Land Environment: Plants and Fungi
Chapter
18
Outline 
	18.1	Onto Land 
	•	Plants are adapted to living on land. Their closest present day relatives are the green algae, which are adapted to living in water.284
	•	All plants undergo a cycle termed alternation of generations, and therefore each plant exists in two forms.286
	18.2	Diversity of Plants 
	•	Nonvascular plants, such as mosses, lack well-developed conducting tissues.287
	•	The life cycle of a fern demonstrates the reproductive strategies of most seedless vascular plants.289
	•	The gymnosperms are seed plants well represented by conifers, cone-bearing plants.291
	•	The reproductive structure of angiosperms, which are also seed plants, is the flower.
292–293
	•	Much of the diversity among flowers comes from specialization for certain pollinators.294
	18.3	The Fungi
	•	Fungi are multicellular eukaryotes different in their biology from both plants and animals.296 
	•	Fungi are saprotrophs, and like the decomposing bacteria, they keep ecological cycles functioning in the biosphere.298
	•	Fungi enter significant mutualistic relationships.299
	•	Fungi are also important for their commercial services and as a source of food.300
Have you thanked a green plant today? Without plants, we and our civilized life could not exist. Plants provide us with our food, either directly or indirectly, and with the oxygen we breathe. Oxygen also rises into the stratosphere and becomes the ozone shield that absorbs ultraviolet rays and makes life on land possible. 
The bulk of the human diet comes from just 12 plants. Wheat is associated with Europe, corn with the Americas, and rice with the Far East. Other significant food crops include white and sweet potatoes, cassava, soybeans, common beans, sugarcane, sugar beets, coconuts, and bananas. 
Eons ago, the bodies of plants became the coal we now burn to produce much of our electricity. The wood of trees is also commonly used as fuel. Then, too, fermentation of plant materials produces alcohol, which can be used directly to fuel automobiles or as a gasoline additive. 
Much of our clothing comes from plants. People still like the feel of cotton next to their skin, while linen from flax makes a stronger cloth. Cotton is used in all sorts of other cloth products, such as towels, sheets, and upholstered furniture. Even the manufacture of rayon depends on cellulose from plant cell walls.
Thousands of other household products come from plants. Wood is used to make furniture as well as the houses that contain the furniture. Jute is used for rope, carpet, insulation, and burlap bags. Coconut oil is found in soaps, shampoos, and suntan lotions. Even toothpastes contain such plant flavorings as peppermint and spearmint. Suffice it to say, it would be impossible to mention all the products from plants that we depend on for our daily needs. 
18.1		Onto Land
Plants are a diverse group that range in size and structure from the smallest aquatic duckweed to the largest giant sequoia. Plants, unlike algae, are adapted for living on land. Still, several types of evidence support the conclusion that plants are related, through evolution, to green algae. Both green algae and plants (1) contain chlorophylls a and b and various accessory pigments, (2) store excess carbohydrates as starch, and (3) have cellulose in their cell walls. Additionally, in recent years biochemists have compared the sequences of DNA bases coding for ribosomal RNA between organisms. The results suggest that plants are most closely related to a group of green algae encrusted with calcium carbonate and, therefore, known as stoneworts, perhaps those in the genus Chara. The split between green algae and plants may have occurred about 500 million years ago (mya).
Today, plants are classified into four groups (Fig. 18.1). It is possible to associate each group with one of four evolutionary events, all of them adaptations to a land existence. First, all the groups of plants protect and nourish a multicellular embryo within the body of the plant (Fig. 18.1a). Since green algae do not protect the embryo as all plants do, this may be the first evolved feature that separates plants from green algae.
The next event was most likely the evolution of vascular tissue. Vascular tissue is essential for the transport of water and solutes from the roots to the leaves of terrestrial plants (Fig. 18.1b). The fossil record indicates that vascular plants evolved about 420 mya.
The third event was the evolution of seeds. Two groups of plants produce seeds (Fig. 18.1c,d). A seed is composed of an embryo and stored organic nutrients within a protective coat (see Fig. 18.10). When you plant a seed, a plant of the next generation emerges; in other words, seeds disperse offspring. Seeds are highly resistant structures well suited to protect a plant embryo from drought, and to some extent from predators, until conditions are favorable for germination. The first seed plants appear in the fossil record about 360 mya.
The fourth event was the evolution of the flower, a reproductive structure (Fig. 18.1d). Flowers attract pollinators such as insects, and they also give rise to fruits that cover seeds. Plants with flowers may have evolved about 160 mya. 
Figure 18.2 traces the evolutionary history of plants and will serve as a backdrop as we discuss the four major groups of plants in the pages that follow. 
Alternation of Generations 
The life cycle of plants is quite different from that of animals. All plants undergo an alternation of generations, which means that each type of plant exists in two forms (Fig. 18.3). One form is the diploid sporophyte, and the other is the haploid gametophyte. The sporophyte (2n) is named for its production of haploid spores by meiosis. A spore is a reproductive cell that develops into a new organism without the need to fuse with another reproductive cell. In the plant life cycle, a spore undergoes mitosis and becomes a gametophyte. The gametophyte (n) is named for its production of gametes. A sperm and egg fuse, forming a zygote that undergoes mitosis and becomes the sporophyte.
You’ll want to make two observations in Figure 18.3. First note that, in plants, meiosis produces haploid spores. This is consistent with the realization that the sporophyte is the diploid generation and spores are -haploid. Second, note that mitosis occurs as a spore becomes a gametophyte, and it also occurs as a zygote becomes a sporophyte. Mitosis must happen in both places in order to have two generations.
The Dominant Generation
When you bring a plant to mind, you’re probably thinking of the dominant generation—the generation that is larger, lasts longer, and is the generation we recognize as the plant. In nonvascular plants, the gametophyte is the dominant generation, and in the other groups of plants, the sporophyte is the dominant generation. In the history of plants, only the sporophyte evolves vascular tissue; therefore, the trend toward sporophyte dominance is an adaptation to life on land. As the sporophyte gains in -dominance, the gametophyte becomes microscopic. It also becomes dependent upon the sporophyte, the generation best adapted to a dry land environment. 
Note the appearance of the generations in Figure 18.4. In mosses (bryophytes), the gametophyte is much larger than the sporophyte. In ferns, the gametophyte is a small, indepen-dent, heart-shaped structure. In contrast, the -female gametophyte of cone-bearing plants (gymnosperm) and flowering plants (angiosperm) is microscopic—it is retained within the body of the sporophyte plant. This protects the female gametophyte from drying out. Also, -the male -gametophyte of seed plants lies within a pollen grain. Pollen grains have strong protective walls and are transported by wind, -insects, or birds to reach the egg. In the life cycle of seed plants, the spores, the gametes, and the zygote are protected from drying out in the land environment. 
18.2		Diversity of Plants
In the evolution of plants, the nonvascular plants evolved before the vascular plants. 
