Mushrooms, morels, and truffles, delights of the gourmet,

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1 26 Kingdom Fungi Fungal spores. The rounded earthstar (Geastrum saccatum) releases a puff of microscopic spores after the sac, which is about 1.3 cm (0.5 in) wide, is hit by a raindrop. This fungus is common in leaf litter under trees throughout North America. Jeff Lepore/Photo Researchers, Inc. KEY CONCEPTS Fungi are eukaryotic heterotrophs that absorb nutrients from their surroundings. A fungus may be a unicellular yeast or a filamentous, multicellular mold consisting of long, branched hyphae that form a mycelium. Most fungi reproduce both asexually and sexually by means of spores. According to current hypotheses, fungi evolved from a unicellular, flagellate protist and diverged into five main groups. Fungi are of major ecological, economic, biological, and medical importance. Mushrooms, morels, and truffles, delights of the gourmet, share a recent common ancestry with baker s yeast, the black mold that forms on stale bread, and the mildew that collects on damp shower curtains. All of these life-forms belong to kingdom Fungi, a diverse group of about 100,000 known species, most of which are terrestrial. Some biologists estimate that there may be as many as 1.5 million species of fungi. Fungi grow best in moist habitats, but they are found universally wherever organic material is available. They require moisture to grow, and they obtain water from a humid atmosphere as well as from the medium on which they live. When the environment becomes dry, fungi survive by going into a resting stage or by producing spores (see photograph) that resist desiccation (drying out). Some fungi grow to enormous size. In Washington State, a fungal clone (Armillaria ostoyae) has been identified that covers more than 1500 acres. This giant fungus, which is mainly underground, developed from a single spore that germinated more than a thousand years ago. The fungus has fragmented and is no longer one continuous body. Like prokaryotes, most fungi are decomposers that obtain nutrients from dead organic matter. They are vital members of ecosystems because they break down the organic compounds found in dead organisms, leaves, garbage, sewage, and other waste. When they break down organic compounds, carbon and other elements are released into the environment, where they are recycled. Many fungi form vital symbiotic relationships. For example, most terrestrial plants have fungal partners that live in close as- 555

2 sociation with their roots. The fungi help the plants obtain phosphate ions and other needed minerals from the soil. In exchange, the plants provide the fungi with organic nutrients. Some fungi live symbiotically with algae and cyanobacteria as lichens. Others are parasites and pathogens that cause disease in animals or plants. CHARACTERISTICS OF FUNGI Learning Objectives 1 Describe the distinguishing characteristics of kingdom Fungi. 2 Describe the body plan of a fungus. 3 Describe the life cycle of a typical fungus, including sexual and asexual reproduction. Although they vary strikingly in size and shape, all fungi (sing., fungus) are eukaryotes; their cells contain membrane-enclosed nuclei, mitochondria, and other membranous organelles. Fungi share certain key characters, including their way of obtaining nutrition. The optimum ph for most species is about 5.6, but various fungi can tolerate and grow in environments where the ph ranges from 2 to 9. Many fungi are less sensitive to high osmotic pressures than are bacteria. As a result, they can grow in concentrated salt solutions or in sugar solutions such as jelly, which discourage or prevent bacterial growth. Fungi also thrive over a wide temperature range. Even refrigerated food may be invaded by fungi. Fungi absorb food from the environment Unlike plants, fungi cannot produce their own organic materials from a simple carbon source (carbon dioxide). Like animals, fungi are heterotrophs. They depend on preformed carbon molecules produced by other organisms. However, fungi do not ingest food and then digest it in the body as animals do. Instead, they infiltrate a food source and secrete digestive enzymes onto it. Digestion takes place outside the body. When complex