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Chapter 31: Fungi - Mighty Mushrooms • Fungi are diverse and widespread • They are essential for the well-being of most terrestrial ecosystems because they break down organic material and recycle vital nutrients Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Outline for fungi: • How do they obtain organic materials? • Body structure • Asexual vs. sexual life cycles • Fungal diversity – we will see examples of fungi • Relationships between fungi and other organisms • Lichens – fungi living along with bacteria or algae • Pathogenic fungi and uses of fungi Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Fungi are heterotrophs that feed by absorption • Despite their diversity, fungi share key traits, most importantly the way in which they derive nutrition • Fungi are heterotrophs but do not ingest their food • They secrete exoenzymes that break down complex molecules, and then they absorb the smaller compounds • Fungi exhibit diverse lifestyles: – Decomposers – Parasites – Mutualistic symbionts Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Representation of a saprobe and its mode of action to get energy source from the environment. Extracellular digestion in a saprobe with a cell wall. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Body Structure • The morphology of multicellular fungi enhances their ability to absorb nutrients • Fungi consist of mycelia, networks of branched hyphae adapted for absorption • Most fungi have cell walls made of chitin • Some fungi have hyphae divided into cells by septa – called septate hyphae, with pores allowing cell-to-cell movement • Coenocytic fungi are fungi whose hyphae lack septa Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The body structure of fungi Reproductive structure Hyphae Spore-producing structures 20 µm Mycelium The two types of hypha in a fungus Cell wall Cell wall Nuclei Pore Septum Septate hypha Nuclei Coenocytic hypha Porous septa in septate hyphae functions similar to gap junctions in animal cells and plamodesmata in plant cells – they allow easy transfer of molecules. In this case, it is transfer of molecules throughout the hypha. Specialized hyphae Nematode Hyphae 25 µm •Some unique fungi have specialized hyphae that allow them to penetrate the tissues of their host •These hyphae then secrete exoenzymes in order to digest outside food sources to absorb the released nutrients Hyphae adapted for trapping and killing prey Plant cell wall Fungal hypha Plant cell Haustorium Haustoria Plant cell plasma membrane Fungi produce spores through sexual or asexual life cycles •Fungi propagate themselves by producing vast numbers of spores, either sexually or asexually Key Heterokaryotic stage Haploid (n) Heterokaryotic (unfused nuclei from different parents) PLASMOGAMY (fusion of cytoplasm) Diploid (2n) KARYOGAMY (fusion of nuclei) Spore-producing structures Zygote Spores ASEXUAL REPRODUCTION Mycelium SEXUAL REPRODUCTION MEIOSIS GERMINATION GERMINATION Spore-producing structures Spores Sexual Reproduction • Plasmogamy is the union of two parent mycelia • In many fungi, the haploid nuclei from each parent do not fuse right away; they coexist in the mycelium, called a heterokaryon • In some fungi, the haploid nuclei pair off two to a cell; such a mycelium is said to be dikaryotic • Hours, days, or even centuries may pass before the occurrence of karyogamy, which is nuclear fusion • During karyogamy, the haploid nuclei fuse, producing diploid cells • The diploid phase is short-lived and undergoes meiosis, producing haploid spores Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Asexual Reproduction •In addition to sexual reproduction, many fungi can reproduce asexually •Many of these species grow as mold, sometimes on fruit, bread, and other foods Asexual Reproduction •Other fungi that can reproduce asexually are yeasts, which inhabit moist environments •Instead of producing spores, yeasts reproduce asexually by simple cell division Many molds and yeasts have no known sexual stage Mycologists have traditionally called these deuteromycetes, or imperfect fungi Parent cell Bud Fungi descended from an aquatic, single-celled, flagellated protist • Systematists now recognize Fungi and Animalia as sister kingdoms • In other words, fungi and animals are more closely related to each other than they are to plants or other eukaryotes • Molecular evidence supports the hypothesis that fungi