Nonvascular Plants
The nonvascular plants, often called the bryophytes, do not have true roots, stems, and leaves—which, by definition, must contain vascular tissue. In bryophytes, the gametophyte is the dominant generation. 
The most familiar bryophytes are the liverworts and mosses, which are low-lying plants. In bryophytes, the gametophyte consists of leafy shoots, which produce the gametes (Fig. 18.5). Flagellated sperm swim in a film of water to reach an egg. After a sperm fertilizes an egg, the resulting zygote becomes an embryo that develops into a sporophyte. The sporophyte is attached to, and derives its nourishment from, the photosynthetic gametophyte. The sporophyte produces windblown spores, an adaptation to life on land.
The common name of several plants implies that they are mosses when they are not. Irish moss is an edible red alga that grows in leathery tufts along northern seacoasts. Reindeer moss, a lichen, is the dietary mainstay of reindeer and caribou in northern lands. Club mosses, discussed later in this chapter, are vascular plants, and Spanish moss, which hangs in grayish clusters from trees in the southeastern United States, is a flowering plant of the pineapple family.
Adaptations and Uses of Bryophytes
The lack of vascular tissue and the presence of swimming sperm largely account for the short height of bryophytes such as mosses. Still, mosses can be found from the Antarctic through the tropics to parts of the Arctic. Although most mosses prefer damp, shaded locations in the temperate zone, some survive in deserts, and others -inhabit bogs and streams. In forests, they frequently form a mat that covers the ground and rotting logs. In dry environments, they may become shriveled, turn brown, and look completely dead. As soon as it rains, the plant -becomes green and resumes metabolic activity. Mosses are even better than flowering plants at living on stone walls, on fences, and in shady cracks of hot, exposed rocks. When bryophytes colonize bare rock, the rock degrades to soil that can be used for their own growth and the growth of other organisms. 
In areas such as bogs, where the ground is wet and acidic, dead mosses, especially Sphagnum, do not decay. The accumulated moss, called peat or bog moss, can be used as fuel. Peat moss is also commercially important in another way. Because it has special nonliving cells that can absorb moisture, peat moss is often used in gardens to improve the water-holding capacity of the soil. The so-called copper mosses live only in the vicinity of copper, and can serve as an indicator of copper -deposits.
Vascular Plants
All the other plants we will study are vascular plants. The vascular plants usually have true roots, stems, and leaves. The roots absorb water from the soil, and the stem conducts water to the leaves. The leaves are fully covered by a waxy cuticle except where it is interrupted by stomata, little pores for gas exchange, the opening and closing of which can be regulated to control water loss. 
Vascular tissue consists of xylem, which conducts water and minerals up from the roots, and phloem, which conducts organic -nutrients from one part of a plant to another (Fig. 18.6). The walls of conducting cells in xylem are strengthened by lignin, an -organic compound that makes them stronger, more waterproof, and resistant to attack by parasites and predators. Only because of strong cell walls can plants reach great heights. 
Seedless Vascular Plants
The seedless vascular plants include club mosses, horsetails, and ferns. Today club mosses are represented in temperate forests by ground pines, so called  because of their limited height and superficial resemblance to a pine tree. Horsetails are taller and have a stem that is interrupted by whorls of slender, green branches. Horsetails live in moist habitats about the globe. This category of plants has vascular tissue but does not produce seeds. Instead, windblown spores disperse the plant to new locations. 
Before the seed plants evolved, the seedless vascular plants were as tall as today’s trees, and they dominated the swamp forests of the Carboniferous period (Fig. 18.7). A large number of these plants died but did not decompose completely. Instead, they were compressed to form the coal that we still mine and burn today. (Oil has a similar origin, but it most likely formed in marine sedimentary rocks and included animal remains.)
Ferns
Ferns are a widespread group of plants that are well known for their attractiveness. The large and conspicuous leaves of ferns, called fronds, are commonly divided into leaflets. The royal fern has fronds that stand about 1.8 meters tall; those of the hart’s tongue fern are straplike and leathery; and those of the maidenhair fern are broad, with subdivided leaflets (Fig. 18.8). 
Adaptations and Uses of FernsFerns are most often found in a moist environment because the water-dependent gametophyte, which lacks vascular tissue, is separate from the sporophyte. -Also, flagellated sperm require an outside source of moisture in which to swim to the eggs (Fig. 18.9). Once established, some ferns, such as the bracken fern, can spread into drier areas because their rhizomes, which grow horizontally in the soil, produce new plants.
At first it may seem that ferns do not have much economic value, but they are frequently used by florists in decorative bouquets and as ornamental plants in the home and garden. Wood from tropical tree ferns is often used as a building material because it resists decay, particularly by termites. Ferns, especially the ostrich fern, are used as food, and in the northeastern United States, many restaurants feature fiddleheads (that season’s first growth) as a special treat. Ferns also have medicinal value; many Native Americans use ferns as an astringent during childbirth to stop bleeding, and the maidenhair fern is the source of a cold medicine.
Seed Plants
Seed plants are the most plentiful plants in the biosphere today. Most trees are seed plants, and so are almost all your garden plants. The major parts of a seed are shown in Figure 18.10. The seed coat and stored food protect the embryo and allow it to survive harsh conditions during a period of dormancy (arrested state), until environmental conditions become favorable for growth. Seeds can even remain dormant for hundreds of years. When a seed germinates, the stored food is a source of nutrients for the growing seedling. The survival value of seeds largely accounts for the dominance of seed plants today.
Seed plants have two types of spores and produce two kinds of gametophytes—male and female. The gametophytes are microscopic and consist of just a few cells. Pollen grains are drought resistant male gametophytes. Pollination occurs when a pollen grain is brought to the vicinity of the female gametophyte by wind or a pollinator. Later, sperm move toward the female gametophyte through a growing pollen tube, and fertilization occurs. Note that no external water is needed to accomplish fertilization. The whole male gametophyte, rather than just the sperm as in seedless plants, moves to the female gametophyte. 
A female gametophyte develops within an ovule, which eventually becomes a seed (Fig. 18.11). The embryo in a seed will be the sporophyte plant. In gymnosperms, the ovules are not completely enclosed by sporophyte tissue at the time of pollination. In angiosperms, the ovules are completely enclosed within diploid sporophyte tissue (an ovary), which becomes a fruit. We shall see that the flower has many advantages, one of which is the production of fruit. 
Gymnosperms
In gymnosperms, ovules and seeds are exposed on the surface of a cone scale (modified leaf). The term gymnosperm means “naked seed.” Ancient gymnosperms, including cycads, were present in the swamp forests of the Carboniferous period (Fig. 18.12). Of these, the conifers have become a dominant plant group today. 
Conifers 
Pines, spruces, firs, cedars, hemlocks, redwoods, cypresses, and others are all conifers (Fig. 18.13). The name conifer signifies plants that bear cones, but other types of gymnosperms are also cone-bearing. The coastal redwood, a conifer native to northwestern California and southwestern Oregon, is the tallest living vascular plant; it may attain nearly 100 meters in height. Another conifer, the bristlecone pine of the White Mountains of California, is the oldest living tree—one specimen is 4,900 years of age.
