Chapter 24 Lecture Outline See separate PowerPoint slides for all figures and tables preinserted into PowerPoint without notes. Copyright © McGraw-Hill Education. Permission required for reproduction or display. 1 A 460 million year old fossil fungus – one of the oldest known 2 Chapter 24 Fungi Chapter Outline: Evolutionary Relationships of the Kingdom Fungi Fungal Bodies and Feeding Fungal Asexual and Sexual Reproduction The Importance of Fungi in Ecology and Medicine Biotechnological Applications of Fungi 3 Evolutionary Relationships of the Kingdom Fungi Eukaryote supergroup Opisthokonta Fungi are most closely related to animals, but diverged over a billion years ago Includes certain protists, Kingdom Animalia, and Kingdom Fungi Fungi arose from protists related to Nuclearia – an amoeba that feeds by engulfing cells True fungi are a monophyletic group of over 100,000 species Does not include slime molds or oomycetes 4 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Supergroup Opisthokonta Basidiomycetes Ascomycetes AM fungi Zygomycetes Microsporidia Chytrids Cryptomycota Nuclearia (protist) Choanoflagellates (protists) Metazoa (animal kingdom) Kingdom Fungi Septate hyphae, dikaryotic hyphae, fruiting bodies Beneficial associations with photosynthetic organisms Primarily terrestrial habitat Rigid chitin cell wall, osmotrophic nutrition KEY Critical innovations Single flagellum 5 BIOLOGY PRINCIPLE Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. All species (past and present) are related by an evolutionary history There are more than 15 fungal phyla, but their relationships and names are still being determined. 6 Fungal cell walls Rigid cell walls are composed of chitin A tough, nitrogen-containing carbohydrate Except for the cryptomycota which lack cell walls Benefit Allows cells to resist high osmotic pressure resulting from feeding by absorption of small organic molecules Drawbacks Cells cannot engulf food due to rigid cell walls Restricts mobility of nonflagellated cells 7 Seven main groups of fungi Cryptomycota Chytrids Microsporidia Zygomycetes AM fungi Ascomycetes Basidiomycetes 8 Cryptomycota The earliest-diverging fungi Occur in soil and water Can produce flagella for reproduction Only fungi to lack a cell wall 9 Chytrids Microscopic aquatic species Have cell walls made of chitin Produce reproductive cells with flagella Only found in chytrids and cryptomycota Loss of flagella linked to ecological transition from aquatic habitats to land 10 Zygomycetes Several lineages of terrestrial fungi Have distinctive large zygotes called zygospores ex: Common black bread molds 11 Microsporidia Pathogens that can only reproduce inside the cells of an animal host. Linked to honeybee decline Very small size (1–4 μm) Single-celled, chitin-walled spores Strong chitin wall helps survival in the environment until they enter the bodies of animals 12 AM fungi Arbuscular mycorrhizal fungi Close symbiotic associations with plant roots Fungus provides plant with minerals Plant provides food for the fungus Fossils suggest that even early plants may have depended on these AM fungal associations 13 Ascomycetes Asci – unique reproductive structures Some ascomycetes cause disease Ecologically important as decomposers example: Edible truffles and morels 14 Basidiomycetes Basidia – club-shaped reproductive cells Very important decomposers and plant symbionts ~30,000 species Varied reproductive structures Mushrooms, puffballs, stinkhorns, shelves, rusts, smuts 16 Fungal Bodies and Feeding Fungi are most closely related to animals and share several opisthokont features Both heterotrophic – cannot produce their own food use absorptive nutrition – secrete enzymes and absorb small organic molecules Both Both store surplus food as the carbohydrate glycogen 17 Unique body form Most have mycelia composed of hyphae Most of the mycelium is diffuse and inconspicuous Fruiting bodies – visible reproductive structures Mushrooms Fruiting are one type bodies produce spores Spores Chitin-walled An reproductive cells adaptation to the terrestrial environment Transported by wind or by animals 18 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Fruiting body above the substrate Mated hyphae Spores Mycelium within substrate (such as soil) Region where hyphae mate, forming a fruiting body Unmated mycelium Different unmated mycelium 19 BIOLOGY PRINCIPLE