molecules are broken down into smaller compounds, the fungus absorbs the predigested food into its body. The fungus is very efficient at absorbing nutrients and growing. It rapidly converts nutrients into new cell material. If excessive amounts of nutrients are available, fungi store them, usually as lipid droplets or glycogen. Fungi have cell walls that contain chitin Like the cells of bacteria, certain protists, and plants, fungal cells are enclosed by cell walls during at least some stage in their life cycle. Fungal cell walls, however, have a different chemical composition from cell walls of other organisms. In most fungi, the cell wall consists of complex carbohydrates, including chitin, a polymer that consists of subunits of a nitrogen-containing sugar (see Fig. 3-11). Chitin is resistant to breakdown by most microorganisms. Chitin is also a component of the external skeletons of insects and other arthropods. Most fungi have a filamentous body plan Mycologists, biologists who study fungi, identify two main types of fungi, based on body plan: yeasts and molds. The simplest fungi are the yeasts, which are unicellular, with a round or oval shape. Yeasts are widely distributed in the soil; on leaves, fruits, and cured meats; and on and in our bodies. Yeasts are essential in making bread and fermenting alcoholic beverages. Some yeasts have been very important in medicine and as model organisms in modern biology (discussed later in this chapter). Most fungi are molds. The vegetative (nonreproductive) body plan of molds consists of long, branched, threadlike filaments called hyphae (sing., hypha) ( Fig. 26-1a and b). Hyphae are an adaptation to the fungal mode of nutrition. Growth occurs at the tips of the hyphae; as the hyphae elongate, the fungus grows into and infiltrates food sources. The fungus absorbs nutrients through its very large surface area. A hypha first develops from a unicellular spore. As hyphae grow, they form a tangled mass or tissuelike aggregation known as a mycelium (pl., mycelia). The cobweblike mold sometimes seen on bread is the mycelium of a fungus. What is not seen is the extensive mycelium that grows into the bread. Depending on environmental conditions, some fungi can alternate between a yeast phase and a phase in which they produce hyphae. In most fungi, hyphae are divided by cross walls, called septa (sing., septum), into individual cells containing one or more nuclei ( Fig. 26-1c and d). As we will discuss, the presence of septa is an important character in the two largest fungal phyla (which include the most complex fungi). The septa of many fungi are perforated by a pore that may be large enough to permit organelles to flow from cell to cell. Some fungi, called coenocytic fungi, lack septa. In these species, nuclear division is not followed by cytoplasmic division. As a result, a coenocytic fungus is one elongated, multinucleated, giant cell ( Fig. 26-1e). Fungi reproduce by spores Most fungi reproduce by means of microscopic spores, reproductive cells that can develop into new organisms ( Fig. 26-2). In most groups, spores are nonmotile. They are dispersed by wind, water, or animals. The air we breathe is filled with hundreds of thousands of fungal spores. Fungi produce spores either sexually or asexually. With asexual reproduction, new fungi are produced quickly, but there is little genetic variability. Sexual reproduction involves meiosis and generates new genotypes. Spores are usually produced on specialized aerial hyphae or in fruiting structures. Positioned up above the ground, spores can 556 Chapter 26

3 Hyphae (c) Dennis Drenner (a) A fungal mycelium, a mass of threadlike hyphae (10 cm [4 in] wide), growing on agar in a culture dish. In nature, fungal mycelia are rarely so symmetric. 