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Fungi Chapter 31 HIT PICTURE FOR HYPERLINK Figure 31.1
Figure 31.1 Can you spot the largest organism in this forest? HIT PICTURE FOR HYPERLINK 1
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Mighty Mushrooms Fungi are essential for the well-being of most terrestrial ecosystems because they break down organic material and recycle vital nutrients About 100,000 species of fungi have been described. (It is estimated there are actually 1.5 million species of fungi.) Despite their diversity, fungi share key traits, most importantly the way in which they derive nutrition
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Nutrition and Ecology Fungi are heterotrophs and absorb nutrients from outside of their bodies Fungi use enzymes to break down a large variety of complex molecules into smaller organic compounds Fungi are heterotrophic by absorption– they secrete hydrolytic enzymes which break down their food which they then absorb into their bodies The versatility of these enzymes contributes to fungi’s ecological success Fungi exhibit diverse lifestyles as decomposers, with some as parasites, and others are mutualists.
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Body Structure The most common body structures are multicellular filaments and single cells (yeasts) Some species grow as either filaments or yeasts; others grow as both The morphology of multicellular fungi enhances their ability to absorb nutrients Fungi consist of mycelia (the collective name for the entire fungi), networks of branched hyphae (individual threads) adapted for absorption A mycelium’s structure maximizes its surface-to-volume ratio Fungal cell walls contain chitin
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Reproductive structure
Figure 31.2 Reproductive structure Hyphae Spore-producing structures Figure 31.2 Structure of a multicellular fungus 60 μm Mycelium (a mass of hyphae)
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Figure 31.3 Most fungi have hyphae divided into cells by septa, with pores allowing cell-to-cell movement of organelles Cell wall Cell wall Pore Septum Nuclei Nuclei (a) Septate hypha (b) Coenocytic hypha Figure 31.3 Two forms of hyphae Coenocytic fungi lack septa and have a continuous cytoplasmic mass with hundreds or thousands of nuclei
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Specialized Hyphae in Mycorrhizal Fungi
Some unique fungi have specialized hyphae called haustoria that allow them to penetrate the tissues of their host 7
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Mycorrhizae are mutually beneficial relationships between fungi and plant roots. Mycorrhizal fungi deliver phosphate ions and minerals to plants. In return, plants provide organic nutrients to the fungi. Most vascular plants have mycorrhizae. Mycorrhizal fungi colonize soils by the dispersal of haploid cells called spores Ectomycorrhizal fungi form sheaths of hyphae over a root and also grow into the extracellular spaces of the root cortex Arbuscular mycorrhizal fungi extend hyphae through the cell walls of root cells and into tubes formed by invagination of the root cell membrane
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Ectomycorrhizal fungi/
Arbuscular mycorrhizal fungi Ectomycorrhizal fungi/ flikr.com -->
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Concept 31.2: Fungi produce spores through sexual or asexual life cycles
Fungi propagate themselves by producing vast numbers of spores, either sexually or asexually Fungi can produce spores from different types of life cycles specifically a mushroom // general fungal reproduction Fungal nuclei are normally haploid, with the exception of transient diploid stages formed during the sexual life cycles Sexual reproduction requires the fusion of hyphae from different mating types Fungi use sexual signaling molecules called pheromones to communicate their mating type
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Plasmogamy is the union of cytoplasm from two parent mycelia
In most 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, nuclear fusion. During karyogamy, the haploid nuclei fuse, producing diploid cells The diploid phase is short-lived and undergoes meiosis, producing haploid spores The paired processes of karyogamy and meiosis produce genetic variation
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PLASMOGAMY (fusion of cytoplasm) Heterokaryotic stage Diploid (2n)
Figure Haploid (n) Another mycelium Heterokaryotic PLASMOGAMY (fusion of cytoplasm) Heterokaryotic stage Diploid (2n) Spore- producing structures KARYOGAMY (fusion of nuclei) SEXUAL REPRODUCTION ASEXUAL REPRODUCTION Mycelium Zygote Spores Figure Generalized life cycle of fungi (step 3) GERMINATION MEIOSIS GERMINATION Spores
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Asexual Reproduction In addition to sexual reproduction, many fungi can reproduce asexually Molds produce haploid spores by mitosis and form visible mycelia 13
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Other fungi that can reproduce asexually are yeasts, which are single cells