Adaptations and Uses of Pine TreesVast areas of northern temperate regions are covered in evergreen coniferous forests. The tough, needlelike leaves of pines conserve water because they have a thick cuticle and recessed stomata. This type of leaf helps them live in areas where frozen topsoil makes it difficult for the roots to obtain plentiful water. 
A substance called resin protects leaves and other parts of the tree from insect and fungal attacks. The resin of certain pines is harvested; the liquid portion, called turpentine, is a paint thinner, while the solid portion is used on stringed instruments. The wood of pines is used extensively in construction, and vast forests of pines are planted for this purpose. The wood consists primarily of xylem tissue that lacks some of the more rigid cell types found in flowering trees. Therefore, it is considered a “soft” rather than a “hard” wood. 
Angiosperms
The angiosperms (meaning covered seeds) are an exceptionally large and successful group of plants, with 240,000 known species—six times the number of species of all other plant groups combined. Angiosperms, also called the flowering plants, live in all sorts of habitats, from fresh -water to desert, and from the frigid north to the torrid tropics. They range in size from the tiny, almost microscopic -duckweed to Eucalyptus trees over 100 meters tall. Most plants in your garden produce flowers, and therefore are angiosperms. In northern climates, the trees that lose their leaves are flowering plants. In subtropical and tropical climates, flowering trees as well as gymnosperms tend to keep their leaves all year. 
Although the first fossils of angiosperms are no older than about 135 million years, the angiosperms probably arose much earlier. Indirect evidence suggests the possible ancestors of angiosperms may have originated as long ago as 160 mya. To help solve the mystery of their origin, botanists have turned to DNA comparisons to find a living plant that is most closely related to the first angiosperms. Their data point to Amborella trichopoda as having the oldest lineage among today’s angiosperms (Fig. 18.14). This small shrub, which has small, cream-colored flowers, lives only on the island of New Caledonia in the South Pacific. 
The Flower
Most flowers have certain parts in common despite their dissimilar appearances. The flower parts, called sepals, petals, stamens, and carpels, occur in whorls (circles) (Fig. 18.15). The sepals, collectively called the calyx, protect the flower bud before it opens. The sepals may drop off or may be colored like the petals. Usually, however, sepals are green and remain -in place. The petals, collectively called the corolla, are quite diverse in size, shape, and color. The petals often attract a particular pollinator. Each stamen consists of a stalk called a filament and an anther, where pollen is produced in pollen sacs. In most flowers, the anther is positioned where the pollen can be carried away by wind or a pollinator. One or more carpels is at the center of a flower. A carpel has three major regions: ovary, style, and stigma. The swollen base is the ovary, which contains from one to hundreds of ovules. The style elevates the stigma, which is sticky or otherwise adapted for the reception of pollen grains. Glands located in the region of the ovary produce nectar, a nutrient that is gathered by pollinators as they go from flower to flower. 
Flowering Plant Life Cycle
In angiosperms, the flower produces seeds enclosed by fruit. The ovary of a carpel contains several ovules, and each of these eventually holds an egg-bearing female gametophyte called the embryo sac. During pollination, a pollen grain is transported by various means from the anther of a stamen to the stigma of a carpel, where it germinates. The pollen tube carries the two sperm into a small opening of an ovule. During double fertilization, one sperm unites with an egg nucleus forming a diploid zygote, and the other sperm unites with two other nuclei forming a triploid (3n) endosperm (Fig. 18.16). In angiosperms, the endosperm is the stored food. 
Ultimately, the ovule becomes a seed that contains a sporophyte embryo. In some seeds, the endosperm is absorbed by the seed leaves, called cotyledons, whereas in other seeds, endosperm is digested as the seed germinates. When you open a peanut, the two halves are the cotyledons. If you look closely, you will see the embryo between the cotyledons. A fruit is derived from an ovary and possibly accessory parts of the flower. Some fruits (e.g., apple) provide a fleshy covering for their seeds, and other fruits provide a dry covering (e.g., pea pod, shell of peanut). 
Adaptations and Uses of Angiosperms
Successful completion of sexual reproduction in angiosperms requires the effective dispersal of pollen and then seeds. Adaptations have resulted in various means of dispersal of pollen and seeds. Wind-pollinated flowers are usually not showy, whereas insect-pollinated flowers and bird-pollinated flowers are often colorful (Fig. 18.17a–c). Night-blooming flowers attract nocturnal mammals or insects; these flowers are usually aromatic and white or cream-colored (Fig. 18.17d). Although some flowers disperse their pollen by wind, many are adapted to attract specific pollinators, such as bees, wasps, flies, butterflies, moths, and even bats, which carry only particular pollen from flower to flower. For example, bee--pollinated flowers are usually blue or yellow and have -ultraviolet shadings that lead the pollinator to the location of nectar at the base of the flower. In turn, the mouthparts of bees are fused into a long tube that is able to obtain nectar from this location. Today, there are some 240,000 species of flowering plants and over 900,000 species of insects. This diversity suggests that the success of angiosperms has contributed to the success of insects and vice versa. 
The fruits of flowers protect and aid in the dispersal of seeds. Dispersal occurs when seeds are transported by wind, gravity, water, and animals to another location. Fleshy fruits may be eaten by animals, which transport the seeds to a new location and then deposit them when they defecate. Because animals live in particular habitats and/or have particular migration patterns, they are apt to deliver the fruit-enclosed seeds to a suitable location for seed germination (initiation of growth) and development of the plant.
Economic Benefits of Plants
One of the primary economic benefits of plants is the use of their fruits as food. Botanists use the term “fruit” in a much broader way than do laypeople. Among the foods mentioned in the introduction, you would have no trouble recognizing a banana as a fruit. A coconut is also a fruit, as are the grains (corn, wheat, rice) and the pods that contain beans or peas (Fig. 18.18). Cotton is derived from the cotton boll, a fruit containing seeds with seed hairs that become textile fibers used to make cloth. 
Other economic benefits of plants include foods and commercial products made from roots, stems, and leaves. Cassava and sweet potatoes are edible roots, while white potatoes are the tubers of underground stems. Most furniture and paper are made from the wood of a tree trunk (Fig. 18.19). Also, the many chemicals produced by plants make up 50% of all pharmaceuticals and various other types of products we can use. The cancer drug taxol originally came from the bark of the Pacific yew tree. Today, plants are even bioengineered to produce certain substances of interest. 
Indirectly, the economic benefits of plants are often dependent on pollinators. Only if pollination occurs can these plants produce a fruit and propagate themselves. In recent years, the populations of bees and other pollinators have been declining worldwide, principally due to a parasitic mite but also due to the widespread use of pesticides. Consequently, some plants are endangered because they have lost their normal pollinators. Because of our dependence on flowering plants, we should protect pollinators! 
