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Chapter 17 The Evolution of Plant and Fungal
Diversity
Green algae (charophytes) are the ancestors of plants
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17.1 Plants have adaptations for life on land
More than 500 million years ago, the algal
ancestors of plants may have carpeted moist
fringes of lakes and coastal salt marshes.
Plants and green algae called charophytes
– are thought to have evolved from a common ancestor,
– have complex multicellular bodies, and
– are photosynthetic eukaryotes.
– Algae do not have tissues like plants
© 2012 Pearson Education, Inc.
17.1 Plants have adaptations for life on land
Life on land offered many opportunities for plant
adaptations that took advantage of
– unlimited sunlight,
– abundant CO2, and
– initially, few pathogens or herbivores.
© 2012 Pearson Education, Inc.
17.1 Plants have adaptations for life on land
But life on land had disadvantages too. On land,
plants must
– maintain moisture inside their cells, to keep from drying
out,
– support their body in a nonbuoyant medium,
– reproduce and disperse offspring without water, and
– obtain resources from soil and air.
© 2012 Pearson Education, Inc.
17.1 Plants have adaptations for life on land
Unlike land plants, algae
– generally have no rigid tissues,
– are supported by surrounding water,
– obtain CO2 and minerals directly from the water
surrounding the entire algal body,
– receive light and perform photosynthesis over most of
their body,
– use flagellated sperm that swim to fertilize an egg, and
– disperse offspring by water.
© 2012 Pearson Education, Inc.
Figure 17.1C
Key
Vascular
tissue
Pollen
Spores
Leaf
Spores
Flagellated
sperm
Alga
Surrounding
water supports
alga. Whole alga
Leaf
performs photosynthesis; absorbs Stem
water, CO2, and
minerals from
the water.
Roots
Flagellated
sperm
Holdfast
(anchors alga)
Seed
Flagellated
sperm
Stem
Leaf
Roots
Moss
Stomata only on sporophytes;
primitive roots anchor plants;
no lignin; no vascular tissue;
fertilization requires moisture
Fern
Stomata; roots anchor
plants, absorb water;
lignified cell walls;
vascular tissue;
fertilization requires
moisture
Stem
Roots
Pine tree
Stomata;
roots anchor plants, absorb water;
lignified cell walls; vascular tissue;
fertilization does not require moisture
17.1 Plants have adaptations for life on land
Land plants maintain moisture in their cells using
– a waxy cuticle and
– cells that regulate the opening and closing of stomata.
Land plants obtain
– water and minerals from roots in the soil and
– CO2 from the air and sunlight through leaves.
Growth-producing regions of cell division, called
apical meristems, are found near the tips of stems
and roots.
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17.1 Plants have adaptations for life on land
In many land plants, water and minerals move up
from roots to stems and leaves using vascular
tissues.
– Xylem
– consists of dead cells and
– conveys water and minerals.
– Phloem
– consists of living cells and
– conveys sugars.
© 2012 Pearson Education, Inc.
17.1 Plants have adaptations for life on land
In all plants, the
– gametes and embryos must be kept moist,
– fertilized egg (zygote) develops into an embryo while
attached to and nourished by the parent plant, and
– life cycle involves an alternation of a
– haploid generation, which produces eggs and sperm, and
– diploid generation, which produces spores within protective
structures called sporangia.
Pines and flowering plants have pollen grains,
structures that contain the sperm-producing cells.
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Fig. 17.2 Plant diversity reflects the evolutionary history of the plant kingdom
Land plants
Hornworts
1
Nonvascular
plants
(bryophytes)
Ancestral
green
alga
Liverworts
Origin of land plants
(about 475 mya)
Mosses
Angiosperms
500
450
400
350
Millions of years ago (mya)
300
0
Vascular plants
Origin of seed plants
(about 360 mya)
Seed
plants
3
Lycophytes (club mosses,
spike mosses, quillworts)
Pterophytes or
Monilophytes (ferns,
horsetails, whisk ferns)
Gymnosperms
Seedless
vascular
plants
2
Origin of vascular plants
(about 425 mya)
17.2 Plant diversity reflects the evolutionary
history of the plant kingdom
Early diversification of plants gave rise to seedless,
nonvascular plants called bryophytes, including
– _____, that we saw in laboratory
– liverworts, and
– hornworts.