25 µm (b) SEM of a mycelium of Blumeria graminis growing on a leaf (darker area underneath the mycelium). G. T. Cole/University of Texas/ Biological Photo Service (d) (e) (c) A hypha divided into cells by septa; each cell is monokaryotic. In some taxa the septa are perforated (as shown). (d) A septate hypha in which each cell is dikaryotic (has two nuclei). (e) A coenocytic hypha. Figure 26-1 Fungus body plan be easily dispersed. Structures in which spores are produced are called sporangia (sing., sporangium). The aerial hyphae of some fungi produce spores in large, complex reproductive structures, referred to as fruiting bodies. The familiar part of a mushroom is a large fruiting body. We do not normally see the bulk of the fungus, a nearly invisible mycelium buried out of sight in the rotting material or soil on which it grows. Many fungi reproduce asexually Yeasts reproduce asexually, primarily by forming buds that pinch off from the parent cell ( Fig. 26-3). Many species of molds also reproduce asexually. Spores are produced by mitosis and then released into the air or water. Conidiophores (from the Greek, meaning dust-bearers ) are specialized hyphae that produce asexual spores called conidia (sing., conidium). The arrangement of conidia on conidiophores varies from species to species. Most fungi reproduce sexually Many mold species reproduce sexually when they come into contact with other mating types. In contrast to the majority of animal and plant cells, most fungal cells contain haploid nuclei. In sexual reproduction, the hyphae of two genetically compatible mating types come together, and their cytoplasm fuses, a process known as plasmogamy. The resulting cell has two haploid nuclei, one from each fungus. This cell gives rise by mitosis to other cells with two nuclei. At some point the two haploid nuclei fuse. This process, called karyogamy, results in a cell containing a diploid nucleus known as a zygote nucleus. In some groups, the zygote nucleus is the only diploid nucleus. In the two largest fungal phyla, the ascomycetes and basidiomycetes (discussed later in this chapter), plasmogamy occurs (the hyphae fuse), but karyogamy (the fusion of the two different nuclei) does not follow immediately. For a time, the nuclei re- Hypha Spore Mycelium Figure 26-2 Animated to form a mycelium Germination of a spore Kingdom Fungi 557

4 Bud development (a) Yeasts can reproduce asexually by budding. fungi, karyogamy (fusion of the two nuclei) occurs, producing a zygote nucleus that undergoes meiosis. Each haploid spore produced can germinate and develop into a new mycelium by mitosis. In asexual reproduction, spore-producing structures develop and spores are produced by mitosis. Fungi communicate chemically by secreting signaling molecules called pheromones. At least one pheromone has been identified in each fungal phylum. The pheromone binds with a compatible receptor on a different mating type. For example, in the phylum Zygomycetes, a pheromone induces the formation of specialized aerial hyphae. Another pheromone causes the tips of aerial hyphae of opposite mating types to grow toward each other and fuse prior to sexual reproduction. Review What characteristics distinguish fungi from other organisms? How does the body of a yeast differ from that of a mold? Draw a generalized life cycle of a fungus. FUNGAL DIVERSITY Learning Objectives SciMat /Photo Researchers, Inc. (b) SEM (color enhanced) of Saccharomyces cerevisiae. Also known as baker's or brewer's yeast, this fungus can ferment sugar, producing carbon dioxide and alcohol in the process. It is used in brewing beer, in wine production, and in baking bread. Note that several yeast cells are budding, a form of sexual reproduction. Figure 26-3 Yeasts are unicellular fungi main separate within the fungal cytoplasm. Hyphae that contain two genetically distinct, sexually compatible nuclei within each cell are described as dikaryotic (see Fig. 26-1d). This condition is referred to as n n rather than 2n, because there are two separate haploid nuclei. Hyphae that contain only one nucleus per cell are described as monokaryotic. The presence of a dikaryotic stage is an important defining character of the ascomycetes and basidiomycetes. Fungi are a diverse group with many variations in their life cycles. Most fungi (but not all) reproduce both asexually and sexually. Figure 26-4 illustrates a generalized life cycle. When a fungal spore comes into contact with an appropriate food source, the spore germinates and begins to grow. A threadlike hypha emerges from the tiny spore and grows at its tip, branching frequently. In many fungi, cells from individuals of two different mating types fuse (plasmogamy), resulting in a dikaryotic mycelium. In most 4 Give arguments to support the hypothesis that fungi are opisthokonts, more closely related to animals than to plants. 