Ecological Benefits of Plants
The ecological benefits of flowering plants are so important that we could not exist without them. Plants produce food for themselves and directly or indirectly for all other organisms in the biosphere. The oxygen they produce is also used by themselves and all organisms that carry out cellular respiration. 
Forests are an important part of the water cycle and the carbon cycle. The roots of trees in particular hold soil in place and absorb water that returns to the atmosphere. Without these functions of trees and other plants, rainwater runs off and contributes to flooding. The absorption of carbon dioxide by plants lessens the amount in the atmosphere. CO2 in the atmosphere contributes to global warming because it and other gases trap heat near the Earth. The burning of tropical rain forests is double trouble for global warming because it adds CO2 to the atmosphere and removes trees that otherwise would absorb CO2. Some plants can also be used to clean up toxic messes. For example, poplar, mustard, and mulberry species take up lead, uranium, and other pollutants from the soil. 
In addition to all the other uses of plants, we should not forget their aesthetic value. Almost everyone prefers to vacation in a natural setting and enjoys the sight of trees and flowers in the yard and in the house.
18.3		The Fungi
Asked whether fungi are animal or vegetable, most would choose vegetable. But this would be wrong because fungi do not have chloroplasts, and they can't photosynthesize. Then, too, fungi are not animals, even though they are chemoheterotrophs like animals. Animals ingest their food, but fungi release digestive enzymes into their immediate environment and then absorb the products of digestion. Also, animals are motile, but fungi are nonmotile and do not have flagella at any stage in their life cycle. The fungal life cycle differs from that of both animals and plants because fungi produce windblown spores during both an asexual and a sexual life cycle.
Table 18.1 contrasts fungi with plants and animals. The many unique features of fungi indicate that although fungi are multicellular eukaryotes (except for the unicellular yeasts), they are not closely related to any other group of organisms. DNA sequence data suggest fungi are distantly related to animals rather than plants.
General Biology of a Fungus
Representative fungi are shown in Figure 18.20. All parts of a fungus are composed of hyphae (sing., hypha), which are thin filaments of cells. The hyphae are packed closely together to form a complex structure, such as a mushroom. However, the main body of a fungus is not the mushroom or the puffball or the morel; these are just temporary reproductive structures. Mold shows best that the main body of a fungus is a mass of hyphae called a mycelium (Fig. 18.21). The mycelium penetrates the soil, the wood, or the bread in which the fungus is growing. Mycelia in the soil can become large enough to cover acres, making them the largest creatures on Earth and earning them the label “the humongous fungus among us.”
Fungal cells typically have thick cell walls, but unlike plants, the fungal cell walls do not contain cellulose. They are made of another polysaccharide in which the glucose monomers contain amino groups (amino sugars) and form a polymer called chitin. This polymer is also the major structural component of the exoskeleton of insects and arthropods, such as lobsters and crabs. The walls dividing the cells of a hypha have pores that allow the cytoplasm to pass from one cell to the other along the length of the hypha. The hyphae give the mycelium quite a large surface area, which facilitates the ability of the mycelium to absorb nutrients. Hyphae move toward a food source by growing at their tips, and the hyphae of a mycelium absorb and then pass nutrients on to the growing tips. 
Black Bread Mold
Multicellular organisms are characterized by specialized cells. Black bread mold demonstrates that the hyphae of a fungus may be specialized for various purposes (Fig. 18.22). In this fungus, horizontal hyphae exist on the surface of the bread; other hyphae grow into the bread, anchoring the mycelium and carrying out digestion; and some form stalks that bear sporangia. A sporangium is a structure that produces spores. 
The mycelia of two different mating types are featured in the center and bottom of Figure 18.22. During asexual reproduction, each mycelium produces sporangia, where spore formation occurs. Spores are resistant to environmental damage, and they are often made in large numbers. Fungal spores are windblown, a distinct advantage for a nonmotile organism living on land. The spores germinate into new mycelia without going through any developmental stages, another feature that distinguishes fungi from animals. 
Sexual reproduction in fungi involves the conjugation of hyphae from different mating types (usually designated 1 and 2). In black bread mold, the tips of 1 and 2 hyphae join, the nuclei fuse, and a thick-walled zygospore results. The zygospore undergoes a period of dormancy before it germinates, producing sporangia. Meiosis occurs within the sporangia, producing spores of both mating types. The spores, which are dispersed by air currents, give rise to new mycelia. In fungi, only the zygote is diploid, and all other stages of the life cycle are haploid, a distinct difference from animals.
Mushroom
When you eat a mushroom, you are eating a fruiting body, whose function is to produce spores. In mushrooms, when the tips of 1 and 2 hyphae fuse, the haploid nuclei do not fuse immediately. Instead, so-called dikaryotic (two nuclei) hyphae form a mushroom consisting of a stalk and a cap. Club-shaped structures called basidia (sing., basidium) project from the gills located on the underside of the cap (Fig. 18.23a). Fusion of nuclei inside these structures is followed by meiosis and production of windblown spores. Each cap of a mushroom produces tens of thousands of spores.
In the soil, a mycelium is absorbing nutrients and may form a ring. Therefore, when the weather turns warm and rain is plentiful, the mycelium may put forth a circle of mushrooms on your lawn. At one time, people believed this ring indicated where fairies had joined hands and danced the night before (Fig. 18.23b).
Ecological Benefits of Fungi
Most fungi are saprotrophs that decompose the remains of plants, animals, and microbes in the soil. Fungal enzymes can degrade cellulose and even lignin in the woody parts of plants. That is why fungi so often grow on dead trees. This also means that fungi can be used to remove excess lignin from paper pulp. Ordinarily, lignin is difficult to extract from pulp and ends up being a pollutant once it is removed.
Along with the bacteria, which are also decomposers, fungi play an indispensable role in the environment by returning inorganic nutrients to photosynthesizers. Many people take advantage of the activities of bacteria and fungi by composting their food scraps or yard waste. When a gardener makes a compost pile and provides good conditions for decomposition to occur, the result is a dark, crumbly material that makes excellent fertilizer. And while the material may smell bad as decomposition is occurring, the finished compost looks and smells like rich, moist earth.
Some fungi eat animals that they encounter as they feed on their usual meals of dead organic remains. For example, the oyster fungus excretes a substance that anesthetizes any nematodes (roundworms) feeding on it (Fig. 18.24). After the worms become inactive, the fungal hyphae penetrate and digest their bodies, absorbing the nutrients. Other fungi snare, trap, or fire projectiles into nematodes and other small animals before digesting them. The animals serve as a source of nitrogen for the fungus.
Mutualistic Relationships
In a mutualistic relationship, two different species live together and help each other out. Lichens are a mutualistic association between a particular fungus and cyanobacteria or green algae (Fig. 18.25). The fungal partner is efficient at acquiring nutrients and moisture, and therefore lichens can survive in poor soils, as well as on rocks with no soil. The organic acids given off by fungi release from rocks the minerals that can be used by the photosynthetic partner. Lichens are ecologically important because they produce organic matter and create new soil, allowing plants to invade the area.
