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Fungi: Recyclers, Pathogens, 31 Parasites, and Plant Partners

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Fungi: Recyclers, Pathogens, 31 Parasites, and Plant Partners Fungi: Recyclers, Pathogens, 31 Parasites, and Plant Partners About 300 million Africans in 25 countries are suffering because of the invasion of crops by witchweed (Striga), a parasitic flowering plant. This parasite has attacked more than two-thirds of the sorghum, maize, and millet crops in sub-Saharan Africa, doing dam- age estimated at U.S. $7 billion each year. In 1991 a team of Canadian scientists began a search for a solution to the Striga problem. By 1995 they had begun fieldwork in Mali. What was their strategy? They had isolated a strain of a fungus, the mold Fusarium oxysporum, that has two out- standing properties. First, it grows on Striga, wiping out a high percentage of the par- Fungus Trumps Plant The fungus asites. Second, it is not toxic to humans, nor does it attack the crop plants on which Fusarium oxysporum is a potent pathogen Striga is growing. Now farmers apply the fungus to their crops and are rewarded by of witchweed (Striga), a parasitic plant that attacks crops.The fungus spores are shown greatly increased crop yields as Striga is held in check. in blue; the fungal filaments are in tan. Both It may be possible to repeat this story—using a fungus to wipe out a particular colors were added to this electron micro- type of flowering plant—in a very different context. graph. A different strain of F. oxysporum preferentially at- tacks coca plants (the source of cocaine). There is a controversial proposal to use F. oxysporum to wipe out the coca plantations of Andean South America and some countries in other parts of the world. Some other fungi attack people, not plants. Every breath we take contains large numbers of fungal spores. Some of those spores can be dan- gerous, and fungal diseases of humans, some of which are as yet incurable, have become a major global threat. However, other fungi are of immense commercial importance to us. Fungi are essential to plants as well. They interact with roots, greatly enhancing the roots’ ability to take up water and mineral nutrients. Fungi and plants probably in- vaded the land together in the Paleozoic era (see Table 22.1). Earth would be a messy place without the fungi. They are constantly at work in forests, fields, and garbage dumps, breaking down the remains of dead organisms (and even manufactured sub- stances, such as some plastics). For almost a billion years, the ability of fungi to decompose organic substances has been essential for life on Earth, chiefly because by breaking down carbon com- 604 CHAPTER THIRTY-ONE pounds, they return carbon and other elements to the envi- The alternation between gametophyte (n) and sporophyte ronment, where they can be used again by other organisms. (2n) generations that evolved in plants (see Chapter 29) is In this chapter we will examine the general biology of the found in only the most basal group of fungi, the chytrids. The kingdom Fungi, which differs in interesting ways from the derived condition, which is found in the other three fungal other kingdoms. We will also explore the diversity of body clades, involves a unique state in which two haploid nuclei forms, reproductive structures, and life cycles among the four are present in a single cell, discussed later in this chapter. As phyla of fungi, as well as the mutually beneficial associations one might expect, the chytrids, which are aquatic, possess of certain fungi with other organisms. As we begin our study, flagellated gametes (or spores). Flagella have been lost in the recall that the fungi and the animals are descended from a terrestrial fungi. common ancestor—molds and mushrooms are more closely The kingdom Fungi consists of four phyla: Chytridiomy- related to us than they are to the flowers we admired in the cota, Zygomycota, Ascomycota, and Basidiomycota. We dis- last chapter. tinguish the phyla on the basis of their methods and struc- tures for sexual reproduction and, to a lesser extent, by criteria such as the presence or absence of cross-walls sepa- General Biology of the Fungi rating their cell-like compartments. This morphologically The kingdom Fungi encompasses heterotrophic organisms with based phylogeny has proved largely consistent with phylo- absorptive nutrition and with chitin in their cell walls. The fungi genies based on DNA sequencing. The term “fungal system- live by absorptive nutrition: They secrete digestive enzymes atics” has an interesting anagram, “fantastic ugly mess,” but that break down large food molecules in the environment, we’ll see that the situation isn’t all that bad. and then absorb the breakdown products. Many fungi are In the sections that follow, we’ll consider some aspects of saprobes that absorb nutrients from dead matter, others are the general