© 2012 Pearson Education, Inc.
17.2 Plant diversity reflects the evolutionary
history of the plant kingdom
These plants resemble other plants in having apical
meristems and embryos retained on the parent
plant, but they lack
– true roots,
– leaves, and
– stems.
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Figure 17.2B
Bryophytes
_____
Liverwort
Hornwort
17.2 Plant diversity reflects the evolutionary
history of the plant kingdom
About 425 million years ago, vascular plants evolved
with lignin-hardened vascular tissues.
The seedless vascular plants include
– lycophytes (including club mosses) and
– Pterophytes or monilophytes (ferns and their relatives).
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Figure 17.2C
Seedless vascular plants
Club moss (a lycophyte).
Spores are produced in the
fern (a pterophyte or
upright tan-colored structures.
monilophyte), spores under ____
17.2 Plant diversity reflects the evolutionary
history of the plant kingdom
The first vascular plants with seeds evolved about
360 million years ago.
A seed consists of an embryo packaged with a food
supply within a protective covering.
© 2012 Pearson Education, Inc.
17.2 Plant diversity reflects the evolutionary
history of the plant kingdom
Vascular plants with seeds include
– gymnosperms (including ginkgo, cycad, and conifer
species) and
– angiosperms (such as flowering trees and grasses).
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17.2 Plant diversity reflects the evolutionary
history of the plant kingdom
Gymnosperms
– have naked seeds that are not produced in fruits and
– include ginkgo, cycad, and conifer species.
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Figure 17.2D
Gymnosperms
Cycad
Ginkgo
Ephedra
(Mormon tea)
A conifer
17.2 Plant diversity reflects the evolutionary
history of the plant kingdom
Angiosperms
– are flowering plants and
– include flowering trees and grasses.
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Figure 17.2E
Angiosperms
A tropical jacaranda tree
Green foxtail, a grass
ALTERNATION
OF GENERATIONS
AND PLANT LIFE CYCLES
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17.3 Haploid and diploid generations alternate in
plant life cycles
Plants have an alternation of generations in
which the haploid and diploid stages are distinct,
multicellular bodies.
– The haploid gametophyte produces gametes (eggs or
sperm) by mitosis.
– Fertilization results in a diploid zygote.
– The zygote develops into the diploid sporophyte, which
produces haploid spores by meiosis.
– Spores grow into gametophytes.
© 2012 Pearson Education, Inc.
Figure 17.3
THE PLANT LIFE CYCLE
Key
Gametophyte
plant (n)
Haploid (n)
Diploid (2n)
Sperm (n)
Egg (n)
Spores (n)
Meiosis
Fertilization
Zygote (2n)
Sporophyte
plant (2n)
17.3 Haploid and diploid generations alternate in
plant life cycles
Gametophytes make up a bed of moss.
– Gametes develop in male and female gametangia.
– Sperm swim through water to the egg in the female
gametangium.
© 2012 Pearson Education, Inc.
17.3 Haploid and diploid generations alternate in
plant life cycles
The zygote
– develops within the gametangium into a mature
sporophyte,
– which remains attached to the gametophyte.
Meiosis occurs in sporangia at the tips of the
sporophyte stalks.
Haploid spores are released from the sporangium
and develop into gametophyte plants.
© 2012 Pearson Education, Inc.
Figure 17.3
A Moss Life Cycle
Key
Haploid (n)
Diploid (2n)
Male
gametangium
Sperm
Spores (n)
Gametophyte plants (n)
Egg
Female
gametangium
Sporangium
Sporophyte
Meiosis
Gametophyte
Fertilization
Zygote
Gametophyte or sporophyte?