Lichens occur in three varieties: compact crustose lichens, often seen on bare rocks or tree bark; shrublike fruticose lichens; and leaflike foliose lichens. Regardless, the body of a lichen has three layers. The fungal hyphae form a thin, tough upper layer and a loosely packed lower layer. These layers shield the photosynthetic cells in the middle layer. Specialized fungal hyphae that penetrate or envelop the photosynthetic cells transfer organic nutrients to the rest of the mycelium. The fungus not only provides minerals and water to the photosynthesizer, but also offers protection from predation and desiccation. Lichens can reproduce asexually by releasing fragments that contain hyphae and an algal cell. At first, the relationship between the fungi and algae was likely a parasite-and-host interaction. Over evolutionary time, the relationship apparently became more mutually beneficial, although how to test this hypothesis is a matter of debate at the present time.
Mycorrhizal fungi form mutualistic relationships with the roots of most plants, helping them grow more successfully in dry or poor soils, particularly those deficient in inorganic nutrients (Fig. 18.26). The fungal hyphae greatly increase the surface area from which the plant can absorb water and nutrients. It has been found beneficial to encourage the growth of mycorrhizal fungi when restoring lands damaged by strip mining or chemical pollution.
Mycorrhizal fungi may live on the outside of roots, enter between root cells, or penetrate root cells. The fungus and plant cells exchange nutrients, with the fungus bringing water and minerals to the plant and the plant providing organic carbon to the fungus. Early plant fossils indicate that the relationship between fungi and plant roots is an ancient one, and therefore it may have helped plants adapt to life on dry land. The general public is not familiar with mycorrhizal fungi, but a few people relish truffles, a mycorrhizal fungus that grows in oak and beech forests. Truffles are considered a gourmet delicacy.
Economic Benefits of Fungi
Fungi help us produce medicines and many types of foods (Fig. 18.27). The mold Penicillium was the original source of penicillin, a breakthrough antibiotic that lead to an important class of cillin antibiotics. Cillin antibiotics have saved millions of lives. 
Yeast fermentation is utilized to make bread, beer, wine, and distilled spirits. During fermentation, yeast produce the carbon dioxide that causes dough to rise when gas pockets are preserved as the bread bakes. Ethanol, also a product of yeast fermentation, is desired for the making of alcoholic beverages. Other types of fungal fermentations contribute to the manufacture of various cheeses and soy sauce from soybeans. Another commercial application of interest is the use of fungi to soften the centers of certain candies.
Fungi as Food
In the United States, the consumption of mushrooms has been steadily increasing. In 2001, total consumption of all mushrooms totalled 1.13 billion pounds—21% greater than in 1991. In addition to adding taste and texture to soups, salads, and omelets, and being used in stir-fry and on salads, mushrooms are an excellent low-calorie meat substitute with great nutritional value and lots of vitamins. Although there are thousands of mushroom varieties in the world, the white button mushroom, Agaricus bisporus, dominates the U.S. market. However, in recent years, sales of brown-colored variants have surged in popularity and have been one of the fastest-growing segments of the mushroom industry. Portabella is a marketing name used by the mushroom industry for the more flavorful brown strains of A. bisporus. The mushroom is brown because it is allowed to open, exposing the mature gills with their brown spores; crimini is the same brown strain, but it is not allowed to open before it is harvested. Non-Agaricus varieties, especially shiitake and oyster, have slowly gained in popularity over the past decade. Shiitake is touted for lowering cholesterol levels and having antitumor and antiviral properties.
Fungi as Disease-Causing Organisms
Fungi cause diseases in both plants and humans.
Fungi and Plant Diseases 
Fungal pathogens, which usually gain access to plants by way of the stomata or a wound, are a major concern for farmers. Serious crop losses occur each year due to fungal disease. As much as a third of the world’s rice crop is destroyed each year by rice blast disease. Corn smut is a major problem in the midwestern United States. Various rusts attack grains, and leaf curl is a disease of fruit trees (Fig. 18.28). 
The life cycle of rusts may be particularly complex, since it requires two different host species to complete the cycle. Black stem rust of wheat uses barberry bushes as an alternate host. Eradication of barberry bushes in areas where wheat is grown helps control this rust. Fungicides are regularly applied to crops to limit the negative effects of fungal pathogens. Wheat rust can also be controlled by producing new and resistant strains of wheat.
Fungi and Human Diseases
As is well known, certain mushrooms are poisonous. The ergot fungus that grows on grain can result in ergotism when a person eats contaminated bread. Ergotism is characterized by hysteria, convulsions, and sometimes death. 
Mycoses are diseases caused by fungi. Mycoses have three possible levels of invasion: Cutaneous mycoses only affect the epidermis; subcutaneous mycoses affect deeper skin layers; and systemic mycoses spread their effects throughout the body by traveling in the bloodstream. Fungal diseases that can be contracted from the environment include ringworm from soil, rose gardener’s disease from thorns, Chicago disease from old buildings, and basketweaver’s disease from grass cuttings. Opportunistic fungal infections now seen in AIDS patients stem from fungi that are always present in the body but take the opportunity to cause disease when the immune system becomes weakened. 
Candida albicans causes the widest variety of fungal infections. Disease occurs when antibacterial treatments kill off the microflora community, allowing Candida to proliferate. Vaginal Candida infections are commonly called “yeast infections” in women. Oral thrush is a Candida infection of the mouth common in newborns and AIDS patients (Fig. 18.29a). In individuals with inadequate immune systems, Candida can move throughout the body, causing a systemic infection that can damage the heart, brain, and other organs.
Ringworm is a group of related diseases caused, for the most part, by fungi in the genus Tinea. Ringworm is a cutaneous infection that does not penetrate the skin. The fungal colony grows outward, forming a ring of inflammation. The center of the lesion begins to heal, giving the lesion its characteristic appearance, a red ring surrounding an area of healed skin (Fig. 18.29b). Athlete’s foot is a form of tinea that affects the foot, mainly causing itching and peeling of the skin between the toes (Fig. 18.29c).
The vast majority of people living in the midwestern United States have been infected with Histoplasma capsulatum. This common soil fungus, often associated with bird droppings, leads in most cases to a mild “fungal flu.” However, about 3,000 of these cases develop into a severe disease. About 50 persons die each year from histoplasmosis when the fungus grows within cells of the immune system. Lesions are formed in the lungs, leaving calcifications that are visible in X-ray images and resemble those of tuberculosis.
Because fungi are more closely related to animals than to bacteria, it is hard to design an antibiotic against fungi that does not also harm humans. Thus, researchers exploit any biochemical differences they can discover between humans and fungi.