5 Give arguments to support the hypothesis that chytrids may have been the earliest fungal group to evolve from the most recent common ancestor of fungi. 6 List distinguishing characteristics, describe a typical life cycle, and give examples of each of the following fungal groups: chytridiomycetes, zygomycetes, glomeromycetes, ascomycetes, and basidiomycetes. For centuries, biologists classified fungi in the plant kingdom. However, some systematists disagreed with this classification, arguing that there are important structural differences between fungi and plants. Like plants, fungi have cell walls; but unlike plants, fungal cell walls do not contain cellulose. Rather, they contain chitin, a polysaccharide found in insect skeletons. The fungal mode of nutrition is also very different from that of plants Whereas plants are photosynthetic autotrophs, fungi, like animals, are heterotrophs. In 1969, R. H. Whittaker proposed that fungi be assigned to a separate kingdom Fungi. As with systematics for other kingdoms, fungal systematics is a challenging and continuously changing process. For example, slime molds and water molds were formerly classified as fungi but have now been assigned to the protist kingdom (see Chapter 25). Fungi are assigned to the opisthokont clade As discussed in Chapter 25, fungi, animals, and a few protists, including the choanoflagellates, form a monophyletic group, the opisthokonts. Both genetic and structural similarities sup- 558 Chapter 26

5 Kathy Merrifield/Photo Researchers, Inc. Runk /Schoenberger, from Grant Heilman (a) Brown rot of peaches is caused by Monilinia fruticola, an ascomycete. Photographed in Oregon. (b) Corn smut on an ear of sweet corn is caused by Ustilago maydis, a basidiomycete. Figure Fungi that cause plant diseases tacked native American chestnut trees, which had no resistance to the fungus. By the late 1940s, the blight had killed or damaged several billion mature trees almost every North American chestnut tree throughout its entire natural range. Plant biologists are working to save the American chestnut tree from extinction. One approach has been to develop a disease-resistant variety by breeding the American chestnut with the Chinese chestnut, a related species that is resistant to chestnut blight. Other biologists are genetically engineering a virus that infects the chestnut blight fungus and reduces its virulence. Basidiomycetes cause smuts and rusts that attack corn, wheat, oats, and other grains ( Fig b). Some fungal parasites, such as the stem rust of wheat, have complex life cycles that involve two or more different host plants and the production of several kinds of spores. For example, wheat rust must infect a barberry plant at one stage in its life cycle. As a result of this discovery, the eradication of barberry plants in wheat-growing regions has reduced infection with wheat rust. Eradication of the barberry has not eliminated wheat rust, however, because the fungus overwinters on wheat at the southern end of the Grain Belt and forms asexual spores. During the spring, wind blows these spores for hundreds of kilometers, reinfecting northern areas of the United States and Canada. Review Some dictionaries erroneously define a morel as a type of mushroom. Why is this definition incorrect? How are fungi important in modern biology and medicine? What are three important fungal diseases of humans? What are three important fungal diseases of plants? SUMMARY WITH KEY TERMS Learning Objectives 1 Describe the distinguishing characteristics of kingdom Fungi (page 556). Fungi are eukaryotic heterotrophs that secrete digestive enzymes onto their food source and then absorb the predigested food. Fungi are characterized by cell walls that contain chitin. 2 Describe the body plan of a fungus (page 556). A fungus may be a unicellular yeast or a filamentous, multicellular mold. The body of most multicellular fungi consists of long, threadlike filaments called hyphae that branch and form a tangled mass called a mycelium. Learn more about mycelium by clicking on the figure in ThomsonNOW. In most fungi, perforated septa, or cross walls, divide the hyphae into individual cells. In some fungi (zygomycetes and glomeromycetes), the hyphae are coenocytic, that is, they form an elongated, multinuclear cell. 3 Describe the life cycle of a typical fungus, including sexual and asexual reproduction (page 556). Most fungi reproduce both sexually and asexually by means of spores. Spores are produced on aerial hyphae. Kingdom Fungi 577