Living organisms grow and develop Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. After a mating process occurs, mated hyphae produce fruiting bodies whose form fosters spore production and dispersal. In suitable sites, spores may germinate, producing new mycelia. Distinctive growth processes Mycelia grow quickly when food is plentiful Grow from the edges as hyphae extend their tips Narrow dimensions and extensive branching provides high surface area for absorption Osmosis important in growth – entry of water produces force for tip extension 21 22 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. (a) Mycelium growing in liquid medium (b) Mycelium growing on flat, solid medium a: © Agriculture and Agri-Food Canada, Southern Crop Protection and Food Research Centre, London ON; b: CDC Mycelium shape depends on substrate Long extensions in soil to reach food-rich areas Spherical in liquid medium Flat disk in petri dish Fungal Asexual and Sexual Reproduction Many fungi reproduce by microscopic spores that grow into a new organism Spores may be asexual clones Or from sexual reproduction with new allele combinations Asexual reproduction is ideal for rapid spread No need to find a mate No fruiting body No meiosis 24 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Conidia Asexual spores grown at the tips of hyphae Aspergillus versicolor Causes skin and lung infections in vulnerable patients 69 µm © Dr. Dennis Kunkel Microscopy/ Visuals Unlimited 25 Medically important fungi that reproduce primarily by asexual means include Athlete’s foot fungus (Epidermophyton floccosum) Infectious yeast (Candida albicans) Budding yeast, Saccharomyces cerevisiae Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Can reproduce either sexually or by asexual budding Daughter cell (bud) Mother cell 26 © Medical-on-Line/Alamy Sexual reproduction Involves union of gametes, zygote formation and meiosis Gametes of most fungi are cells from hyphal branches Mating types differ biochemically Hyphal branches fuse between compatible mating types 27 Most sexual organisms have plasmogamy (fusion of gametes’ cytoplasm) followed by karyogamy (fusion of gametes’ nuclei) In fungi, after plasmogamy, nuclei may remain separate for a long time Mycelium is dikaryotic or heterokaryotic Some fungi persist as dikaryons, producing clones that can live for hundreds of years Dikaryotic mycelia are functionally diploid Eventually, dikaryotic mycelia produce fruiting bodies, the next stage of reproduction 28 Fruiting bodies Mated mycelia will produce a fleshy fruiting body when conditions are right Fruiting structure disperse haploid spores that grow into mycelia If a haploid mycelium meets a compatible mating type, they fuse (mate) and the cycle repeats Fruiting body structures aid spore dispersal Puffballs puff spores out onto wind currents Stinkhorns stink, and attract flies that carry off spores are underground – but their scent attracts animals that dig them up Truffles 29 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Spores in a sticky matrix (a) Fruiting bodies adapted for dispersal of spores by wind (b) Fruiting body adapted for dispersal of spores by insects a: © Felix Labhardt/Taxi/Getty Images; b: © Bob Gibbons/ardea.com 30 31 Many fungi produce substances in the fruiting body to deter consumption Toxins can cause liver failure requiring a transplant Hallucinogenic or psychoactive substances Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Ergot 32 © David Q. Cavagnaro/Getty Images The Importance of Fungi in Ecology and Medicine Decomposer fungi are essential components of the Earth’s ecosystems Work with bacteria Only certain bacteria and fungi can break down cellulose Release minerals to the soil and water 33 Some fungi are predators trapping tiny soil nematodes Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Nematode Hyphal loop © N. Allin & G.L. Barron/Biological Photo Service 34 Fungal pathogens Crop diseases caused by 5000 species Rust spores can be spread on the wind Human diseases Dermatophytes – athlete’s foot, ringworm Pneumocystis jiroveci and Cryptococcus neoformans infect people with weakened immune systems (as in AIDS) Blastomyces dermatitidis, Coccidioides immitis, and Histoplasma capsulatum infect the lungs In nature, fungal pathogens play important ecological role in controlling other species Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Wheat leaf tissue Puccinia graminis spores 0.1 mm (left): © Nigel Cattlin/Photo Researchers, Inc.; (right): © Herve Conge/ISM/Phototake 36 BIOLOGY PRINCIPLE Biology affects our society Recent analyses indicate that environmental changes linked to human activities correlate with increases in the incidence of new fungal pathogen infestations that threaten human health and agricultural sustainability. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. © Nigel Cattlin/Photo Researchers, Inc. Beneficial fungal associations Fungal associations with photosynthetic partners can be mutualistic Symbioses where both partners benefit Some animal species farm fungus for food Leaf-cutter ants, termites, beetles, salt marsh snail Mycorrhizal fungi associate with plant roots Lichens are partnerships between fungi and photocynthetic algae or bacteria 38 Mycorrhizae Association between the hyphae of certain fungi and the roots of most seed plants More than 80% of terrestrial plants have mycorrhizae Plants receive increased supply of water and mineral nutrients Fungi get organic food molecules from the plants Two most common types are endomycorrhizae (within roots) and ectomycorrhizae (on roots) 39 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Seedling root Mycorrhizal hyphae © Dr. D.P. Donelley and Prof. J.R. Leake, University of Sheffield, Department of Animal & Plant Sciences 40 Endomycorrhizae Fungal hyphae penetrate the spaces between root cell walls and plasma membranes and grow along the outer surface of the plasma membrane Arbuscular mycorrhizae (AM) form highly branched structures with high surface area Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Hyphae Arbuscules Cell wall Plasma membrane 49 µm (a) Micrograph of arbuscular mycorrhizae Root cells (b) Hyphae growing between cell walls and plasma membranes a: © Mark Brundrett 41 Ectomycorrhizae Coat root surface and grow between cells of roots Some species of oak, beech, pine, and spruce trees will not grow unless their ectomycorrhizal partners are also present Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Ectomycorrhizal hyphae Root cells Ectomycorrhizal hyphae coating a root tip (a) Ectomycorrhizal fruiting body (b) SEM of ectomycorrhizal hyphae (c) Hyphae invading intercellular spaces a: © Jacques Landry, Mycoquebec.org; b: © Courtesy of Larry Peterson and Hugues Massicotte 42 EVOLUTIONARY CONNECTIONS Comparison of genomes reveals how basidiomycete metabolism diversified Basidiomycete genomes show diverse metabolic pathways that help utilize organic carbon from plants Some decompose cellulose and lignin Break down dead trees, woody debris, leaf litter Some break down similar materials from animal dung Other species evolved ectomycorrhizal associations with living plants What genes are different to give species different capabilities? EVOLUTIONARY CONNECTIONS White rot fungi – decompose both cellulose and lignin Complex enzymatic pathways to break down the many types of chemical bonds Energetically expensive but give access to cellulose White rot fungal metabolism arose 300 mya during the Carboniferous Brown rot fungi – break down cellulose, leave lignin This new set of enzymes is why there are no major plant carbon deposits since then! Evolved from white-rot fungi but lost genes to degrade lignin, saving energy by not producing those enzymes Ectomycorrhizal fungi evolved in turn from the brown-rot fungi Lichens Partnerships of particular fungi and certain photosynthetic green algae and/or cyanobacteria 25,000 lichen species Not all descended from a common ancestor At least five separate fungal lineages Three major forms – crustose, foliose, fruticose Photosynthetic partner provides organic food molecules and oxygen Fungal partner provides carbon dioxide, water, and minerals 45 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. (a) Crustose lichen (b) Foliose lichen (c) Fruticose lichen (d) Microscopic view of a cross section of a lichen a: © Joe McDonald/Corbis; b: © Lee W. Wilcox; c: © Ed Reschke/Getty Images; d: © Lee W. Wilcox 46 Biotechnological Applications of Fungi Fungi convert inexpensive organic compounds into citric acid, glycerol, and antibiotics Distinctive flavor of blue cheese Saccharomyces cerevisiae for bread, beer and wine Replace chemical procedures that generate harmful waste products Wood pulp bleaching 47