biology of the fungi, including their body struc- parasites that absorb nutrients from living hosts (Figure 31.1), ture and its intimate relationship with their environment, and still others are mutualists that live in intimate association their nutrition, and some special aspects of their unusual sex- with other organisms. ual reproductive cycles. The production of chitin, a polysaccharide, is a synapo- morphy (shared derived trait) for fungi, choanoflagellates, and animals. That is, its presence in fungi is the evidence that Some fungi are unicellular all fungi are more closely related to animals than any fungi Unicellular forms are found in all of the fungal phyla. Uni- are to plants. Chitin is used in the cell walls of fungi, but it is cellular members of the Zygomycota, Ascomycota, and Ba- used in other ways in animals. The use of chitin in cell walls sidiomycota are called yeasts. Yeasts may reproduce by bud- is a synapomorphy for fungi, and it allows us to distinguish ding, by fission, or by sexual means (Figure 31.2). Their between the fungi and the basal eukaryotes (protists) that re- means of reproduction help us to place them in their appro- semble them. Some protists that were formerly confused with priate phyla, as we will see below. fungi include the slime molds (see Figures 28.31 and 28.32) and water molds (oomycetes; see Figure 28.23). The body of a multicellular fungus is composed of hyphae (a) Fungus (b) Fungal fruiting body Most fungi are multicellular. The body of a multicellular fungus is called a mycelium (plural, mycelia). It is composed of rapidly growing individual tubular filaments called 31.1 Parasitic Fungi Attack Other Living Organisms (a) The gray masses on this ear of corn are the parasitic fungus Ustilago maydis,commonly called corn smut. (b) The tropical fungus whose fruit- ing body is growing out of the carcass of this ant has developed from a spore ingested by the ant.The spores of this fungus must be ingested by insects before they will germinate and develop.The grow- ing fungus absorbs organic and inorganic nutrients from the ant’s body, eventually killing it, after which the fruiting body produces a new crop of spores. FUNGI: RECYCLERS, PATHOGENS, PARASITES, AND PLANT PARTNERS 605 Nuclei Cell wall Pore Septum Septa are not complete: Pores allow movement Saccharomyces sp. of organelles and other materials between cell- 31.2 Yeasts Are Unicellular Fungi Unicellular members of the fun- like compartments. gal phyla Zygomycota, Ascomycota, and Basidiomycota are known as yeasts. Many yeasts reproduce by budding—mitosis followed by (a) Coenocytic hypha (b) Septate hypha asymmetrical cell division—as those shown here are doing. 31.3 Most Hyphae Are Incompletely Divided into Separate Cells (a) Coenocytic hyphae have no septa between their nuclei. (b) Even in septate hyphae, the septa do not block the movement of organelles within the hypha. hyphae (singular, hypha). Within hyphae of two clades, in- complete cross-walls called septa (singular, septum) divide the hypha into separate cells. Pores in the septa allow or- ganelles—sometimes even nuclei—to move in a controlled phae may be widely dispersed to forage for nutrients over a way between cells (Figure 31.3). Other hyphae are coenocytic large area, or they may clump together in a cottony mass to and have no septa. exploit a rich nutrient source. Sometimes, when sexual spores Certain modified hyphae, called rhizoids, anchor chytrids are produced, the mycelium becomes reorganized into a fruit- and some other fungi to their substratum (the dead organ- ing (reproductive) structure such as a mushroom. ism or other matter upon which they feed). These rhizoids The way in which a parasitic fungus attacks a plant illus- are not homologous to the rhizoids of plants because they are trates the absorptive role of fungal hyphae (Figure 31.4). The not specialized to absorb nutrients and water. Parasitic fungi hyphae of a fungus invade a leaf through the stomata, through may possess modified hyphae that take up nutrients from wounds, or in some cases, by direct penetration of epidermal their host. cells. Once inside the leaf, the hyphae form a mycelium. Some The total hyphal growth of a mycelium (not the growth hyphae produce haustoria, branching projections that push of an individual hypha) may exceed 1 km per day. The hy- into the living plant cells, absorbing the nutrients within the cells. The haustoria do not break through the plant cell plasma mem- Grass cells branes; they simply press into the cells, with the membrane fitting them like a glove. Fruiting structures may form, either within the plant body or on its surface. Fungal hyphae SporeStoma Hypha Fungal spores Elongating hyphae pass Some hyphae 31.4 A Fungus Attacks a Leaf The white structures in the micro- germinate on the through stomata into penetrate cells surface of the leaf. the interior of the leaf.
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