The Chapter in Review
Summary
18.1		Onto Land
Plants probably evolved from a multicellular, freshwater green alga about 500 mya. Whereas algae are adapted to life in the water, plants are adapted to living on land. During the evolution of plants, four significant events are associated with adaptation to a land existence:
Alternation of Generations
All plants have an alternation of generations life cycle, in which each type of plant exists in two forms, the sporophyte (2n) and the gametophyte (n): 
18.2		Diversity of Plants
Nonvascular Plants
Bryophytes, represented by the mosses, are plants with the following characteristics:
	•	There is no well-developed vascular tissue.
	•	The gametophyte is dominant, and flagellated sperm swim in external moisture to the egg.
	•	The sporophyte is dependent on the gametophyte.
	•	Windblown spores disperse the gametophyte.
Vascular Plants
In vascular plants, the dominant sporophyte has two kinds of well-defined conducting tissues. Xylem is specialized to conduct water and dissolved minerals, and phloem is specialized to conduct organic nutrients and hormones.
Seedless Vascular PlantsThe seedless vascular plants grew to enormous sizes during the Carboniferous period. Today, they include club mosses (e.g., ground pines), horsetails, and ferns. 
Ferns best represent seedless vascular plants. They have the following characteristics: 
	•	The sporophyte generation is dominant and produces windblown spores.
	•	The gametophyte is separate and independent. Flagellated sperm swim in external moisture to the egg.
Seed PlantsSeed plants have male and female gametophytes. Gametophytes are reduced in size. The female gametophyte is retained within an ovule, and the male gametophyte is the mature pollen grain. The ovule becomes the seed, which contains a sporophyte embryo, food, and a seed coat. 
	•	Gymnosperms are cone-bearing plants, represented by the pine tree. They have “naked seeds” because the seeds are not enclosed by fruit, as are those of flowering plants.
	•	Angiosperms are the flowering plants. Angiosperm reproductive organs are in the flower. Pollen is produced in pollen sacs inside an anther. Pollen is transported by wind, or from flower to flower by birds, insects, or bats. Ovules in the ovary become seeds, and the ovary becomes the fruit. Thus, angiosperms have “covered seeds.” 
18.3		The Fungi
Fungi include unicellular yeasts and multicellular mushrooms and molds.
General Biology of a Fungus
Fungi are neither animals nor plants. 
	•	The body of a fungus is composed of thin filaments of cells, called hyphae, that form a mass called a mycelium.
	•	The cell wall contains chitin.
	•	Fungi produce windblown spores during both asexual and sexual reproduction.
Ecological Benefits of Fungi
	•	Fungi are saprotrophs that carry on external digestion. As decomposers, fungi keep ecological cycles going in the biosphere.
	•	Lichens (fungi plus cyanobacteria or green algae) are primary colonizers in poor soils or on rocks.
	•	Mycorrhizal fungi grow on or in plant roots and help the plant absorb minerals and water.
Economic Benefits of Fungi
	•	Fungi help produce foods and medicines, as well as serving as a source of food themselves.
Fungi as Disease-Causing Organisms
	•	Fungal pathogens of plants include blasts, smuts, and rusts that attack crops of great economic importance, such as rice and wheat.
	•	Human diseases caused by fungi include thrush, ringworm, and histoplasmosis.
Thinking Scientifically
	1.	Bare-root pine tree seedlings transplanted into open fields often grow very slowly. However, pine seedlings grow much more vigorously if they are dug from their native environment and then transplanted into a field, as long as some of the original soil is retained on the seedlings. Why is it so important to retain some native soil on the seedlings?
	2.	Some orchids produce flowers that have modified petals resembling female wasps. Male wasps emerge from their pupal stage a week earlier than females. The male wasps are attracted to the flowers even though they are not rewarded with nectar. Why do you suppose the male wasps visit the flowers? Why is this an especially effective pollen dispersal strategy?
Testing Yourself
Choose the best answer for each question.
	1.	Which of the following is not a plant adaptation to land?
	a. recirculation of water
	b. protection of embryo in maternal tissue
	c. development of flowers
	d. creation of vascular tissue
	e. seed production
	2.	Plant spores are
	a. haploid and genetically different from each other.
	b. haploid and genetically identical to each other.
	c. diploid and genetically different from each other.
	d. diploid and genetically identical to each other.
	3.	Label the parts of the generalized plant life cycle in the following illustration.
For questions 4–10, identify the group(s) to which each feature belongs. Each answer in the key may be used more than once. Each question may have more than one answer.
Key:
	a. nonvascular plants
	b. ferns
	c. gymnosperms
	d. angiosperms
	4.	Produce lignin in cell walls.
	5.	Exhibit alternation of generations. 
	6.	Produce seeds. 
	7.	Lack true roots, stems, and leaves. 
	8.	Produce ovules that are not completely surrounded by sporophyte tissue. 
	9.	Produce swimming sperm. 
	10.	Produce flowers. 
	11.	Label the parts of the flower in the following illustration.
	12.	A fruit is derived from
	a. the corolla.
	b. an ovary.
	c. the stamen.
	d. an ovule.
	e. the calyx.
	13.	Which of the following statements about fungi is false?
	a. Most fungi are multicellular.
	b. Fungal cell walls are composed of cellulose.
	c. Fungi are nonmotile.
	d. Fungi digest their food before ingesting it.
	e. Fungi have haploid and diploid stages in their sexual life cycle.
	14.	The spores produced by a fruiting body are
	a. haploid and genetically different from each other.
	b. haploid and genetically identical to each other.
	c. diploid and genetically different from each other.
	d. diploid and genetically identical to each other.
	15.	Which of the following is a true statement?
	a.	
People don’t eat mushrooms because they might be poisonous.
	b. Penicillin is derived from a fungus.
	c. The alcohol from yeast fermentation makes bread rise.
	d. Fungi are prokaryotes like bacteria.
	e. Fungi ingest their food in the same way animals do. 
	16.	A fungal spore
	a.	 contains an embryonic organism.
	b. germinates directly into an organism.
	c. is always windblown.
	d. is most often diploid.
	e. Both b and c are correct.
	17.	Lichens
	a. can live on bare rocks.
	b. need a nitrogen source in order to live.
	c. are parasitic on trees.
	d. can reproduce asexually.
	e. Both a and d are correct.
	18.	Mycorrhizal fungi 
	a. are a type of lichen.
	b. help plants gather solar energy.
	c. help plants gather inorganic nutrients.
	d. All of these are correct.
	19.	Tinea infections are generally found
	a. in the eye.
	b. in cartilage.
	c. on the skin.
	d. in the vagina.
	20.	Which of the following diseases is (are) caused by Candida?
	a. oral thrush
	b. athlete’s foot
	c. vaginal yeast infection
	d. ringworm
	e. Both a and c are correct.
	21.	The gametophyte is the dominant generation in
	a. ferns.
	b. mosses.
	c. gymnosperms.
	d. angiosperms.
	e. More than one of these are correct.
	22.	A seed is a mature
	a. embryo.
	b. ovule.
	c. ovary.
	d. pollen grain.