and animals diverged from a common unicellular, flagellated ancestor • Fungi probably evolved before the colonization of land by multicellular organisms • The oldest undisputed fossils of fungi are only about 460 million years old • Fungi were among the earliest colonizers of land, probably as symbionts with early land plants Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings •Molecular analysis has helped clarify evolutionary relationships between fungal groups, although areas of uncertainty remain Arbuscular mycorrhizal fungi Sac fungi Club fungi Zygomycota Glomeromycota Ascomycota Basidiomycota •Fungi phylogeny is the subject of much research Chytridiomycota Fungi have radiated into a diverse set of lineages Chytrids Zygote fungi Chytrids • Chytrids (phylum Chytridiomycota) are found in freshwater and terrestrial habitats • They can be saprobic or parasitic • Molecular evidence supports the hypothesis that chytrids diverged earliest in fungal evolution • Chytrids are unique among fungi in having flagellated spores, called zoospores Flagellum Hyphae Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Zygomycetes • The zygomycetes (phylum Zygomycota) exhibit great diversity of life histories • They include fast-growing molds, parasites, and commensal symbionts • The zygomycetes are named for their sexually produced zygosporangia • The life cycle of black bread mold (Rhizopus stolonifer) is fairly typical of the phylum • Zygosporangia, which are resistant to freezing and drying, can survive unfavorable conditions • Some zygomycetes, such as Pilobolus, can actually “aim” their sporangia toward conditions associated with good food sources Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Life cycle of black bread mold (Rhizopus stolonifer) Key Haploid (n) Heterokaryotic (n + n) Diploid (2n) PLASMOGAMY Rhizopus growing on bread Mating type (+) Gametangia with Mating haploid nuclei type (–) 100 µm Young zygosporangium (heterokaryotic) SEXUAL REPRODUCTION Dispersal and germination Sporangia Sporangium ASEXUAL REPRODUCTION MEIOSIS Dispersal and germination 50 µm Mycelium KARYOGAMY Diploid nuclei Zygosporangium (heterokaryotic) Pilobolus can “aim” their sporangia toward conditions associated with good food sources •Microsporidia are unicellular parasites of animals and protists •They are now classified as zygomycetes Host cell nucleus Developing microsporidian Spore 10 µm Microsporidia Glomeromycetes •The glomeromycetes (phylum Glomeromycota) were once considered zygomycetes, but are now classified in a different phyla •Glomeromycetes form a distinct type of endomycorrhizae called arbuscular mycorrhizae Ascomycetes • Ascomycetes (phylum Ascomycota) live in marine, freshwater, and terrestrial habitats • The phylum is defined by production of sexual spores in saclike asci, usually contained in fruiting bodies called ascocarps • Ascomycetes vary in size and complexity, from unicellular yeasts to elaborate cup fungi • Ascomycetes reproduce asexually by enormous numbers of asexual spores called conidia • Conidia are not formed inside sporangia; they are produced asexually at the tips of specialized hyphae called conidiophores Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Ascomycetes The cup-shaped ascocarps (fruiting bodies) of Aleuria aurantia give this species its common name: orange peel fungus. The edible ascocarp of Morchella esculenta, the succulent morel is often found under trees in orchards. 10 µm Tuber melanosporum is a truffle, an ascocarp that grows underground and emits strong odors. These ascocarps have been dug up and the middle one sliced open. Neurospora crassa feeds as a mold on bread and other food (SEM). Ascomycetes life cycle Key Haploid (n) Dikaryotic (n + n) Conidia; mating type (–) Diploid (2n) Dispersal Germination ASEXUAL REPRODUCTION Mating type (+) Mycelium PLASMOGAMY Ascus (dikaryotic) Mycelia Conidiophore Dikaryotic hyphae extended from ascogonium SEXUAL REPRODUCTION Germination Dispersal Asci Diploid nucleus (zygote) Eight ascospores Four haploid nuclei Ascocarp KARYOGAMY MEIOSIS Basidiomycetes •Basidomycetes (phylum Basidiomycota) include mushrooms and shelf fungi Maiden veil fungus (Dictyphora), a fungus with an odor like rotting meat •The phylum is defined by a clublike structure called a basidium, a transient diploid stage in the life cycle •The life cycle of a basidiomycete usually includes a long-lived dikaryotic mycelium •In response to environmental stimuli, the