Gametophyte or sporophyte?
17.3 Haploid and diploid generations alternate in
plant life cycles
Fern gametophytes are small and inconspicuous.
Gametophytes produce flagellated sperm that swim
to the egg and fertilize it to produce a zygote.
The zygote initially develops within the female
gametangia but eventually develops into an
independent sporophyte.
© 2012 Pearson Education, Inc.
17.3 Haploid and diploid generations alternate in
plant life cycles
Sporangia develop on the underside of the leaves
of the sporophyte.
Within the sporangia, cells undergo meiosis to
produce haploid spores.
Spores are released and develop into
gametophytes.
© 2012 Pearson Education, Inc.
Figure 17.3
A Fern Life Cycle
Key
Gametophyte
plant (n)
Haploid (n)
Diploid (2n)
Male
gametangium
Spores
Sperm
Female
gametangium
Egg
Meiosis
Mature
sporophyte
Fertilization
Zygote
New sporophyte
growing from the
gametophyte
17.4 Seedless vascular plants dominated vast
“coal forests”
Two groups of seedless plants formed vast ancient
forests in low-lying wetlands during the
Carboniferous period (360–299 million years ago):
– lycophytes (such as club mosses) and
– Pterophytes or monilophytes (such as ferns).
When these plants died, they formed peat deposits
that eventually formed coal.
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Figure 17.4
17.4 Seedless vascular plants dominated vast
“coal forests”
As temperatures dropped during the late
Carboniferous,
– glaciers formed,
– the climate turned drier,
– the vast swamps and forests began to disappear, and
– wind-dispersed pollen and protective seeds gave seed
plants a competitive advantage.
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17.5 Pollen and seeds are key adaptations to life
on land
A pine tree is a sporophyte.
Tiny gametophytes grow in sporophyte cones.
The ovule is a key adaptation, a protective device
for all the female stages in the life cycle, as well as
the site of
– pollination,
– fertilization, and
– embryonic development.
© 2012 Pearson Education, Inc.
17.5 Pollen and seeds are key adaptations to life
on land
A sperm from a pollen grain fertilizes an egg in the
female gametophyte.
The zygote develops into a sporophyte embryo.
The ovule becomes the seed with
– stored food and
– a protective seed coat.
The seed is a key adaptation for life on land and a
major factor in the success of seed plants.
© 2012 Pearson Education, Inc.
Figure 17.5
Longitudinal
section of
ovulate cone
Longitudinal
section of
pollen cone
Sporangia
Figure 17.5B
Seed coat
Spore wall
Sporangium (2n)
(produces spore)
Female
gametophyte (n)
Egg nucleus (n)
Spore
wall
Ovulate cone
Spore (n)
Male gametophyte
(within a germinated
pollen grain) (n)
Pollen grain (n)
Discharged
sperm nucleus (n)
Food
supply
Pollen tube
Embryo (2n)
(new sporophyte)
17.6 The flower is the centerpiece of angiosperm
reproduction
Flowers house separate male and female sporangia
and gametophytes.
Flowers are the sites of
– pollination and
– fertilization.
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17.6 The flower is the centerpiece of angiosperm
reproduction
Flowers usually consist of
– sepals, which enclose the flower before it opens,
– petals, which attract animal pollinators,
– stamens, which include a filament and anther, a sac at
the top of each filament that contains male sporangia
and releases pollen, and
– Carpels or pistils, the female reproductive structure,
which produce eggs.
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17.8 The flower is the centerpiece of angiosperm
reproduction
Ovules develop into seeds.
Ovaries mature into fruit.