	23.	A mushroom is like a plant because it
	a. is a multicellular eukaryote.
	b. produces spores. 
	c. is adapted to a land environment. 
	d. is photosynthetic.
	e. All but d are correct. 
Go to www.mhhe.com/maderessentials for more quiz questions.
Bioethical Issue
Bristlecone pines are among the oldest trees on Earth. In 1964, a graduate student working in the southwestern United States took core samples from several trees to determine their age. One tree was found to be over 4,000 years old. When the student’s coring tool broke, the U.S. Forest Service gave him permission to cut down the tree in order to accurately determine its age. The tree was found to be 4,862 years old—the oldest known living creature on Earth.
Did the student do anything wrong, scientifically or morally, considering that he was given permission to cut down the tree? Who should be responsible for protecting unique trees like the bristlecone pine?
Understanding the Terms
alternation of generations286
angiosperm286, 292
anther292
bryophyte286
calyx292
carpel292
cone291
conifer291
corolla292
cotyledon293
double fertilization293
endosperm293
fern289
filament292
flower285
frond289
fruit293
fungus (pl., fungi)296
gametophyte286
gymnosperm286, 291
lichen299
lignin288
mycelium296
mycorrhizal fungi299
nonvascular plant287
ovary292
ovule290
petal292
phloem288
pollen grain290
pollen tube293
pollination290
saprotroph298
seed285
sepal292
spore286
sporophyte286
stamen292
stigma292
style292
vascular plant288
vascular tissue285
xylem288
Match the terms to these definitions:
a.	_______________	Structure composed of embryo and stored food within a protective coat. 
b.	_______________	Part of a flower that produces pollen. 
c.	_______________	Water-conducting vascular tissue. 
d.	_______________	Fern leaves. 
e.	_______________	Male gametophytes in seed plants.
f.	_______________	Mass of fungal filaments.
g.	_______________	Decomposer.
h.	_______________	Generation that produces spores in a plant. 
i.	_______________	Structure in flowering plants derived from the ovary. 

Without plants, there would be no animals.
Twelve plants, wheat among them, stand between humans and starvation. 
Cotton is a textile fiber that accounts for about half of all the fabric sold in the world. 
Figure 18.2Evolution of plants.
The evolution of plants is marked by four significant events: protection of the embryo, evolution of vascular tissue, evolution of the seed, and evolution of the flower. 
Figure 18.1Representatives of the four major groups of plants.
Check Your Progress
	1.	List the similarities between green algae and plants.
	2.	
List the four evolutionary events that allowed plants to successfully inhabit land.
Answers:1. Both green algae and plants contain chlorophylls a and b plus accessory pigments, store carbohydrates as starch, and have cellulose in their cell walls.2. Protection of the embryo within the body of the plant, the presence of vascular tissue, seed production, and the presence of flowers.
Check Your Progress
	1.	Explain what is meant by alternation of generations. 
	2.	What generation is dominant among mosses? In other groups of plants? Give an explanation for this trend of generation dominance among plants. 
Answers:1. Every plant exists in two forms. The diploid sporophyte produces haploid spores by meiosis. The haploid gametophyte produces gametes.
2. The gametophyte is dominant among mosses, and the sporophyte is dominant among the other groups of plants. The sporophyte evolved vascular tissue, and vascular tissue is a distinct advantage in a land environment. 
Figure 18.5Mosses.
In bryophytes, exemplified by mosses, the gametophyte is the dominant generation. Sperm swim from male shoots to female shoots, and the zygote develops into an attached sporophyte. The sporophyte produces haploid spores that develop into gametophyte shoots.
Figure 18.4Reduction in the size of the -gametophyte.
Notice the reduction in the size of the gametophyte among these representatives of today’s plants. This trend occurred as plants became adapted for life on land.
Figure 18.3Alternation of generations.
Figure 18.8Diversity of ferns.
Figure 18.9Fern life cycle.
 The sporophyte is dominant in ferns.  In the fern shown here, sori are on the underside of a fern. Each sorus, protected by an indusium, contains sporangia, in which meiosis occurs and spores are produced and then released.  A spore germinates into a prothallus (the gametophyte), which has sperm-bearing and egg-bearing structures on its underside.  Fertilization takes place when moisture is present, because the flagellated sperm must swim in a film of water to the egg.  The resulting zygote begins its development inside an archegonium, and eventually a young sporophyte -becomes visible. The young sporophyte develops, and fronds appear.
Figure 18.6Vascular tissue.
The roots, stems, and leaves of vascular plants, such as this flowering plant, have vascular tissue (xylem and phloem). Xylem transports water and minerals; phloem transports organic compounds.
Figure 18.7The Carboniferous period.
Growing in the swamp forests of the Carboniferous period were treelike club mosses (left), treelike horsetails (right), and lower, fernlike foliage (left). When the trees fell, they were covered by water and did not decompose completely. Sediment built up and turned to rock, the pressure of which caused the organic material to become coal, a fossil fuel that helps run our industrialized society today.
Figure 18.12Cycads.
Cycads are an ancient group of gymnosperms that are threatened today because they are slow growing. Herbivorous dinosaurs of the Mesozoic era most likely fed on cycad seeds. 
Figure 18.13Conifers.
a. Pine trees are the most common of the conifers. The pollen cones (male) are smaller than the seed cones (female) and produce plentiful pollen. A cluster of pollen cones may produce more than one million pollen grains. Other conifers include (b) the spruces, which make beautiful Christmas trees, and (c) junipers, which have fleshy seed cones.
Figure 18.10Seed anatomy.
A split bean seed showing the seed coat, embryo, and stored food (cotyledon). 
Figure 18.11Production of a seed.
Development of the male gametophyte (above) begins when a microspore mother cell undergoes meiosis to produce microspores, each of which becomes a pollen grain. Development of the female gametophyte (below) begins in an ovule, where a megaspore mother cell undergoes meiosis to produce megaspores, only one of which will undergo mitosis to become the female gametophyte. During pollination, the pollen grain is carried to the vicinity of the ovule. The pollen grain germinates, and a nonflagellated sperm travels in a pollen tube to the egg produced by the female gametophyte. Following fertilization, the zygote becomes the embryo; tissue within the ovule becomes the stored food, and the ovule wall becomes the seed coat.
Check Your Progress
	1.	What structure in seed plants becomes a seed? 
	2.	What are the components of a seed? 
	3.	What is the difference between pollination and fertilization?
Answers:1. Ovule.2. A seed contains an embryo, stored food, and a seed coat.3. Pollination is the delivery of the male gametophyte to the region of the female gametophyte. Fertilization occurs when the sperm from the male gametophyte unites with the egg in the female gametophyte.
Figure 18.16Life cycle of flowering eudicot.
Figure 18.14Amborella trichopoda.
Molecular data suggest this plant is most closely related to the first flowering plants.
Figure 18.15Generalized flower.