Instead of producing spores, yeasts reproduce asexually by simple cell division and the pinching of “bud cells” from a parent cell Some fungi can grow as yeasts and as filamentous mycelia 10 μm Parent cell Bud
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Many molds and yeasts have no known sexual stage
Mycologists have traditionally called these deuteromycetes These fungi are reclassified once their sexual stage is discovered Mycologists can now also use genomic techniques to classify fungi Fungi and animals are more closely related to each other than they are to plants or other eukaryotes
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Fungi have radiated into a diverse set of lineages
Figure 31.10 Chytrids (1,000 species) Fungi have radiated into a diverse set of lineages Hyphae 25 μm Zygomycetes (1,000 species) Glomeromycetes (160 species) Fungal hypha 25 μm Ascomycetes (65,000 species) Figure Exploring fungal diversity Basidiomycetes (30,000 species)
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Chytrids Flagellum 4 μm Chytrids (phylum Chytridiomycota) are found in terrestrial, freshwater, and marine habitats including hydrothermal vents They can be decomposers, parasites, or mutualists Molecular evidence supports the hypothesis that chytrids diverged early in fungal evolution Chytrids are unique among fungi in having flagellated spores, called zoospores
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Video: Phlyctochytrium Zoospore Release
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Zygomycetes The zygomycetes (phylum Zygomycota) exhibit great diversity of life histories They include fast-growing molds, parasites, and commensal symbionts The life cycle of black bread mold (Rhizopus stolonifer) is fairly typical of the phylum Its hyphae are coenocytic with septa found only where reproductive cells are found Horizontal hyphae spread out over food, penetrate it, digest it and absorb the nutrients In the asexual phase, hundreds of asexual sporangia produce haploid spores at the tips of upright hyphae 19
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Gametangia with haploid nuclei
Figure 31.12 Haploid (n) Heterokaryotic (n + n) Diploid (2n) PLASMOGAMY Mating type (−) Gametangia with haploid nuclei Mating type (+) 100 μm Young zygosporangium (heterokaryotic) Rhizopus growing on bread New mycelia SEXUAL REPRODUCTION Dispersal and germination of spores Zygo- sporangium KARYOGAMY Sporangia Figure The life cycle of the zygomycete Rhizopus stolonifer (black bread mold) Diploid nuclei Sporangium ASEXUAL REPRO- DUCTION MEIOSIS Mycelium 50 μm
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The zygomycetes are named for their sexually produced zygosporangia
Zygosporangia are the site of karyogamy and then meiosis Zygosporangia, which are resistant to freezing and drying, can survive unfavorable conditions Some zygomycetes, such as Pilobolus, can actually “aim” and shoot their sporangia toward bright light 0.5 mm
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Figure 31.14 Glomeromycetes The glomeromycetes (phylum Glomeromycota) were once considered zygomycetes They are now classified in a separate clade Glomeromycetes form arbuscular mycorrhizae Figure Arbuscular mycorrhizae 2.5 μm
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Ascomycetes Ascomycetes (65,000 species) Ascomycetes (phylum Ascomycota) live in marine, freshwater, and terrestrial habitats Ascomycetes produce sexual spores in saclike asci contained in fruiting bodies called ascocarps Ascomycetes are commonly called sac fungi Ascomycetes vary in size and complexity from unicellular yeasts to elaborate cup fungi and morels
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Morchella esculenta Tuber melanosporum Figure 31.15
Figure Ascomycetes (sac fungi) Morchella esculenta Tuber melanosporum
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Ascomycetes include plant pathogens, decomposers, and symbionts (About 40% of ascomycete species live with green algae or cyanobacteria in mutualistic associations called lichens.) Ascomycetes reproduce asexually by enormous numbers of asexual spores called conidia = naked spores that develop in long chains or clusters at the tips of special hyphae called conidiophores Conidia are not formed inside sporangia; they are produced asexually at the tips of specialized hyphae called conidiophores Neurospora crassa, a bread mold, is a model organism with a well-studied genome 25
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Sexual Reproduction in Ascomycetes
Plasmogamy occurs between two parental fungi. Hyphae are extended that are partitioned by septa into dikaryotic cells, each with two haploid nuclei representing two parents. The cells at the tip of these dikaryotic hyphae develop into asci. Within an ascus, karyogamy combines the two parental genomes, and meiosis forms four genetically different nuclei forming eight ascospores. 26
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Figure 31.17 The life cycle of Neurospora crassa, an ascomycete.