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Figure 17.6B
Stigma
Style
_____
Ovary
Anther
Filament
_____
_____
Ovule
Receptacle
_____
17.6 The angiosperm plant is a sporophyte with
gametophytes in its flowers
Key events in a typical angiosperm life cycle
1. Meiosis in the anthers produces haploid spores that form
the male gametophyte (pollen grains).
2. Meiosis in the ovule produces a haploid spore that forms
the few cells of the female gametophyte, one of which
becomes the egg.
3. Pollination occurs when a pollen grain lands on the
stigma. A pollen tube grows from the pollen grain to the
ovule.
4. The tube carries a sperm that fertilizes the egg to form a
zygote.
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17.6 The angiosperm plant is a sporophyte with
gametophytes in its flowers
Key events in a typical angiosperm life cycle,
continued
5. Each ovule develops into a seed, consisting of
– an embryo (a new sporophyte) surrounded by a food supply and
– a seed coat derived from the integuments.
6. While the seeds develop, the ovary’s wall thickens,
forming the fruit that encloses the seeds.
7. When conditions are favorable, a seed germinates.
© 2012 Pearson Education, Inc.
Figure 17.7
Anther
1
Pollen grains (n)
(male gametophytes)
Meiosis
2
3
Egg within
a female
gametophyte (n)
Stigma
Pollen grain
Pollen tube
Meiosis
Ovary
Sporophyte
(2n)
Ovule
Ovule
containing
female sporangium
(2n)
Sperm
Germination
7
Seeds
Food
supply
6
Fertilization
Seed coat
Fruit
(mature ovary)
5
Seed
Embryo (2n)
4
Zygote
(2n)
Key
Haploid (n)
Diploid (2n)
17.8 The structure of a fruit reflects its function
in seed dispersal
Fruits are
– ripened ovaries of flowers and
– adaptations that disperse seeds.
Seed dispersal mechanisms include relying on
– wind,
– hitching a ride on animals, or
– fleshy, edible fruits that attract animals, which then
deposit the seed in a supply of natural fertilizer at some
distance from the parent plant.
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Figure 17.8A-C
Fruit
Seed
dispersal
17.9 Angiosperms sustain us—and add spice to
our diets
Most human food is provided by the fruits and
seeds of angiosperms.
– Corn, rice, wheat, and other grains are dry fruits.
– Apples, cherries, tomatoes, and squash are fleshy fruits.
– Spices like pepper (Piper nigrum)
and cumin are also angiosperm
fruits.
© 2012 Pearson Education, Inc.
17.10 EVOLUTION CONNECTION: Pollination
by animals has influenced angiosperm
evolution
About 90% of angiosperms use animals to transfer
their pollen.
– Birds are usually attracted by colorful flowers, often red,
but without scent.
– Most beetles are attracted by fruity odors, but are
indifferent to color.
– Night-flying bats and moths are usually attracted by
large, highly scented flowers that are often white.
– Wind-pollinated flowers typically produce large amounts
of pollen.
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Figure 17.10B
17.11 CONNECTION: Plant diversity is vital to the
future of the world’s food supply
Some new crops may come from the hundreds of
species of nutritious fruits, nuts, and grains that
people gather and use locally.
Figure 17.11a-0 Sugar plums (left) and African plums (right), two wild fruits that may be ripe
for domestication
DIVERSITY OF FUNGI
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17.12 Fungi absorb food after digesting it outside
their bodies
Fungi
– are absorptive heterotrophic eukaryotes,
– secrete powerful enzymes to digest their food externally,
and
– acquire their nutrients by absorption.
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17.12 Fungi absorb food after digesting it outside
their bodies
Most fungi consist of a mass of threadlike hyphae
making up a mycelium.
Hyphal cells
– are separated by cross-walls with pores large enough for
organelles to cross, and
– have a huge surface area to secrete digestive enzymes
and absorb food.
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Figure 17.12A
Figure 17.12B
Reproductive
structure
Hyphae
Spore-producing
structures (tips of hyphae)
Mycelium
17.12 Fungi absorb food after digesting it outside
their bodies
Fungal hyphae
– are surrounded by a cell wall made of chitin instead of
cellulose as in plants and some algae.