6 When a fungal spore lands in a suitable spot, the spore germinates. When fungi of two different mating types meet, their hyphae fuse, a process called plasmogamy. The cytoplasm fuses, but the nuclei remain separate. The fungi enter a dikaryotic (n n) stage in which each new cell formed has one nucleus of each type. Karyogamy, fusion of the nuclei, takes place in the hyphal tip and results in a diploid (2n) zygote nucleus. Meiosis takes place, producing four genetically different haploid (n) nuclei. Each nucleus becomes part of a spore. When the spores germinate, they form new mycelia by mitosis. Genetically similar asexual spores can be produced by mitosis. When these spores germinate, they also develop into mycelia. 4 Give arguments to support the hypothesis that fungi are opisthokonts, more closely related to animals than to plants (page 558). Like animals, some fungi have flagellate cells for example, chytrid gametes and spores; the flagellate cells propel themselves with a single posterior flagellum. Like animal cells, fungal cells have platelike cristae in their mitochondria. Based on chemical and structural characters, fungi are classified, along with animals and choanoflagellates, as opisthokonts. 5 Give arguments to support the hypothesis that chytrids may have been the earliest fungal group to evolve from the most recent common ancestor of fungi (page 558). Chytrids, or chytridiomycetes, produce flagellate cells at some stage in their life cycle. No other fungi have flagella. Thus, chytrids probably were the earliest fungi to evolve from a flagellate protist, the common ancestor of all fungi. 6 List distinguishing characteristics, describe a typical life cycle, and give examples of each of the following fungal groups: chytridiomycetes, zygomycetes, glomeromycetes, ascomycetes, and basidiomycetes (page 558). Chytrids reproduce both asexually and sexually. Their gametes and zoospores are flagellate. Allomyces, a common chytrid, spends part of its life as a multicellular haploid thallus and part as a multicellular diploid thallus. The haploid thallus produces two types of flagellate gametes that fuse. Both plasmogamy and karyogamy occur, producing a flagellate zygote. The diploid thallus bears zoosporangia that produce diploid zoospores and resting sporangia in which haploid zoospores form by meiosis. The haploid zoospores form new haploid thalli. Zygomycetes, such as the black bread mold, Rhizopus, form a haploid thallus that produces both asexual spores and sexual spores. Asexual spores germinate and form new thalli. In sexual reproduction, hyphae of two different haploid mating types form gametangia. Plasmogamy occurs as the gametangia fuse. Karyogamy occurs, and a diploid zygote is formed, from which a zygospore develops. Meiosis produces recombinant haploid zygospores. When zygospores germinate, each hypha develops a sporangium at its tip. Spores are released and develop into new hyphae. Microsporidia, now considered zygomycetes, are opportunistic pathogens that penetrate and infect animal cells with their long, threadlike polar tubes. Glomeromycetes (phylum Glomeromycota) are symbionts that form intracellular associations called mycorrhizae with the roots of many plants. Because they extend their hyphae into root cells, glomeromycetes are endomycorrhizal fungi. The most common endomycorrhizae are called arbuscular mycorrhizae because the hyphae inside the root cells form branched, tree-shaped structures known as arbuscules. Glomeromycetes have coenocytic hyphae. They reproduce asexually with large, multinucleate spores called blastospores. Ascomycetes produce asexual spores called conidia; sexual spores called ascospores are produced in asci. The asci line a fruiting body called an ascocarp. Haploid mycelia of opposite mating types produce septate hyphae. Plasmogamy occurs, and nuclei are exchanged. A dikaryotic n n stage occurs in which hyphae