia, and as an insecticide in Europe. It is ectomycorrhizal with many trees. A toxic dose for an adult is one cap and a fatal dose is 15 caps. The toxins are muscimol and ibotenic acid. There is no antidote. If properly prepared (detoxified) it can be safely consumed and there are traditions in Siberia, Japan and the Americas of eating Amanita. btw, morels are also toxic if not cooked properly. The volva or surrounding membrane of developing mushrooms is discintive. Deadly webcap (Cortinarius rubellus formerly speciosissimus) The death cap (Amanita phalloides), produces α-amanitin and there is no known antidote. Native to Europe. Especially associated with beeches, birches, pines, chestnuts, spruces. It is highly toxic and responsible for the majority of fungal poisonings worldwide. Toxicity is not reduced by cooking or drying, 1/2 of a cap is fatal for an adult. Initial symptoms are gastrointestinal and resolve within 2-3 days. Death occurs 6 to 16 days after ingestion due to liver and kidney damage. α-amanitin Jack o’Lantern mushroom (Omphalotus olearius) looks similar to a chanterelle. Not fatal but causes cramps vomiting and diarrhea. The gills are bioluminescent due to luciferase and luciferin. The gills glow (hence the name). It glows blue-green. Cortinarius mushrooms produce orellanine. If ingested, the first symptoms usually do not appear for 2–3 days and can in some cases take as long as 3 weeks. The first symptoms are similar to the common flu (nausea, vomiting, stomach pains, headaches, myalgia, etc), these symptoms are followed by early stages of renal failure (immense thirst, frequent urination, pain on and around the kidneys) and eventually decreased or nonexistent urine output and other symptoms of renal failure occur. If left untreated death will follow. orellanine Maintaining the dikaryotic stage: at each cell division two nuclei need to be sorted into each cell. This is done either by making the septum in the plane of the spindle fibers (left) or a clamp connection is formed (right). 7 3/8/17 A more complex septum may also permit basidiomycetes to have more complicated structures. Basidioma have many forms brackets Puff balls septum hyphal wall dolipore parenthesome pore ER Coral fungi In addition to the Hymenomycetes, the Basidiomycota contains classes Ustilagomycetes (smuts) and Urediniomycetes (rusts) that do not make large fruiting structures. These include some of the most destructive fungal pathogens. Basidiomyctes make huge numbers of spores example a shelf fungus can make 3 billion spores per day for 6 months. How are they released from the basidioma? droplet sterigma water film spore a compression By ballistospore release - the projection secretes sugars, attracting water from the air. When the droplet spreads around the b spore it compresses the sterigma rebound which acts as a spring to launch the spore. Corn smut 8
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