Some fungi
– are parasites and
– obtain their nutrients at the expense of living plants or
animals.
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17.13 Fungi produce spores in both asexual and
sexual life cycles
In many fungi, sexual fusion of haploid hyphae
leads to a heterokaryotic stage, in which cells
contain two genetically distinct haploid nuclei.
– Hours or centuries may pass before parental nuclei fuse
to form a short-lived diploid phase.
– Zygotes undergo meiosis to produce haploid spores.
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17.13 Fungi produce spores in both asexual and
sexual life cycles
In asexual reproduction, spore-producing structures
arise from haploid mycelia that have undergone
neither a heterokaryotic stage or meiosis.
Many fungi that reproduce sexually can also
produce spores asexually.
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17.13 Fungi produce spores in both asexual and
sexual life cycles
Molds are any rapidly growing fungus that
reproduces asexually by producing spores.
Yeasts are single-celled fungi that reproduce
asexually by cell division or budding.
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Figure 17.13
Key
Haploid (n)
Heterokaryotic (n + n)
(unfused nuclei)
Diploid (2n)
Heterokaryotic
stage
Fusion of nuclei
1
Fusion of cytoplasm
Spore-producing
structures
Spores
(n)
Zygote
(2n)
Sexual
reproduction
Asexual Mycelium
reproduction
4
2
Meiosis
Spore-producing
structures
Germination
3
Germination
Spores (n)
17.13 Fungi produce spores in both asexual and sexual life cycles
The life cycle of a black bread mold is typical of zygomycetes.
Hyphae reproduce asexually by producing spores in sporangia
at the tips of upright hyphae. When food is depleted, the
fungus reproduces sexually. Mycelia of different mating types
join and produce a zygosporangium, a thick-walled cell
containing heterokaryotic nuclei from two parents that can
tolerate dry, harsh conditions. When conditions are
favorable, the
parental nuclei fuse
to form diploid
zygotes, which
undergo meiosis
producing haploid
spores.
17.14 Fungi are classified into five groups
There are over 100,000 described fungi species.
Suspected but as yet undescribed species may
number as many as 1.5 million.
Sexual reproductive structures are often used to
classify fungi.
Fungi and animals may have diverged
– from a flagellated ancestor
– more than 1 billion years ago.
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17.16 Fungi are classified into five groups
Chytrids are the
– only fungi with flagellated spores and
– earliest lineage of fungi.
Chytrid fungi are
– common in lakes, ponds, and soil and
– linked to the widespread decline of amphibian species.
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Figure 17.14A
Chytrids
Zygomycetes
(zygote fungi)
Glomeromycetes
(arbuscular
mycorrhizal fungi)
Ascomycetes
(sac fungi)
Basidiomycetes
(club fungi)
17.14 Fungi are classified into five groups
Zygomycetes, or zygote fungi
– are characterized by their protective zygosporangium,
where zygotes produce haploid spores by meiosis.
– This diverse group includes fast-growing molds that
attack
– bread
– peaches,
– strawberries,
– sweet potatoes, and
– some animals.
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17.14 Fungi are classified into five groups
Glomeromycetes
– form a distinct type of mycorrhizae, in which hyphae that
invade plant roots in a symbiotic relationship
– Mycorrhizal fungi absorb phosphorus and other essential materials
from the soil and make them available to the plant and sugars
produced by the plant through photosynthesis nourish the
mycorrhizal fungi.
– About 80% of all plants have symbiotic partnerships with
glomeromycetes.
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17.14 Fungi are classified into five groups
Ascomycetes, or sac fungi
– form saclike structures called asci, which produce
spores in sexual reproduction,
– live in marine, freshwater, and terrestrial habitats, and
– range in size from unicellular yeast to elaborate morels
and cup fungi.