Conidia; mating type () Key Dispersal Haploid (n) Germination Mating type () Dikaryotic (n n) ASEXUAL REPRODUCTION Diploid (2n) Hypha PLASMOGAMY Ascus (dikaryotic) Conidiophore Mycelia Dikaryotic hyphae Mycelium Germination SEXUAL REPRODUCTION KARYOGAMY Dispersal Diploid nucleus (zygote) Asci Eight ascospores Ascocarp Figure The life cycle of Neurospora crassa, an ascomycete. Four haploid nuclei MEIOSIS 27
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Basidiomycetes Basidomycetes (phylum Basidiomycota) include mushrooms, puffballs, and shelf fungi Some basidiomycetes form mycorrhizae, and others are plant parasites The phylum is defined by a clublike structure called a basidium, a transient diploid stage in the life cycle The basidiomycetes are also called club fungi Many basidiomycetes are decomposers of wood Basidiomycetes (30,000 species)
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Shelf fungi Puffballs Maiden veil fungus Figure 31.17
Figure Basidiomycetes (club fungi) Puffballs Maiden veil fungus
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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 called basidiocarps Mushrooms are examples of basidiocarps The numerous basidia in a basidiocarp are sources of sexual spores called basidiospores
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1 μm PLASMOGAMY Dikaryotic mycelium Mating type (−) Mating type (+)
Figure 31.18 PLASMOGAMY Dikaryotic mycelium Mating type (−) Mating type (+) Gills lined with basidia Haploid mycelia SEXUAL REPRODUCTION Basidiocarp (n + n) Dispersal and germination Basidiospores (n) Basidia (n + n) Basidium Basidium with four basidiospores Figure The life cycle of a mushroom-forming basidiomycete Basidium containing four haploid nuclei KARYOGAMY MEIOSIS Haploid (n) Dikaryotic (n + n) Diploid (2n) 1 μm Basidiospore Diploid nuclei
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Basidiomycetes can produce mushrooms quickly
Some species may produce “fairy rings”
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Concept 31.5: Fungi play key roles in nutrient cycling, ecological interactions, and human welfare
Fungi interact with other organisms as decomposers, mutualists, and pathogens: Fungi are efficient decomposers of organic material including cellulose and lignin. They perform essential recycling of chemical elements between the living and nonliving world. Fungi form mutualistic relationships with plants, algae, cyanobacteria, and animals-- all of these relationships have profound ecological effects
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Fungus-Plant Mutualisms
Mycorrhizae are enormously important in natural ecosystems and agriculture Plants harbor harmless symbiotic endophytes, fungi that live inside leaves or other plant parts Endophytes make toxins that deter herbivores and defend against pathogens Most endophytes are ascomycetes
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Endophyte not present; pathogen present (E−P+)
Figure 31.20 Results Endophyte not present; pathogen present (E−P+) Both endophyte and pathogen present (E+P+) 30 15 20 10 Leaf mortality (%) Leaf area damaged (%) 10 5 Figure Inquiry: Do fungal endophytes benefit a woody plant? E−P+ E+P+ E−P+ E+P+
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Fungus-Animal Mutualisms
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 use the digestive power of fungi by raising them in “farms”
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Lichens Lichens are important pioneers on new rock and soil surfaces
A lichen is a symbiotic association between a photosynthetic microorganism and a fungus Millions of photosynthetic cells are held in a mass of fungal hyphae The photosynthetic component is green algae or cyanobacteria which occupy an inner layer below the lichen surface The fungal component is most often an ascomycete The symbioses are so complete that lichens are given scientific names The algae provide carbon compounds, cyanobacteria also provide organic nitrogen, and fungi provide the environment for growth The fungi of lichens can reproduce sexually and asexually--Asexual reproduction is by fragmentation or the formation of soredia, small clusters of hyphae with embedded algae 37