mycelium reproduces sexually by producing elaborate fruiting bodies call basidiocarps Fly agaric (Amanita muscoria), a common species in conifer forests in the northern hemisphere Puffballs emitting spores Shelf fungi, important decomposers of wood Mushrooms are examples of basidiocarps Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Dikaryotic mycelium PLASMOGAMY •The numerous basidia in a basidiocarp are sources of sexual spores called basidiospores Mating type (–) •Asexual reproduction is much less common in basidiomycetes than in ascomycetes Haploid mycelia Mating type (+) SEXUAL REPRODUCTION Gills lined with basidia Basidiocarp (dikaryotic) Dispersal and germination Basidiospores Basidium with four appendages Basidium Basidia (dikaryotic) Basidium containing four haploid nuclei KARYOGAMY MEIOSIS Key Diploid nuclei 1 µm Basidiospore Haploid (n) Dikaryotic (n + n) Diploid (2n) Fungi have a powerful impact on ecosystems and human welfare • Fungi are efficient decomposers - They perform essential recycling of chemical elements between the living and nonliving world • Fungi form symbiotic relationships with plants, algae, and animals • All of these relationships have profound ecological effects by helping each other survive • Mycorrhizae are enormously important in natural ecosystems and agriculture • They increase plant productivity Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Ecological associations of fungi – their lifestyles • Influence fungi can have on other living organisms in the environment – Symbiotic relationship: Organisms that live in close nutritional relationship, required by one or both organisms – Types • Mutualism – both organisms benefit (mutualistic symbionts) • Commensalism – one organism benefits, other not harmed • Parasitism – host/microbe relationship, microbe benefits, host is harmed – Non-symbiotic relationship: Organisms are free-living, and do not rely on each other for survival – Types • Synergism – shared metabolism between members, not required • Antagonism - competition between microorganisms where some organisms growth is inhibited or they are destroyed by others Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mycorrhizae • Mycorrhizae are mutually beneficial relationships between fungi and plant roots • Ectomycorrhizal fungi form sheaths of hyphae over a root and also grow into the extracellular spaces of the root cortex • Endomycorrhizal fungi extend hyphae through the cell walls of root cells and into tubes formed by invagination of the root cell membrane • The fungi aid the plant’s uptake of nutrients and increase the surface area for water uptake. (root hairs also increase the total surface area of a plant’s roots!) • These relationships between the fungi and the plant roots allow vascular plants to grow tall. Without the fungi, the plants wouldn’t grow as tall as they would not take up nutrients as easily. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Fungi have a powerful impact on ecosystems and human welfare Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Fungus-Animal Symbiosis • Some fungi share their digestive services with animals • These fungi help break down plant material in the guts of cows and other grazing mammals • Many species of ants and termites use the digestive power of fungi by raising them in “farms” Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Lichens A fruticose (shrub-like) lichen A foliose (leaf-like) lichen Crustose (crust-like) lichens •Lichens are a symbiotic association of millions of photosynthetic microorganisms held in a mass of fungal hyphae Ascocarp of fungus Soredia Lichens Fungal hyphae Algal layer •The fungal component of a lichen is most often an ascomycete •Algae or cyanobacteria occupy an inner layer below the lichen surface to complete the lichen formation with the fungus Algal cell 10 µm Fungal hyphae Pathogens •About 30% of known fungal species are parasites, mostly on or in plants •Animals are much less susceptible to parasitic fungi than are plants •The general term for a fungal infection in animals is mycoses Corn smut on corn Tar spot fungus of sycamore leaves Ergots on rye Practical Uses of Fungi •Humans eat many fungi and use others to make many different cheeses, alcoholic beverages, and bread. •Genetic research on fungi is leading to applications in biotechnology •Antibiotics produced by fungi treat bacterial infections Staphylococcus Penicillium Zone of inhibited growth