– Some ascomycetes live with green algae or
cyanobacteria in symbiotic associations called lichens.
© 2012 Pearson Education, Inc.
Figure 17.14D
Ascomycetes
Edible morels
Cup fungus
17.14 Fungi are classified into five groups
Basidiomycetes, or club fungi,
– include common mushrooms, puffballs, and shelf
fungi and
– are named for their club-shaped, spore-producing
structure called a basidium.
These fungi include
– important forest decomposers and
– particularly destructive plant parasites called rusts
and smuts.
© 2012 Pearson Education, Inc.
Figure 17.14E
Basidiomycetes
Mushrooms
A puffball
Shelf fungi
17.17 Fungal groups differ in their life cycles and
reproductive structures
The life cycle of a black bread mold is typical of
zygomycetes.
Hyphae reproduce asexually by producing spores
in sporangia at the tips of upright hyphae.
© 2012 Pearson Education, Inc.
17.15 CONNECTION: Fungi have enormous
ecological benefits
Fungi
– supply essential nutrients to plants through symbiotic
mycorrhyizae,
– along with prokaryotes are essential decomposers in
ecosystems, breaking down organic matter and
restocking the environment with vital nutrients essential
for plant growth, and
– may also be used to digest petroleum products to clean
up oil spills and other chemical messes.
17.16 CONNECTION: Fungi have many practical
uses
Fungi have many practical uses for humans.
– We eat mushrooms and cheeses modified by fungi.
– Yeasts produce alcohol and cause bread to rise.
– Some fungi provide antibiotics that are used to treat
bacterial disease.
Figure 17.16A
Figure 17.16B
Penicillium
(mold)
Staphylococcus
aureus (bacteria)
Zone of
inhibited
growth
17.17 Lichens are symbiotic associations of fungi
and photosynthetic organisms
Lichens consist of algae or cyanobacteria within a
mass of fungal hyphae.
– Many lichen associations are mutualistic.
– The fungus receives food from its photosynthetic
partner.
– The fungal mycelium helps the alga absorb and retain
water and minerals.
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17.17 Lichens are symbiotic associations of fungi
and photosynthetic organisms
Lichens are important pioneers on new land, where
they help to form soil.
Lichens are sensitive to air pollution, because they
obtain minerals from the air.
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Figure 17.17A Two species of lichens commonly found on coastal rocks
Figure 17.17b
Algal cell
Fungal
hyphae
Figure 17.17C Reindeer moss, a lichen fed on by caribou in the
Arctic Tundra
17.18 SCIENTIFIC THINKING: Mycorrhizae may
have helped plants colonize land
About 80% of all plant species establish symbioses
with glomeromycetes, mycorrhizal fungi that form
bushy structures called arbuscules in root cells,
which increase surface area for nutrient absorbtion.
The fungi obtain sugar from root cells. Both benefit
in the symbiosis called _______________
The presence of mycorrhizal associations in almost
all major lineages of present-day plants suggests an
ancient origin for plant-fungus symbioses.
Fossils and molecular biology provide evidence for
this ancient association
17.19 Parasitic fungi harm plants and animals
Of the 100,000 known species of fungi, about 30%
are either parasites or pathogens in or on plants.
About 80% of plant diseases are caused by fungi.
– Between 10 and 50% of the world’s fruit harvest is lost
each year to fungal attack.
– Dutch elm disease destroyed 70% elm trees across the
eastern USA
– A variety of fungi, including smuts and ergot, infect grain
crops.
© 2012 Pearson Education, Inc.
Figure 17.19B
Figure 17.19C
Ergots
17.19 CONNECTION: Parasitic fungi harm
plants and animals
Only about 500 species of fungi are parasitic on
animals.
Skin diseases include
– ringworm, named because it appears as circular red
areas on the skin,
– athlete’s foot, also caused by the ringworm fungus,
– vaginal yeast infections, and
– deadly lung diseases.
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