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Figure 31.22 A foliose (leaflike) lichen A fruticose (shrublike) lichen Figure Variation in lichen growth forms Lichens are sensitive to pollution, and their death can be a warning that air quality is deteriorating Crustose (encrusting) lichens
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Ascocarp of fungus Soredia Fungal hyphae Algal layer 50 μm
Figure 31.23 Ascocarp of fungus Soredia Fungal hyphae Algal layer 50 μm Figure Anatomy of an ascomycete lichen Fungal hyphae Algal cell
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Fungi as Parasites About 30% of known fungal species are parasites or pathogens, mostly on or in plants Each year, 10% to 50% of the world’s fruit harvest is lost due to fungi Some fungi that attack food crops are toxic to humans Ergot of rye is caused by an ascomycete, and produces toxins. More than 40,000 people died from an epidemic of ergotism during the middle ages Ergotism is characterized by gangrene, nervous spasms, burning sensations, hallucinations, and temporary insanity.Ergots contain lysergic acid, the raw material for LSD
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(b) Tar spot fungus on maple leaves
Figure 31.24 (b) Tar spot fungus on maple leaves Figure Examples of fungal diseases of plants (a) Corn smut on corn (c) Ergots on rye
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Animals are much less susceptible to parasitic fungi than are plants
The general term for a fungal infection in animals is mycosis The chytrid Batrachochytrium dendrobatidis might be the cause of the recent decline in amphibians worldwide Ringworm and athlete’s foot are examples a human mycoses Systemic mycoses spread through the body For example, coccidioidomycosis produces tuberculosis-like symptoms Some mycoses are opportunistic For example, Candida albicans, which causes yeast infections 42
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Yellow-legged frogs killed by B. dendrobatidis infection N
Figure 31.25 California Sixty Lake Basin 2 004 ▲ Yellow-legged frogs killed by B. dendrobatidis infection N 05 2 Key Boundary of chytrid spread Figure Amphibians under attack Lake status in 2009: Frog population extinct Treatment lake: frogs treated with fungicides and released 06 2 08 2 07 2
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Practical Uses of Fungi
Most people have eaten mushrooms The fruiting bodies of certain mycorrhizal ascomycetes, truffles, are prized by gourmets for their complex flavors. The distinctive flavors of certain cheeses come from the fungi used to ripen them. The ascomycete mold Aspergillus is used to produce citric acid for colas. The yeast Saccharomyces cerevisiae is the most important of all cultured fungi, and is available in many strains as baker’s and brewer’s yeast. Contributing to medicine, some fungi produce antibiotics used to treat bacterial diseases. The first antibiotic discovered was penicillin, made by the common mold Penicillium. Fungi play an important role in molecular biology and biotechnology. Researchers use Saccharomyces to study the molecular genetics of eukaryotes. 44
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Genetic research on fungi is leading to applications in biotechnology
For example, scientists are using Saccharomyces to study homologs of the genes involved in Parkinson’s and Huntington’s diseases For example, insulin-like growth factor can be produced in the fungus Saccharomyces cerevisiae For example, Gliocladium roseum, a fungus that produces hydrocarbons similar to diesel fuel, could be used to produce biofuels
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Distinguishing Features
Figure 31.UN07 Fungal Phylum Distinguishing Features Chytridiomycota (chytrids) Flagellated spores Zygomycota (zygomycetes) Resistant zygosporangium as sexual stage Glomeromycota (arbuscular mycorrhizal fungi) Arbuscular mycorrhizae formed with plants Ascomycota (ascomycetes) Sexual spores (ascospores) borne internally in sacs called asci; vast numbers of asexual spores (conidia) produced Figure 31.UN07 Summary of key concepts: fungi lineages Basidiomycota (basidiomycetes) Elaborate fruiting body (basidiocarp) containing many basidia that produce sexual spores (basidiospores)
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Figure 31.1a Figure 31.1a What is the largest organism in this forest? (part 1: mushroom cluster)
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