form and produce asci and an ascocarp. Karyogamy occurs, followed by meiosis. The recombinant nuclei divide by mitosis, producing eight haploid nuclei that develop into ascospores. When the ascospores germinate, they can form new mycelia. Ascomycetes include yeasts, cup fungi, morels, truffles, and pink, brown, and blue-green molds. Some ascomycetes form mycorrhizae; others form lichens. Basidiomycetes produce sexual spores called basidiospores on the outside of a basidium. Basidia develop on the surface of gills in mushrooms, a type of basidiocarp (a fruiting body). Hyphae in this phylum have septa. Plasmogamy occurs with the fusion of two hyphae of different mating types. A dikaryotic secondary mycelium forms. Then a basidiocarp develops, and basidia form. Karyogamy occurs, producing a diploid zygote nucleus. Meiosis produces four haploid nuclei that become basidiospores. When basidiospores germinate, they form haploid primary mycelia. Basidiomycetes include mushrooms, puffballs, bracket fungi, rusts, and smuts. Explore fungus life cycles by clicking on the figures in ThomsonNOW. 7 Explain the ecological significance of fungi as decomposers (page 570). Most fungi are decomposers that break down organic compounds in dead organisms, leaves, garbage, and wastes into simpler nutrients that can be recycled. 8 Describe the important ecological role of mycorrhizae (page 570). Mycorrhizae are mutualistic relationships between fungi and the roots of plants. The fungus supplies water and nutrient minerals to the plant, and the plant secretes organic compounds needed by the fungus. Glomeromycetes form endomycorrhizal associations with roots. Ascomycetes and basidiomycetes form ectomycorrhizae with tree roots; they do not penetrate the root cells. 9 Characterize the unique nature of a lichen (page 570). A lichen is a symbiotic combination of a fungus and a photoautotroph (an alga or cyanobacterium). The photoautotroph provides the fungus with organic compounds, shelter, water, and minerals. Lichens have three main growth forms: crustose, foliose, and fruticose. Learn more about lichens by clicking on the figure in ThomsonNOW. 10 Summarize some of the ways that fungi impact humans economically (page 574). Mushrooms, morels, and truffles are foods; yeasts are vital in the production of beer, wine, and bread; certain fungi are used to produce cheeses and soy sauce. Fungi are also used to make citric acid and other industrial chemicals. 578 Chapter 26

7 11 Summarize the importance of fungi to biology and medicine; describe how fungi infect plants and humans, identifying at least three fungal plant diseases and three fungal animal diseases (page 574). Modern biologists use the yeast Saccharomyces cerevisiae and other fungi as model organisms for research in molecular biology and genetics. Fungi are also being investigated for the biological control of insects, such as the mosquitoes that transmit malaria. Fungi are used to make many medications, including penicillin and other antibiotics. Fungi are opportunistic pathogens in humans. They cause human diseases, including ringworm, athlete s foot, candidiasis, and histoplasmosis. Some fungi produce mycotoxins, such as aflatoxins, which can cause liver damage and cancer. Fungal hyphae infect plants through stomata. Hyphal branches called haustoria penetrate plant cells and obtain nourishment from the cytoplasm. Fungi cause many important plant diseases, including wheat rust, Dutch elm disease, and chestnut blight. TEST YOUR UNDERSTANDING 1. Fungi produce cell walls containing the polymer. (a) chitin (b) cyclosporine (c) cellulose (d) peptidoglycan (e) lysergic acid 2. Which of the following fungi does not have a mycelium? (a) black bread mold (b) yeast (c) decomposer cup fungus (d) cultivated mushroom (e) Penicillium 3. The condition described as n n is (a) monokaryotic (b) diploid (c) a primary mycelium (d) coenocytic (e) dikaryotic 4. With the exception of chytridiomycetes, fungi are generally disseminated by (a) water currents (b) fragmentation of hyphae (c) soredia (d) airborne spores (e) flagellate zoospores 5. Which statement is not true of the chytrids? (a) they are simple aquatic fungi (b) they produce motile cells with single, posterior flagella (c) they undergo both