A flower has four main parts: sepals, petals, stamens, and carpels. A stamen has a filament and an anther. A carpel has an ovary, a style, and a stigma. The ovary contains ovules.
Check Your Progress
	1.	List the components of the stamen. Where is pollen formed?  
	2.	List the components of the carpel. Which part becomes a seed? The fruit? 
Answers:1. The stamen contains the anther and the filament. Pollen forms in the pollen sacs of the anther.2. The carpel contains the stigma, style, and ovary. An ovule in the ovary becomes a seed, and the ovary becomes the fruit. 
Figure 18.19Wooden furniture. 
Maple, walnut, and mahogany furniture is made from the wood of these trees.
Figure 18.17Pollinators.
a. A bee-pollinated flower is a color other than red (bees can’t see red). b. Butterfly-pollinated flowers are wide, allowing the butterfly to land. c. Hummingbird-pollinated flowers are curved back, allowing the bird’s beak to reach the nectar. d. Bat-pollinated flowers are large and sturdy and able to withstand rough treatment.
Figure 18.18Pea flower and the development of a pea pod.
a. Pea flower. b. Pea flower still has its petals soon after fertilization. c. Petals fall and the pea pod is quite noticeable. d. Pea pod is clearly visible developing from an ovary.
Check Your Progress
	1.	Which groups of plants produce seeds? Give examples of each group.
	2.	What features of the flowering plant life cycle are not found in any other group?
	3.	Which are more showy, wind-pollinated flowers or animal-pollinated flowers? Why? 
Answers:1. Gymnosperms (cone-bearing, such as cycads and pine trees) and angiosperms (flowering plants, such as fruit trees and garden plants) produce seeds.2. Presence of an ovary leads to production of seeds enclosed by a fruit. Animals are often used as pollinators.3. Animal-pollinated flowers are showy, and in different ways, such as color and fragrance, attract their particular pollinators. 
Check Your Progress
	1.	What are some economic benefits of flowering plants? 
	2.	What are some ecological benefits of flowering plants?
Answers:1. Use of fruits as food, such as beans and rice; use of fruits as textile fibers, such as cotton; use of wood as a building material for construction and furniture; and use of chemicals flowering plants produce as pharmaceuticals, such as taxol.2. Flowering plants are producers in ecosystems; forests help keep ecological cycles functioning and keep the environment healthy.
Figure 18.21Body of a fungus.
a. The body of a fungus is called a mycelium. b. A mycelium contains many individual chains of cells, and each chain is called a hypha.
Figure 18.20Diversity of fungi.
a. Scarlet hood, an inedible mushroom. b. Spores exploding from a puffball. c. Common bread mold. d. A morel, an edible fungus.
Figure 18.22Life cycle of black bread mold.
During asexual reproduction, sporangia produce asexual spores (purple and blue arrows). During sexual reproduction, two hyphae tips fuse, and then two nuclei fuse, forming a zygote that develops a thick, resistant wall (zygospore). When conditions are favorable, the zygospore germinates, and meiosis within a sporangium produces windblown spores (yellow arrows). 
Figure 18.24Carnivorous fungus.
Fungi like this oyster fungus, a type of bracket fungus, grow on trees because they can digest cellulose and lignin. If this fungus meets a roundworm in the process, it immobilizes the worm and digests it also. The worm is a source of nitrogen for the fungus. 
Figure 18.23Sexual reproduction produces mushrooms.
a. Fusion of + and 2 hyphae tips results in hyphae that form the mushroom (a fruiting body). The nuclei fuse in clublike structures attached to the gills of a mushroom, and meiosis produces spores. b. A circle of mushrooms is called a fairy ring.
Check Your Progress
	1.	Contrast sexual reproduction in black bread mold with sexual reproduction in a mushroom.
	2.	How do fungi contribute to ecological cycling?
Answers:1. In black bread mold, meiosis produces spores in sporangia. In a mushroom, meiosis produces spores in clublike structures attached to the gills.2. When fungi secrete enzymes that digest the remains of plants and animals in the environment, inorganic nutrients are returned to plants.   
Check Your Progress
	1.	What organisms make up a lichen? Why is the relationship mutualistic?
	2.	What is the ecological importance of mycorrhizal fungi? 
Answers:1. A lichen is composed of a fungus and a cyanobacterium or a green alga. Each partner gives to the other. The fungus provides protection, water, and minerals, and the photosynthesizer provides organic nutrients.
2. Mycorrhizal fungi allow plants to colonize degraded land, and this helps restore the area.
Figure 18.25Lichens. 
a. Morphology of a lichen. b. Examples of lichens.
Figure 18.26Plant growth experiment.
A soybean plant without mycorrhizal fungi grows poorly compared to two others infected with different strains of mycorrhizal fungi.
Figure 18.28Plant fungal disease.
Peach leaf curl causes lesions on the leaf by invading the leaf, as shown in (a) the photo and (b) the diagram.
Figure 18.27Commercial importance of fungi.
Delicacies such as various types of mushrooms are fungi, and some of our other favorite foods require the participation of fungi to produce them. Significant medicines have also come from fungi. 
Figure 18.29Human fungal diseases.
a. Thrush, or oral candidiasis, is characterized by the formation of white patches on the tongue. b. Ringworm and (c) athlete’s foot are caused by Tinea spp.
Check Your Progress
	1.	List the ways in which humans use fungi.
	2.	Why is a white button mushroom white and a portabella mushroom brown?
	3.	What are some common fungal diseases of humans? When is a fungal disease opportunistic?
Answers:1. Fungi are used to process food and to make various types of medicines. Mushrooms are an important source of food.2. A white button mushroom has not been allowed to mature. A portabella mushroom is mature, and spores are present.3. Common fungal diseases include Candida (yeast) infections, ringworm, athlete’s foot, and histoplasmosis. 
An opportunistic fungal disease occurs in a person with a weakened immune system.

Table 18.1	How Fungi Differ from Plants and Animals	

Feature	Fungi	Plants	Animals	
Nutrition	Chemoheterotrophic by absorption	Photosynthetic	Chemoheterotrophic by ingestion	
Movement	Nonmotile	Nonmotile	Motile	
Body	Mycelium of hyphae	Specialized tissues/organs	Specialized tissues/organs	
Adult chromosome number	Haploid	Haploid/diploid  	Diploid	
Cell wall	Composed of chitin	Composed of cellulose	No cell wall	
Reproduction	Spores	Spores/gametes	Gametes	
 Check Your Progress
	1.	Compare and contrast nutrition in fungi with that in animals. 
	2.	Describe the relationship between a mycelium and hyphae.
Answers:1. Both fungi and animals are chemoheterotrophs, but animals ingest their food, while fungi absorb organic nutrients from the environment.2. A mycelium is a mass of hyphae (filaments of cells).
a. 
Male cycad with a pollen-
bearing cone.
b. 
Female cycad with a seed-bearing cone.
Maple wood
Mahogany wood
Walnut wood