sexual and asexual reproduction (d) the black bread mold is a representative of this group (e) they were the earliest fungi to evolve 6. Which statement is not true of the zygomycetes? (a) many members of this group form endomycorrhizae with tree roots (b) their sexual spores are called zygospores (c) they undergo both sexual and asexual reproduction (d) plasmogamy and karyogamy occur (e) they have coenocytic hyphae 7. Which statement is not true of the ascomycetes? (a) their sexual spores are produced in asci (b) their asexual spores are produced in basidia (c) some species in this group are yeasts (d) their asexual spores are called conidia (e) some species cause serious plant diseases 8. The ascomycete life cycle typically includes (a) mainly diploid thalli (b) the formation of a thick zygosporangium (c) the production of eight haploid ascospores within an ascus (d) production of microsporidia (e) the production of ascospores, zoospores, and conidia at different stages 9. Which statement is not true of the basidiomycetes? (a) they have a diploid thallus that produces zoospores (b) their sexual spores are called basidiospores (c) they produce a secondary mycelium with n n hyphae (d) mushrooms and bracket fungi are examples of this group (e) this group includes both edible and poisonous species 10. The familiar portion of a mushroom is actually a large fruiting body called a(an) (a) ascocarp (b) basidium (c) basidiocarp (d) gametangium (e) ascus 11. Glomeromycetes (a) reproduce mainly by sexual spores called glomerospores (b) are characterized by unique structures called polar tubes (c) include some species that associate with cyanobacteria to form lichens (d) include many opportunistic pathogens that cause human disease (e) form arbuscular endomycorrhizae with tree roots 12. A symbiotic association between a photoautotroph and a fungus is called a(an) (a) arbuscular endomycorrhiza (b) ectomycorrhiza (c) lichen (d) pathogenic agent (e) lignin decomposing agent 13. Fungi are (a) eukaryotes and opisthokonts (b) prokaryotes and opisthokonts (c) flagellate and dikaryotic (d) prokaryotic pathogens (e) heterotrophs with cellulose cell walls 14. Mutualistic relationships between fungi and the roots of plants are called (a) lichens (b) mycorrhizae (c) pathogenic associations (d) parasitic haustoria (e) mycotoxic symbioses 15. Amanita virosa (a) is the mushroom commonly cultivated for food (b) is the yeast that ferments wine and beer (c) produces the unique flavor of many cheeses (d) is a highly toxic mushroom (e) has been ingested for its hallucinogenic properties 16. Which of the following describes how a fungus infects a plant? (a) it infiltrates leaves with lichens (b) it forms relationships by attaching mycorrhizae to stems (c) it secretes powerful digestive juices onto the leaves (d) it uses haustoria to penetrate stem tissue (e) its hyphae enter leaves through a stoma 17. Mycotoxins (a) are released by most lichens (b) typically cause mild allergic reactions (c) can harm the liver (d) reduce the effects of sick-building syndrome (e) are produced mainly by chytrids 18. Saccharomyces cerevisiae (a) is used by biologists as a model eukaryotic cell (b) was the first prokaryote whose genome was sequenced (c) has an extremely complex genome with more than 6 million genes (d) is a mold used to produce antibiotics (e) is a common chytrid Kingdom Fungi 579

8 CRITICAL THINKING 1. Explain the statement Mushrooms are like the tips of icebergs. If you do not see mushrooms in your lawn, can you conclude that no fungi live there? Why or why not? 2. Biologists have discovered that many mycorrhizal fungi are sensitive to a low ph. What human-caused environmental problem may prove catastrophic for these fungi? How may this problem affect their plant partners? 3. Evolution Link. How are the life cycles of the marine alga Ulva (see Fig ) and Allomyces similar? What hypotheses do these similarities suggest about their evolutionary relationship? 4. Evolution Link. Justify (a) classifying fungi as opisthokonts; (b) classifying microsporidia as fungi; (c) grouping ascomycetes and basidiomycetes as sister clades. Additional questions are available in ThomsonNOW at login 580 Chapter 26

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